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
Information for Contributors to Fieldiana
>ciper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
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
Accepted July 2, 1996
Published August 31, 1998
PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
1998 Field Museum of Natural History
PRINTED IN THE UNITED STATES OF AMERICA
Table of Contents
Warfe Davis vii
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
VIII. TRENDS IN SPECIALIZATION 26
IX. GEOGRAPHICAL DISTRIBUTION AND
X. TAXONOMIC TREATMENT 34
Brunfelsia Linnaeus 34
Artificial Key to the Sections, Species,
and Infraspecific Taxa of
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
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.
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
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
23. Distribution of Brunfelsia brasiliensis
and B. chiricaspi 63
24. Brunfelsia brasiliensis subsp. bras-
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
35. Brunfelsia macrocarpa 94
36. Brunfelsia mire 96
37. Distribution of Brunfelsia latifolia, B.
macrocarpa, B. obovata, and B.
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-
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
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
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
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
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
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-
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-
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-
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
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.
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-
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-
J. R. Press
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
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
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-
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
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
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.
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-
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).
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
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-
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
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
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
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
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
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-
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 Solanaceae 1
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.
PLOWMAN: REVISION OF BRUNFELSIA
TABLE 1 . Summary of historical classification of Brunfelsia.
Franciscea (excluding Brunfelsia)
D. Don (1829)
Brunfelsia (including Franciscea)
G. Don (1837)
Brunfelsia (including Franciscea)
Bentham and Hooker (1873)
Brunfelsia (including Franciscea)
Brunfelsia (including Franciscea)
Brunfelsia (including Franciscea)
Brunfelsia (including Franciscea)
Brunfelsia (including Franciscea)
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-
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 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
All known species of Brunfelsia are woody
shrubs, a few of which will become small trees.
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
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).
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-
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
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
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
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
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
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
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
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.
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-
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.
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
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
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
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 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
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-
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.
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
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.
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
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-
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.
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
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-
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-
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-
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-
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.
FIG. 6. Fruit of Brunfelsia americana.
PLOWMAN: REVISION OF BRUNFELSIA
TABLE 2. Chromosome counts in Brunfelsia.
B. calycina (= B. pauciflora)
B. calycina (- B. pauciflora)
2n = 22
n = 11
n = 12
In = 22
2n = 22
2n = 22
n = 11
n = 12
2n = 22
2n = 22
Darlington and Janaki Ammal (1945)
Darlington and Janaki Ammal (1945)
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-
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-
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
CN <N * (N "1 '
1 1 1 1 1
CS <N ro fN <
^7 T |? |? 7 ^
js JS o .c "" M
a. o, as a< <u <u
CS 03 CS 03 "3 23
C C J2 C _g _g
<u <u 3 <u o o
2 2 S J E^ E^
_d IS S cs ca
S| E E"H
Se ^ 3 N N
,. ,5 ^ .0 U U
O^ ^* co co ^N ^N
CN CN CO CO ^ -^
C C C C C C
OS CQ CB OS C3 03
E E E E E E
cu cu cu (X cu a.
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c s e c c c
S! SI 2 S^^ "^S QJ
551 'WS 5
CIQ 0^ 1^ *i; C ^S
CQ QQ QQ CQ QQ ^c,
related genera (Browallia grandiflora sensu Gott-
schalk (1954) (probably B. speciosa Hook.) and
Nierembergia coerulea Gill, ex Miers).
From the preliminary results obtained in this
study, it may now be safely predicted that the base
chromosome number in Brunfelsia is x = 11. Ra-
tera's (1944) count of n = 12 in B. australis was
not confirmed because no flowering material was
available for this species. This count, however,
should be repeated at the earliest possible oppor-
tunity. Both section Brunfelsia and section Fran-
ciscea have 11 pairs. No material from section
Guianenses was available.
The most common chromosome number for the
Solanaceae is 2n = 24. There has been consider-
able debate about whether it can be assumed that
12 pairs represent the base number for the family.
Goodspeed (1954) presented evidence that the
primitive number for the family may be six pairs
(2n = 12), from which higher numbers were de-
rived by secondary association.
Related genera of the tribe Salpiglossideae have
various chromosome numbers. Four genera
(Streptosolen, Salpiglossis, Browallia, Brunfelsia)
have 1 1 pairs (2n = 22). Lower numbers occur in
Petunia (2n = 14), Nierembergia (2n = 18), and
Schizanthus (2n = 20). The endemic Australian
genus Duboisia has 30 pairs (2n 60) (Darling-
ton & Wylie, 1955; Madhavadian, 1968).
The Salpiglossideae are thought to be derived,
at least in part, from the Cestreae, in which tribe
the following chromosome numbers have been
derived: Cestrum L. and Vestia Willd., eight pairs
(2n = 16); Fabiana Ruiz and Pavon, nine pairs
(2n = 18); and Nicotiana L., eight, nine, ten, 12,
18, 19, 20, 21, and 24 pairs, which represents a
well-known aneuploid series (Goodspeed, 1954).
Because the cytology of several important genera,
such as Juanulloa Ruiz and Pavon, Markea Rich.,
and Sessea Ruiz & Pavon of the Cestreae and An-
thocercis and Anthotroche of the Salpiglossideae,
have not been studied, it would be risky to draw
any phylogenetic conclusions on the basis of
chromosome number alone. It seems reasonable,
however, to suggest that Brunfelsia and other gen-
era of the Salpiglossideae originated from an an-
cestral 12-paired precursor through aneuploidy.
VI. Hybridization Experiments
As an aid to investigating the breeding systems
and interspecific relationships in Brunfelsia, a pro-
PLOWMAN: REVISION OF BRUNFELSIA
gram of artificial hybridization was undertaken.
Plants obtained from both wild and cultivated
sources were grown at the Biological Laboratories
Greenhouse, Harvard University, and at North-
eastern University Greenhouses in Woburn, Mas-
There are several inherent difficulties in at-
tempting artificial hybridization in woody peren-
nials such as Brunfelsia. Although germination is
relatively rapid, the time required from seed to
flower varies from a minimum of 6 months to
several years. In Brunfelsia flowering may be spo-
radic or rare under greenhouse conditions, and in
most species only a few flowers per plant are open
at any one time.
Seventeen clones, representing ten species of
the genus, were available for making crosses, in-
cluding five each from section Franciscea and
section Brunfelsia. Other species were grown but
could not be induced to flower. A major problem
in making crosses was the lack of simultaneous
flowering in desired species. Some species (B.
americana, B. pauciflora) flowered almost contin-
uously throughout the year; others flowered only
occasionally or not at all. Attempts to store pollen
under refrigeration for future crosses were unsuc-
Pollination was accomplished by excising the
corolla limb and uppermost part of the tube to
expose the anthers and pistil. Pollen was then
transferred using a small brush. The anthers usu-
ally dehisce before the complete unfolding of the
corolla lobes and scatter the pollen on the inside
of the tube. The stigma is receptive just before the
flower opens and remains so for 2-3 days.
Because of the small number of plants avail-
able, the results of the breeding experiments were
generally inconclusive. The majority of crosses
made, whether inter- or intraspecific, produced no
fruit. The flowers withered and persisted (B.
americana, B. australis) or dropped off in 1-3
days (B. densifolia, B. lactea, B. pauciflora).
The data obtained from the breeding program
are summarized in the following paragraphs. The
actual records of the crosses are not included be-
cause they are lengthy and, for the most part, neg-
atively repetitious. The following species were in-
cluded in the program: B. americana, B. australis,
B. densifolia, B. grandiflora, B. lactea, B. latifol-
ia, B. nitida, B. pauciflora, B. pilosa, and B. un-
dulata. Reciprocal crosses were made between
these where possible.
All species were artificially self-pollinated, al-
though selfing appears to take place naturally. The
dehiscing anthers lie very close to the stigma, and
loose pollen may easily fall upon it. However, no
viable seed was produced from any of the self-
pollinated individuals, indicating that these spe-
cies are self- incompatible. Fruits were occasion-
ally produced spontaneously in B. pilosa and B.
densifolia, but these contained completely aborted
Mature fruit containing viable seeds was pro-
duced in the following intraspecific crosses be-
tween nonclonal individuals within the following
species: B. latifolia X B. latifolia, B. pauciflora X
B. pauciflora, B. americana X B. americana, and
B. densifolia X B. densifolia. However, even in
these species many of the crosses did not take or
produced aborted or hollow seeds. This may be
attributed to various factors. In some cases failure
to set seed may have been due to fluctuations in
greenhouse conditions or the use of certain pes-
ticides. Certain plants (B. americana, B. pauciflo-
ra) were of cultivated origin and may have been
clones of individuals highly selected for ornamen-
tal characters; reproductive irregularities are not
uncommon in these forms. Lastly, these failures
may have been the result of innate incompatibility
factors as yet uninvestigated in the genus.
The failure of interspecific crosses suggests that
all the species investigated are reproductively iso-
lated. In one case this may be significant in view
of specific delimitation. No viable seeds were pro-
duced in crosses between B. australis and B. pi-
losa, two species of southern Brazil that were for-
merly considered conspecific with B. uniflora.
This lends support to the view that B. australis
and B. pilosa should be considered distinct spe-
cies. Additional crosses involving these two spe-
cies as well as B. uniflora would be useful in con-
firming and further clarifying the relationships
VII. Ecology and Reproductive
Brunfelsia species are found in a variety of hab-
itats. Species of section Brunfelsia of the Antil-
lean region are frequently encountered in rocky
woodlands or thickets on limestone (B. malifor-
mis, B. membranacea, B. splendida) or serpentine
soils (B. densifolia, B. shaferi Britton & P. Wilson,
B. sinuatd), often in relatively dry situations. Oth-
er species grow in humid, montane rain forests
(B. portoricensis) and in cloud forests at higher
elevations (B. lactea, B. jamaicensis). Most of the
species of this section are limited in ecological
tolerance and restricted to relatively small and lo-
cal areas. Only B. americana, which also has the
widest ecological amplitude, occurs on more than
one island (see Appendix II for a list of species
in section Brunfelsia and their distributions).
The basic ecology of section Brunfelsia, partic-
ularly the Cuban species, is virtually unknown. As
in the case of many plants of limited distribution,
many of the Antillean species are now in grave
danger of extinction because of the destruction of
their habitats. Several species in Jamaica are
known from a few local populations (e.g., B. ja-
maicensis, B. membranacea, B. plicata, B. splen-
didd) that are rapidly being encroached upon by
human activities. The only known wild population
of B. undulata, on the north coast of Jamaica, was
recently eliminated by the construction of a cattle
path. Brunfelsia picardae Krug. & Urb. of Haiti
has not been collected since 1928. Little is known
of the many Cuban endemics, the majority of
which are rare and known from only one or a few
The three Puerto Rican endemics are currently
protected from man's destruction because they are
now all included within the boundaries of national
parks. However, the only known population of B.
americana in Puerto Rico is precariously located
in a pasture near Coamo.
If the incredible increase in human population
in the West Indies continues at its present rate, the
sheer pressure of human needs will surely lead to
destruction of the few remaining natural areas.
Many of the Antillean species of Brunfelsia have
large, showy flowers of potentially great horticul-
tural value. Several of these have already been
successfully introduced into cultivation (B. amer-
icana, B. nitida, B. undulata), but the remainder
are in danger of being lost completely. In addi-
tion, several of these plants have been shown to
contain alkaloids or to be of interest medicinally.
In this study, an effort was made to distribute
seeds and cuttings of as many wild species as pos-
sible to tropical botanical gardens and conserva-
tories to preserve them in cultivation.
The six species of section Guianenses all grow
in low-elevation tropical rain forests in continen-
tal South America; they are found in the shrubby
stratum of the understory and at the margins of
rivers. Species of this group have not been studied
in the field, and little is known of their biology.
They are mostly known from only a few collec-
The South American species of section Fran-
ciscea are primarily shrubs or small trees of the
understory of humid tropical rain forests; those
species that have adapted to other environments
are considered more specialized. This group
grows primarily in mountainous regions, up to
about 2,000 m of altitude. Exceptions are found
in several species that appear to be restricted to
very low elevations (B. australis, B. imatacana
Plowman, B. latifolia, B. macrocarpa) and in B.
uniflora, which reaches 3,300 m in the Bolivian
A number of species of section Franciscea
have become specialized in their ecological re-
quirements. Brunfelsia latifolia grows only on
sandy restingas, a maritime formation of low
shrubs on the coast of southeastern Brazil. Brun-
felsia obovata is adapted to semiaquatic environ-
ments and grows in marshes (brejos), often in
standing water, or along flooded river margins
from southeastern to central Brazil. In southern
Brazil, B. cuneifolia and B. pilosa are restricted
more or less to the region of the Araucaria forests
of Parana, Santa Catarina, and Rio Grande do Sul,
and B. rupestris grows only in the campo rupestre
habitat of the Serra do Espinha9o in Minas Gerais,
Brazil. Brunfelsia australis, the southernmost spe-
cies of the genus, grows in gallery forests and in
small patches of woodland found on raised
ground in the low-lying Parand River basin.
Two species have a wider ecological amplitude
and are found in varied habitats, including sec-
ondary forest growth and disturbed areas. Brun-
felsia brasiliensis grows in the humid forests of
Rio de Janeiro, especially in well-drained sites
such as outcrops and rocky peaks. It also flour-
ishes in the drier mountainous states of Minas
Gerais and Sao Paulo, often persisting in second-
ary growth woodlands and in gallery forests.
Brunfelsia uniflora is a wide-ranging species that
occurs in the rain forests of the Serra do Mar; in
the drier central and northeastern states of Brazil;
on the dry coast of Venezuela, and, lastly, in the
high-altitude cloud forests of the Bolivian Andes.
This is perhaps the most widespread species in
Brazil because of its ability to survive in disturbed
Certain species of sections Franciscea and Gui-
anenses are of such local occurrence that they
may also be considered endangered. This is par-
ticularly true of B. latifolia, which is precariously
restricted to the region around the city of Rio de
PLOWMAN: REVISION OF BRUNFELSIA
Janeiro, and of B. dwyeri, on the upper slopes of
Cerro Jefe in central Panama. Both of these areas
are rapidly being developed for housing and farm-
ing, respectively. A large portion of Cerro Jefe
has already been decimated for building poultry
farms. Little is known of the effect of human ac-
tivities on other South American species because
they have not been studied in the field. The danger
of their extinction is less serious, however, than
for the West Indian species because of the usually
wider distribution of the continental species.
The seeds of Brunfelsia species germinate with-
in 2-4 weeks of planting and are relatively short-
lived. They will not tolerate desiccation. Most
species are slow-growing woody shrubs that vary
in the time required to mature from seedling to
flowering plant. Under greenhouse conditions
plants mature from 6 months (B. latifolid) to 2 or
more years after germination. Several species are
known to flower while the plants are still small,
even though they may eventually become trees in
nature. Little is known about the longevity of in-
dividual plants, but there is evidence that they
grow for at least 20 years (B. grandiflora). Veg-
etative reproduction may also occur to a limited
extent by the rooting of broken branches on the
forest floor. This has been observed in B. chiri-
caspi (Plowman 2081).
The flowering period in Brunfelsia may be
short, lasting only a few weeks, or extend more
or less continuously throughout most of the year.
Flowering specimens have been collected from
the type locality of B. grandiflora subsp. schul-
tesii in January, May, and September. The major-
ity of the South American species flower between
October and December.
The fruits of Brunfelsia require several months
to a year for ripening and may remain on the plant
even after the seeds are fully mature, until the
fruits are dried and brown. Dehiscence of the
fruits is discussed under "Dispersal Mecha-
The flowers of Brunfelsia are well adapted for
insect pollination, specifically by moths and but-
terflies. No other kinds of insects have been ob-
served to visit the flowers. The three basic forms
of the corolla found in the genus have apparently
evolved through selection for specific groups of
pollinators: section Franciscea for butterflies, sec-
tion Brunfelsia for hawk moths, and section Gui-
anenses probably for other moths [or perhaps
bees Eds.]. All the species have a disc at the
base of the ovary that secretes abundant nectar in
the base of the tube. So far as is known, they are
Species of section Franciscea have flowers
with a relatively short tube and a brilliant violet
limb with a white eye at the throat, which pre-
sumably serves as a nectar guide. The somewhat
raised, fleshy rim at the throat may function as a
landing platform for the visiting insect. The violet
color of the flower fades to pale lavender or pure
white with age and may signal the insect that the
flowers are no longer receptive. The flowers are
rarely scented in this group, although two species
(B. australis, B. imatacand) are exceptionally fra-
grant with a scent resembling that of jasmine.
Butterflies in four different families have been
collected on the flowers of B. grandiflora, includ-
ing species of Euptychia (Nymphalidae), Eurema
(Pieridae), Euselasia (Lycaenidae), and the Cal-
podes group (Hesperidae) (see Table 4). In south-
ern Colombia the most frequent visitor of these
was an unidentified species of Euptychia. 4
Species of Brunfelsia sect. Brunfelsia, in con-
trast, possess a much longer corolla tube than
those of section Franciscea. They are usually
white upon opening and fade to yellow with age.
They bloom at night and are highly perfumed, be-
ginning to produce scent just before sunset. The
fragrance of a few flowers is capable of pervading
a relatively large area of forest. These flowers
have no landing platform as found in section
Franciscea, which suggests that the insect hovers
Although no insect pollinators have been seen
on the flowers of the Antillean species, these are
typically pollinated by moths. In view of the
length of the corolla tube (up to 14 cm), it may
be said with some certainty that hawk moths
(Sphingidae) are responsible for pollination in this
group. Similarly constructed, fragrant white flow-
ers in other families of plants are known to be
pollinated by hawk moths, such as Ipomoea alba
Garcke (Convolvulaceae), Aquilegia longissima
A. Gray (Ranunculaceae), Habenaria leucophaea
A. Gray (Orchidaceae), Hippobroma longiflora G.
4 Identified by Prof. John Burns, Museum of Com-
parative Zoology, Harvard University.
TABLE 4. Pollinators of Brunfelsia grandiflora.
Euptychia sp. 1
Euptvchia sp. 2
Rio de Janeiro, Brazil
Rio de Janeiro, Brazil
Don (Campanulaceae), and species of Randia L.
(Rubiaceae), Quisqualis L. (Combretaceae), and
Brugmansia Pers. (Solanaceae).
The pollinators of section Guianenses are not
known. The flowers in this group have a short
tube but are white and have no landing platform;
both B. amazonica and B. chocoensis are reputed
to be fragrant, suggesting that they may also be
pollinated by moths. This remains to be confirmed
in the field.
The fruits of Brunfelsia are basically capsular
and of two general types: a more or less thin-
walled capsule that becomes dry and brittle with
age or a more or less thick-walled capsule, the
valves of which may be fleshy, leathery, or bony
at maturity. Both types of fruit occur in all three
sections of the genus.
The fruits of most species of Brunfelsia do not
dehisce to release the seeds as is common in most
capsules; at best they are tardily dehiscent. They
frequently remain on the plant well after the seeds
are ripe, becoming dry and brown, in which state
of brittleness they may then break apart on the
plant [but see discussion of fruit types in Solanum
in Symon (1979) Eds.]. Alternatively, the whole
capsule may fall to the ground, where the outer-
most layer of the pericarp rots away, leaving the
thin, brittle endocarp. This eventually breaks in
half along the lines of dehiscence to release the
Most of the South American species of Brun-
felsia have capsules that are green or brown at
maturity and included in the dark green calyx. As
a result, the fruits are very inconspicuous; fruiting
specimens are almost never collected. In the ab-
sence of any pronounced color, odor, or edible
flesh, it seems unlikely that the fruits could be
attractive to animals. These species apparently
have no special adaptations for dispersal, a situ-
ation that has been pointed out for many shrubs
and trees of the primary rain forest (Richards,
The thin-walled, capsular species may be dis-
persed over short distances by water. The air spac-
es created by placental breakdown and the lack of
dehiscence permit these fruits to float. After they
have fallen to the ground, they may be carried
away from the parent plant during heavy rains,
especially along river courses.
Other species in South America and in the West
Indies have rather large, orange or yellow fleshy
fruits that are probably dispersed by birds or
mammals. Examples of these are B. chocoensis,
B. densifolia, B. guianensis, B. macrocarpa, and
B. nitida. Two of these (B. chocoensis, and B.
macrocarpa) are said to be edible and probably
have a sweet flesh that would attract animals.
However, animal dispersal has not been observed
in the field.
In addition to pollinators, other insects are in-
timately associated with the genus Brunfelsia.
One of the most interesting is the butterfly genus
Methona (Rhopalocera, Nymphalidae), the larvae
of which feed exclusively on the leaves of Brun-
felsia species. This and related groups have been
studied by Dr. Gerardo Lamas of the Museo de
Historia Natural, Lima, Peru (Lamas, 1973).
Methona (Fig. 7) is a genus of medium-sized
butterflies (wing span of 7-10 cm) consisting of
seven species and nine subspecies. The larvae are
PLOWMAN: REVISION OF BRUNFELSIA
FIG. 7. Methona themisto (type specimen at BMNH).
gregarious and warningly colored (Fig. 8). The
species of Methona are all tropical forest insects
and are distributed in the same areas of South
America where Brunfelsia occurs. The greatest
concentration of Methona species is found in the
Andes; three species occur in the Amazon basin,
two occur in northeastern Brazil, and one occurs
in southeastern Brazil. A disjunct population, con-
sidered a distinct subspecies of M. confusa, lives
only on Cerro Jefe in Panama (Lamas, 1973).
Until the present, the documentation of the
Brunfelsia-Methona relationship has been patchy,
primarily because of the lack of voucher speci-
mens of the host plants. Larvae of M. themisto
megisto have been collected on B. australis in Mi-
siones Province of northeastern Argentina (Plow-
man 2731} as well as in Buenos Aires and Uru-
guay. Reports from the latter two sites were made
from cultivated plants that were incorrectly iden-
tified as B. pauciflora, B. uniflora, and B. latifolia,
even though none of these species are known this
far south in the wild or in cultivation. The same
species of Methona has been collected in Tucu-
man in western Argentina. Although no species
of Brunfelsia occurs there naturally, B. australis
is cultivated in gardens and must serve as the
Other species of Methona have also been found
on Brunfelsia species: M. singularis in Paraiba in
northeastern Brazil and M. confusa confusa in
Manaus in the Amazon basin. Without voucher
specimens, it is difficult to say which species of
Brunfelsia were involved.
The geographical distribution of Methona ex-
hibits a striking parallel to that of the South Amer-
ican brunfelsias. The butterfly, however, does not
FIG. 8. Caterpillars of Methona confusa on leaf of Brunfelsia chiricaspi.
PLOWMAN: REVISION OF BRUNFELSIA
TABLE 5. Brunfelsia species recorded as larval food
plants for species of Methona (Lepidoptera: Nymphali-
M. M. M. M.
Species confusa megisto singularis themisto
occur in the West Indies, suggesting that it arose
in South America after Brunfelsia sect. Brunfelsia
became cut off from its South American ances-
tors. Not enough data are available at the present
time to indicate whether there may be any spe-
cies-specific relationships between Brunfelsia and
Methona. A summary of the species of Brunfelsia
fed upon by the larvae of Methona is presented
in Table 5 [more detailed information can be
found in Drummond and Brown (1987) Eds.].
The capsules of Brunfelsia species in South
America are also fed upon by Lepidopteran lar-
vae. The flesh of the young fruit and the seeds
are eaten by tiny larvae 5 of the family Gelechi-
idae. These have been found in the following spe-
cies: B. australis (Plowman 2723), B. brasiliensis
subsp. macrocalyx (Dusen) Plowman (Plowman
2891, 2906), and B. grandiflora subsp. schultesii
(Plowman 2404). Further identification of the in-
sects has not been possible. They apparently lay
their eggs on or near the developing fruits. The
larvae then burrow through the pericarp and de-
vour the seeds and placental tissue. 6
Protection from this predation may be the ori-
gin of certain specializations in the genus. The
fruits of several species (B. macrocarpa, B. hy-
drangeiformis, B. pauciflora) are completely en-
closed in the often coriaceous calyx, which may
offer some defense against the insect larvae. The
thick, leathery valves in the capsules of B. dwyeri
may serve a similar function.
A third insect-plant relationship has been ob-
5 Identified by Dr. Donald Duckworth, U.S. National
Museum. Washington, D.C.
6 The taxon-specific relationships between members
of the Gelechiidae and other Solanaceae have been well-
documented in the temperate zone but not in the tropics
(see Foott, 1967; Scholtz, 1978; Solomon & McNaugh-
ton, 1979) Eds.
served in southern Colombia in B. grandiflora
subsp. schultesii. Two species of ants 7 in the gen-
era Campanotus (Formicinae) (Plowman 2416)
and Pheidole (Myrmicinae) (Plowman 2151) form
temporary bivouacs among the fruit clusters of the
plants. Associated with the ants are two different
Homopteran insects (one in the family Psillidae,
the other unidentified) that feed on the plant and
apparently are tended by the ants. Pheidole man-
ufacture a nest of "carton" (cellulosic material)
among the fruits that may serve to protect their
charges. This may represent a symbiotic relation-
ship between the ants and Homopterans, but it is
not clear what advantage, if any, this may give to
VIII. Trends in Specialization
In attempting to understand the evolutionary re-
lationships among the sections and species of
Brunfelsia, it is necessary to determine which
characters are primitive or unspecialized and
which are derived and specialized. Levels of spe-
cialization may be observed in several organs of
the plants, including leaves, flowers, and fruits, as
well as in habitat preferences and other ecological
factors. Because the species of section Brunfelsia
have not been studied in depth, the following dis-
cussion is limited primarily to the South American
species of section Guianenses and section Fran-
Of the South American brunfelsias, the flowers
of section Guianenses are the least specialized
and possibly ancestral to section Brunfelsia. The
corolla tube is short in section Guianenses and
open at the mouth with relatively narrow corolla
lobes. The corolla is white or greenish white, sug-
gesting pollination by moths.
Section Guianenses occurs primarily in the
Amazon basin with disjuncts in Choco and Bahia.
There is a distinct tendency in this group toward
axillary, few-flowered inflorescences on the older
branchlets, although terminal flowers are also pro-
duced. Two species (B. chocoensis, B. guianensis)
have capsules with fleshy valves, and two others
(B. amazonica, B. clandestina) have a dry, thin-
walled capsule. The fruit of the remaining two
species (B. martiana, B. burchellii) is unknown.
Evolution in the type of fruit, however, shows no
7 Identified by Prof. E. O. Wilson, Department of Bi-
ology, Harvard University.
overall direction because different types of fruit
(dry vs. fleshy, etc.) have probably arisen repeat-
edly under local selection pressures.
Section Guianenses is a small group that occurs
in a relatively uniform environment with few ma-
jor differences separating the species. It is there-
fore not surprising that trends of specialization are
difficult or impossible to detect within the group.
Section Franciscea apparently arose as a sec-
tion by becoming isolated in the Andean region,
where the flowers coevolved with butterflies. The
flowers are bright purple or violet with a distinct
white or yellow eye. The corolla tube is short and
somewhat constricted at the apex into a thickened
ring. The limb is often large and showy with
broad, rounded lobes.
The inflorescence of section Franciscea varies
from a many-flowered, lax or compact cyme to a
solitary terminal flower. Most of the species bear
a few-flowered terminal cyme; the number of
flowers per inflorescence may vary considerably
in a single species (see "General Morphology").
Evolution in the inflorescence must first of all
be considered in terms of the entire plant and pol-
lination strategy. Species with several to many
flowers in the inflorescence tend to have fewer
flower-bearing branches (B. chiricaspi, B. hydran-
geiformis, B. mire). Species with only one to three
flowers are generally more profusely branched,
with many of the branchlets bearing flowers (B.
australis, B. cuneifolia, B. uniflora}. In the first
case, the flowers are aggregated into a large single
mass (see Fig. 30); in the second, the entire shrub
may be diffusely covered with many single flow-
ers (see Fig. 45). Both modes of organization
must be equally attractive to pollinators, although
there may be different butterflies associated with
the different forms of inflorescence. This associ-
ation, however, has not been observed.
There is also a tendency for the one-flowered
species to have larger flowers (B. macrocarpa, B.
pilosa). Among the species with several (one to
ten) flowers, there are many intermediate situa-
tions with respect to the number of flowers per
branch, the number of branches, and the size of
flowers. In adapting to local ecological conditions,
available pollinators, etc., the species of section
Franciscea have evolved a mosaic pattern of in-
florescence organization within which these gen-
eral trends can be perceived.
Attempts to ascertain the "primitive" condition
of the inflorescence, i.e., whether it may have
been one-, few-, or many-flowered, branched or
unbranched, etc., leads to several problems. In
terms of flower and fruit characters, two species
appear to be least specialized, namely B. uniflora
and B. grandiflora. Both of these occur in the An-
dean region, as well as in eastern Brazil, indicat-
ing, as shown below [and in Plowman (1979)
Eds.], that both species are relatively old for the
genus. However, B. uniflora, as its name suggests,
has but a single flower terminating the branch; B.
grandiflora may have few to many flowers in a
more or less branched inflorescence. We cannot,
therefore, decide which of these may have a more
primitive inflorescence solely on the basis of as-
There has been considerable controversy over
the origin of inflorescence types in the Angio-
sperms (for discussion see Constance, 1955; Law-
rence, 1965). According to Lawrence, the inflores-
cence is a branch system in which a solitary flower
is the terminal unit of each branch. Some workers
believe that this single terminal flower is the prim-
itive inflorescence from which more complex
forms were derived through increased lateral
branching. Others, including Rickett (1944), whose
views have been widely repeated, interpreted the
sequence in the opposite direction. Rickett consid-
ered the primitive condition to be a compound,
branched inflorescence, specifically a compound
dichasium, from which few- or solitary-flowered
inflorescences were derived through suppression of
lateral branches. He derived various inflorescence
types, including the cincinnus, by reduction and
condensation of the branch system. The solitary-
or two- to three-flowered cyme could easily be de-
rived from the cincinnus by mere suppression of
the lateral branchlets. Accordingly, Lawrence con-
cluded that "the modern or present-day inflores-
cence represented by a solitary flower in almost all
plants is a product of reduction, a reduction that
may have resulted from any one of many multi-
flowered inflorescence types, whose origin may be
indicated only by comparative studies of inflores-
cences of related taxa."
One genus in the Solanaceae has been thor-
oughly studied from this point of view. Good-
speed, in The Genus Nicotiana (1954), postulated
that an open paniculate inflorescence (a broad
thyrse) is primitive in this group, from which a
wide array of inflorescences has been derived, in-
cluding glomerules and false racemes.
In conclusion and with reservations, I would
designate the pluriflorous, branched inflores-
cences of B. grandiflora as the probable ancestral
type for section Franciscea, from which fewer-
flowered forms such as B. bonodora (Veil.) J.F.
PLOWMAN: REVISION OF BRUNFELSIA
Macbr. and B. uniflora have been derived by re-
duction. Condensed, many-flowered inflores-
cences such as are found in B. mire and B. hy-
drangeiformis could easily be derived from that
of B. grandiflora by a shortening of the axis with
a concomitant increase in the number of lateral
Specializations in the calyx of Brunfelsia have
occurred both between and within the sections. In
section Brunfelsia the calyx is usually small and in
fruit appears only at the base. The same is true of
section Guianenses with the exception of B. ama-
zonica and B. clandestina, in which the calyx par-
tially encloses the thin-walled fruit. In section
Franciscea, the latter situation is the rule: the calyx
persists tightly appressed to the developing fruit,
enclosing only the lower half of the fruit (B. aus-
tralis, B. grandiflora) or far exceeding it in length
(B. hydrangeiformis, B. pauciflord). Even in the
fleshy-fruited members of this section (B. macro-
carpa, B. dwyeri), the calyx envelops the fruit and
in the former species becomes very large. The ca-
lyx in these species remains attached to the fruit
when it falls from the plant.
The relative length of the calyx at anthesis var-
ies considerably both within species (B. bras-
iliensis) and among them. However, a longer ca-
lyx in relation to the corolla tube indicates in-
creased specialization, possibly as protection
against robber bees or other insects that lance the
base of the corolla tube to obtain nectar. Another
feature of the calyx that is considered derived is
the inflated and five-angled condition found pri-
marily in B. boliviano, B. cuneifolia, and B. ob-
The characters of the corolla were discussed
above. The corollas of each of the three sections
are distinctive, as demonstrated above, but show
little variation or evolution within each section.
However, minor differences do occur. In section
Brunfelsia there has been some differentiation in
the size of the corolla limb relative to the tube
and in the color of the corolla. In section Fran-
ciscea. although somewhat variable in size and to
a lesser degree in color, the corolla is relatively
uniform, as it also is in section Guianenses.
Equally conservative is the form and relative
size and position of the stamens and pistil. Within
each section, there has been apparently little
change. A tendency toward exsertion of the up-
permost pair of anthers has arisen convergently in
all three sections, but the adaptive significance of
this, if any, is not known.
Several different types of fruit are found in the
genus Brunfelsia and are distributed in all three
sections. These have been classified variously as
dehiscent or indehiscent berries; as fleshy, leath-
ery, or dry capsules; or as drupes. After examin-
ing most of the species for which fruits are
known, I would suggest that the basic or primitive
fruit in the genus is a tardily dehiscent capsule
with leathery valves. This basic form has under-
gone several modifications in response to partic-
ular environmental conditions and selection pres-
sures for dispersal.
One of the most generally encountered derived
characters is the loss of dehiscence, which may
be partial or complete, in both thin- and thick-
walled types. However, in many species, and es-
pecially in section Franciscea, the septicidal su-
tures (lines of dehiscence) remain on the fruit.
There may be two additional sutures that divide
the valves, apparently a vestige of previous four-
way dehiscence. This mechanism is seen today in
capsules of other genera in the Salpiglossideae,
such as Salpiglossis, Browallia, and Streptosolen,
as well as in Nicotiana and Sessea of the Cestreae.
In Brunfelsia dwyeri the sutures are cruciform and
prominently indented in the surface of the fruit.
A septicidal line of dehiscence is readily observed
in the endocarp of many species, which is usually
cartilaginous and brittle on drying. As the mature
fruit rots, the endocarp, which is often more du-
rable than the mesocarp, remains and eventually
breaks along the sutures through the agencies of
weathering and predation. This I believe to be a
vestige of the ancestral capsular form.
It is of interest here to compare the view of
other workers in the field of Angiosperm evolu-
tion. Hutchinson (1969) believed that there is a
general morphological trend within the Angio-
sperms in which the capsule has preceded the ber-
ry and drupe. Corner (1964), in summarizing his
"Durian Theory," stated that "the primitive fruits
of flowering plants were dehiscent arillate carpels
or, in the syncarpous ovary, capsules. These fruits
had bright red or orange walls and black seeds
covered with red or orange edible arils." From
this original type he derived the dry capsule with
dry seeds, indehiscent types that led eventually to
the fleshy berry, and the stone fruits and nuts by
lignification of part or all of the fruit wall.
Similar trends can be readily observed in the
Solanaceae operating at various taxonomic levels,
including tribe, subtribe, genus, and species. The
family is characterized by two main fruit types,
the berry and the capsule. Both types occur in
four of the five tribes of Wettstein (1895). The
fifth tribe, represented solely by Nicandra, bears
a "dry berry."
Besides Brunfelsia, other genera show hetero-
morphism in their fruits. The genus Lycium, mo-
nographed by Hitchcock (1932) and pointed out
by Croizat (1962), has mostly fleshy, tomato-like
berries except in two species, L. ameghinoi Spe-
gazzini and L. californicum Nutt., which bear cap-
sules. An anomalous species, L. macrodon A.
Gray, has a curious intermediate type of fruit.
A second example is found in the genus Cestrum
and its close relative Sessea. Francey (1935, 1936)
emphasized that the only "good" character to sep-
arate these two genera is the nature of the fruit,
which in Cestrum is baccate and in Sessea a four-
valved capsule. However, even this character, he
adds, breaks down in S. farinosa (Urb. & Ekman)
Francey, the young fruit of which is baccate, and
in C. strigillatwn Ruiz & Pavon and C. lindenii
Dunal, in which the young fruit bears a cruciform
suture at its apex, a feature typical of Sessea.
In that flowers and fruits tend to evolve as in-
dependent units (Davis & Hey wood, 1963), it is
not surprising to find both capsules and berries at
different levels in an evolutionary scheme based
primarily on floral morphology. Irrespective of
their relative primitiveness, the occurrence of two
types of fruits in the same subtribe or genus in-
dicates differential evolutionary rates. The fruit
has continued to change under varying conditions
of climate, habitat, and selection for dispersal, and
presumably different types of fruits may arise
more than once, indicating a mosaic rather than a
directional pattern of evolution. Constance (1955)
summarized this phenomenon as follows: "anal-
ogous dehiscent or indehiscent, dry or fleshy, and
one- or several-seeded fruit types can arise any-
where along this sequence, presumably as a result
of selection pressure of biological demands." De-
tailed morphological studies of fruits in all three
sections of Brunfelsia along with ecologically ori-
ented field work will add much to our understand-
ing of the dynamic relationships between these
basic fruit types.
Specialized features are also found in the vege-
tative organs, which in many cases can be related
directly to environmental conditions. Leaf size and
texture are particularly variable according to exist-
ing climatic conditions. Species of humid tropical
forests typically have relatively large leaves (B.
chiricaspi, B. hydrangeiformis, B. imatacana, B.
pauciflora, B. plicatd). Those of the more exposed,
drier habitats frequently exhibit smaller, often more
coriaceous leaves (B. brasiliensis, B. latifolia, B.
nitida, B. obovata, B. rupestris, B. splendida).
There are many intermediate situations as well as
species that show either type because they occur in
both habitats (B. uniflord). Several of the Antillean
species that inhabit serpentine soils have thick-co-
riaceous, linear leaves (B. linearis in Cuba, B. den-
sifolia in Puerto Rico, Fig. 4). Other species that
grow in high-altitude cloud forests have developed
thick-coriaceous, very shiny leaves (B. lactea in
Puerto Rico (Fig. 5), B. dwyeri in Panama, B. un-
iflora in Bolivia).
Another specialization in vegetative characters
is the crowded or subverticillate arrangement of
the leaves at the end of the stem (B. hydrangei-
formis, B. mire). This generally appears in con-
junction with a dense or capituliform inflores-
cence and very large leaves. The verticillate ar-
rangement of the leaves in B. rupestris is of a
slightly different nature, with crowded groups of
leaves occurring at intervals along the stem.
An example of convergent evolution has ap-
parently taken place in the leaves and fruits of B.
macrocarpa and B. chocoensis of the Pacific low-
lands of Colombia. These species belong to sec-
tion Franciscea and section Guianenses, respec-
tively. The leaves of these species are very similar
in size, shape, and venation, and both species
have fleshy, yellow fruits. However, the flowers
indicate that they belong to different phyletic
lines, although both are somewhat anomalous in
The digression of various species from the trop-
ical forest environment and subsequent adaptation
to more specialized ecological zones, such as
rocky outcrops (B. brasiliensis), sandy shores (B.
latifolia), marshes (B. obovata), campo rupestre
habitat (B. rupestris), or coastal rocks (B. ameri-
cana), are also indicative of evolutionary special-
ization. These aspects are discussed in "Ecology
and Reproductive Biology." Further details are
found in "Taxonomic Treatment" under individ-
ual species entries. The trends of specialization in
the genus are summarized in Table 6.
IX. Geographical Distribution and
The current distribution of species and subspe-
cies, correlated with morphological, cytological,
8 Plowman (1979) provides an in-depth discussion of
the biogeography of Brunfelsia; the material in this sec-
tion is intended as a summary only. Eds.
PLOWMAN: REVISION OF BRUNFELSIA
TABLE 6. Trends in specialization of Brunfelsia.
Plants of humid, tropical forests
Leaves scattered on stem
Leaves large, soft-leathery
Inflorescence terminal and subterminal
Inflorescence lax, branched
Flowers few-many, small
Calyx shorter than corolla tube
Calyx terete, narrow
Calyx present at base of fruit only
Calyx not accrescent
Corolla tube straight, gradually dilat-
ed, and open at apex
Corolla lobes white or greenish, not
Corolla lobes narrow, not overlapping,
Capsule valves leathery or fleshy
Plants of other habitats
Leaves crowded at branch tips or subverticillate
Leaves small, coriaceous
Inflorescence exclusively terminal or axillary
Inflorescence compact, sessile, and unbranched
Flowers solitary or few, large
Calyx equaling corolla tube
Calyx angled, inflated
Calyx partially or totally enclosing fruit
Calyx more or less accrescent
Corolla tube curved and inflated at apex, then slightly constricted
and thickened to form ring
Corolla changing color (white to yellow, purple to white)
Corolla lobes broad, overlapping, undulate to crenate
Capsule valves thick, fleshy or bony, or thin-walled and dry
and other characters, provides us with the basis
for erecting hypothetical phylogenies that indicate
the possible historical relationships among pres-
ent-day species. Unfortunately, no fossil remains
of Brunfelsia or any related Solanaceae have been
found to corroborate the relationships proposed
here, nor have in-depth phylogenetic analyses
In Brunfelsia evidence from morphology and
geographical distribution can be interpreted and
evaluated against the background of the geologi-
cal history and biogeographical patterns in other
plants and in animals to infer the history of the
genus. Such a scheme must be conjectural consid-
ering the paucity of data on several fronts, es-
pecially the all-important gaps in distributional
patterns, the result of inadequate collecting. Nev-
ertheless, the available data are full of implica-
tions that demand thoughtful consideration.
The major divisions of Brunfelsia, represented
by three sections, appeared as the result of an ear-
ly divergence in the ancestral, pre-Brunfelsia lin-
eage, which in turn arose from primitive Salpig-
lossoid stock somewhere in South America. These
three lines of divergence are evident today in the
basic differences in floral morphology among the
three groups and are associated with distinct and
mostly allopatric areas of distribution. This sug-
gests that the early differentiation of the ancestral
pre-Brunfelsia was accompanied by geographical
isolation in the three major areas in which they
are found today.
The three sections of Brunfelsia are centered in
three major areas: section Brunfelsia in the West
Indies; section Guianenses in northern South
America, primarily in the Amazon basin; and sec-
tion Franciscea throughout most of tropical South
America, excluding desert regions and the low-
land forest of the Amazon.
All the species occurring in the West Indies be-
long to Brunfelsia sect. Brunfelsia (Fig. 9). This
group, which exhibits both primitive and ad-
vanced characters, apparently diverged very early
from a South American progenitor that was pos-
sibly similar to present-day section Guianenses.
The most likely route of entry of the genus into
the Antilles seems to be via the inner arc of the
Lesser Antilles. These islands, which were formed
in the late Cretaceous or early Tertiary period,
have been volcanically active since the Eocene
epoch to the present (Malfait & Dinkelman, 1972)
[but see references cited in Liebherr, 1988, and
Page & Lydeard, 1994, for competing theories for
the origin of the Antilles Eds.].
The hypothesis that the Lesser Antilles was the
entry route of Brunfelsia into the Caribbean is
based primarily on the present distribution of spe-
cies. The group most closely related to the Antil-
lean species is section Guianenses, which has its
center of diversity in northern South America.
The only species of Brunfelsia that occurs to-
day in the Lesser Antilles is B. americana, the
least specialized and most widely distributed spe-
cies of the genus. It is also found in Puerto Rico
and the easternmost tip of Hispaniola. This plant
has fleshy, bright orange fruits that may be carried
6 endemic/6 total 1 endemic/2 total
FIG. 9. Distribution of section Brunfelsia.
by birds from island to island. The agency of
long-distance dispersal by birds cannot be ignored
in considering the migration of Brunfelsia into the
Caribbean, although a path of "island hopping"
seems more logical. Brunfelsia americana occurs
as far south in the Lesser Antilles as St. Lucia;
currently prevailing arid conditions apparently ex-
clude it from the southernmost islands of the
chain. Although it is cultivated in St. Vincent and
Trinidad, B. americana is not known to occur nat-
urally in these islands at the present time.
No species are known from Central America
north of Panama. This greatly detracts from the
PLOWMAN: REVISION OF BRUNFELSIA
possibility that Brunfelsia may have entered the
Antilles via some former land connection with
Central America. Such a connection has been sug-
gested with regard to the Nicaraguan Rise, a shal-
low area in the Caribbean extending from Nica-
ragua toward Jamaica [but see papers on Carib-
bean biogeography cited above Eds.].
About 20 species of section Brunfelsia are
known and are distributed throughout the major
islands of the Caribbean, excluding the Bahamas,
the eastern "limestone" islands, and the southern-
most Lesser Antilles. There is a high degree of
endemism on the largest islands, with about ten
101 BWWLL HALL
1 9 1999
FIG. 10. Distribution of sections Guianenses (dark gray shading) and Franciscea (pale gray shading); areas where
both sections occur are shown in black.
endemic species in Cuba, six in Jamaica, one in
Hispaniola, and three in Puerto Rico. Only B.
americana is found on more than one island.
This radiation of species, typical of island evo-
lution, appears to have arisen from reproductive
and ecological isolation of populations. Many of
the species are restricted to very local areas and
appear to be effectively allopatric. Nothing is
known of the breeding systems or pollinator-
flower relationships that may have played a major
role in reproductive isolation in these plants, es-
pecially among the sympatric species. 9 A partic-
ularly fertile area in this regard is the Oriente
Province of Cuba, in which eight endemic species
have been reported. These questions, however, are
9 V. Fuentes, of the Botanic Garden, Havana, is pres-
ently working on this group; his results will be of great
interest in resolving some of the questions posed here.
beyond the scope of the present study, in which
only South American species and species com-
plexes are treated in depth.
Brunfelsia sections Guianenses and Franciscea
are essentially South American. Two species do
extend into Panama but are clearly of South
American origin. For the most part the two sec-
tions displace each other geographically (Fig. 10).
Section Guianenses, with six species, occupies the
lower Amazon basin, extending north into the
Guianas, south to Bahia in Brazil, and west almost
to the Colombian border. One species is isolated
in the Choco region of northeastern Colombia.
Section Franciscea, which includes 15 species, is
much more widespread, occurring over a large
portion of the South American continent in a di-
versity of habitats. These species are found from
the Caribbean coast south to about 30 latitude
and from sea level to about 3300 m altitude.
Most species of section Franciscea are found
in three major geographical provinces, corre-
sponding to the "regional centers" recognized by
Tryon (1972) for major areas of speciation in
ferns. The most outstanding of these, in terms of
numbers of species, is southeastern Brazil, which
has 1 1 species of Brunfelsia, nine of which are
endemic. The second major area is the Andean
region, which has five species, three of which are
endemic. The third region more or less agrees
with the Guiana center of Tryon and includes
parts of Guyana, Venezuela, and Roraima Terri-
tory in Brazil. Two species are found here; one is
endemic, and the other is widely distributed.
The patterns of distribution of the species dis-
cussed here may reflect to some extent the amount
of collecting in the different areas, especially
where gaps in distributions are found in poorly
collected areas such as the Amazon basin [see
Nelson et al., 1990, for a discussion of this prob-
lem Eds.] Recent intensive collecting sponsored
by the New York Botanical Garden (New York,
N.Y.) and the Institute Nacional de Pesquisas da
Amazonia (Manaus, Brazil) have added many
new distribution records in formerly unknown ar-
eas. It is certain that the species that occur there
will be shown to be more widely distributed as
these areas become floristically better known.
It is here believed [see Plowman, 1979 Eds.]
that most of the present distributions of Brunfelsia
in the circum-Amazonian region, and probably in
other parts of South America, are the result of
climatic and geological changes that occurred dur-
ing the Pleistocene epoch or more recently. Vuil-
leumier (1971), in a review article, summarized
these events in view of their effects on the flora
and fauna of the continent, with particular em-
phasis on the role of glaciation. 10
Distribution patterns of Brunfelsia species oc-
curring in the Andes correspond surprisingly well
to the refugia of Haffer, Vanzolini, and Williams
[see Plowman, 1979 Eds.]. It will be some time
before these theories can be tested and retested by
correlation with distribution patterns in other
groups of organisms and with the ever-increasing
body of knowledge of the geological and climatic
history of the South American continent. But for
the moment, the cyclic fragmentation and expan-
sion of forests and their component species pre-
sents a working model for understanding the iso-
10 The refugium hypothesis to which Plowman is al-
luding here and its present status are covered in a variety
of works (e.g.. Nelson et al., 1990; Mallet, 1993). Eds.
lating barriers that have created the patterns of
differentiation and geographical distribution in
As here interpreted, the distribution and hypo-
thetical relationships of Brunfelsia sect. Francis-
cea suggest a complex picture of speciation, mi-
gration, and extinction of populations. Some spe-
cies are confined to relatively small areas in one
of the three major centers of distribution. Others
are found in more than one of these distant areas,
either as the same species or as vicarious pairs. It
is these plants that offer clues to the history of
their origin and distribution over wide areas.
An outstanding aspect of the distribution in sec-
tion Franciscea is the relationship between the
Andean and southeastern Brazilian centers, which
have one species and two pairs of vicarious spe-
cies in common. In floral and fruit characters
these species are relatively unspecialized with the
exception of B. hydrangeiformis. In terms of the
vegetative and inflorescence characters, B. gran-
diflora and B. bonodora are the least specialized
and are considered to be relatively old species that
may have given rise to others.
The present bicentric distribution of these spe-
cies suggests that the two centers may have been
in contact in the past. Several workers (Good-
speed, 1954; Smith, 1962; Vuilleumier, 1971)
have offered the opinion that the intervening re-
gion (eastern Bolivia, Chaco, Mato Grosso) may
have been wetter and cooler during the past, per-
haps in conjunction with periods of glaciation or
other climatic cycles. A more favorable climate
would have permitted the development of tropical
forests and an avenue for migration and gene ex-
change between the Andean region and south-
Today this region is hot and dry and largely
covered with cerrado vegetation, an arid forma-
tion of shrubs and stunted trees. It is an inhospi-
table habitat for wet tropical forest species, serv-
ing as an effective barrier to gene flow. The final
uplift of the Andes was apparently responsible for
the rain shadow now existing in the Chaco-Mato
Grosso area. According to L. B. Smith (1962), the
southward flow of heat from the Amazon and the
northward encroachment of aridity from the south
via the Chaco depression resulted in a climatic
barrier that prevented further gene exchange be-
tween the two regions, creating many of the pres-
ent-day disjunctions of species between them.
During the hypothetical period of more or less
continuous forest between the Andes and southern
Brazil, species were free to migrate in either di-
PLOWMAN: REVISION OF BRUNFELSIA
rection. At this time the primitively Andean spe-
cies migrated into the Brazilian highlands and to
the Serra do Mar, where a secondary center of
There is evidence that some of these species
eventually migrated back to the Andes during a
period of forest expansion and were later severed
from their parent populations by the unfavorable
climate that ensued. This appears to have occurred
in B. uniflora.
In many groups of plants, the area in which the
greatest diversity of species occurs is also the area
of origin of the group. This idea was particularly
favored by Vavilov (1950), who worked with the
wild progenitors of cultivated plants. Cain (1971),
however, has pointed out that this by no means
holds true for all groups of plants and has pro-
posed the origin of secondary centers of specia-
tion as follows:
It is entirely possible, then, that a phyletic stock that has
had its origin elsewhere, may, through migration, en-
counter a region in which there are numerous available
ecological niches that are unsaturated, that is, in which
competition pressure is low. Such a region may provide
a variety of habitats with at least partial evolutionary
radiation. It is apparent that such a region of polymor-
phism is not necessarily indicative of the original center
of origin or of dispersal, but is a fortuitously derived
center of differentiation. It may constitute a secondary
center of evolution and of dispersal.
To summarize the hypothetical relationship in
Brunfelsia sect. Franciscea, I suggest that this
group first appeared in the Andean sector of South
America and subsequently migrated to southeast-
ern Brazil. Here the species encountered extreme-
ly favorable and diverse environments in which
they became differentiated into several additional
species. Secondary and perhaps repeated contacts
with the Andean populations occurred as the new-
ly evolved species migrated far and wide in pe-
riods of optimal climatic conditions. During un-
favorable periods, large-scale extinction of popu-
lations occurred, creating the disjunct distribu-
tions that are seen today. Section Guianenses
apparently originated in northern South America,
probably in the Guiana-northeast Amazon region,
and spread throughout the lower Amazon basin
and to the Atlantic coast of Bahia. Other popu-
lations became isolated in the Choco region, giv-
ing rise to the now disjunct B. chocoensis.
X. Taxonomic Treatment
Brunfelsia L., Sp. PL: 191. 1753, ("Brunsfelsia");
Gen. PL ed. 5:87. 1754, non Braunfelsia Par.
1894. Lectotype species designated by Knapp in
Jarvis et al. (1993): B. americana L. Based on
B. flore albo, fructu croceo, molli. Plumier, PL
Am. 12. 1703; Nov. PL Am. Gen. t. 22. 1703.
Franciscea Pohl, PL Bras. Ic. 1: 1, t. 1-7. 1826. Lec-
totype species: F. uniflora Pohl. (See note under
synonymy of section Franciscea.)
Brunfelsiopsis (Urb.) Kuntze in Post & Kuntze, Lex.
81. Dec. 1903 ("1904"). Type species: Brunfel-
sia densifolia Krug & Urb.
Evergreen shrubs or small trees, 0.3-10 m tall.
Branches few to many, naked or leafy, terete.
Branchlets terete or somewhat angled in cross
section, glabrous or pubescent, the epidermis
splitting lengthwise. Leaves alternate, with 5/13
phyllotaxy, scattered or subverticillate, simple,
exstipulate, short petiolate, blade 3-30 cm long,
0.1-5 cm wide, elliptic, oblong, lanceolate, obo-
vate to oblanceolate, rarely spathulate, ovate, lin-
ear or linear-oblong, apically acuminate, acute,
blunt or rounded, entire at margin, basally nar-
rowed, cuneate or blunt, glabrous to pilose on up-
per surface, glabrous to velutinous on lower sur-
face, firmly membranaceous to thickly coriaceous,
lateral nerves pinnate, 3-12, spreading, arcuately
anastomosing near margin, veinlets finely reticu-
late. Inflorescence a cincinnal cyme or reduced
to a single flower, terminal or axillary, subsessile,
dense and compact, rarely lax and somewhat
branched, with 1-many flowers. Bracts 1-3 per
flower, 1-10 mm long, linear to lanceolate, ca-
ducous. Flowers white turning yellow, reddish
purple, or violet-purple fading to pale violet or
white, with or without fragrance. Calyx gamosep-
alous, 3-35 mm long, tubular to campanulate, gla-
brous or variously pubescent or glandular, light to
dark green or purplish, membranaceous to sub-
coriaceous, 5-dentate, teeth 1-10 mm long, sub-
erect, subequal, valvate in bud, triangular, ovate
or lanceolate, apically acute, acuminate or trun-
cate, minutely glandular-ciliolate; calyx in fruit
persistent, enclosing capsule or present only at
base, scarcely or rarely greatly accrescent, often
becoming coriaceous and dotted with lenticels,
striately nerved. Corolla gamopetalous, hypocra-
teriform, slightly zygomorphic with bilateral sym-
metry; tube 1.5-13 cm long, 1-5 mm in diameter,
1-24 times as long as calyx, cylindric, straight or
slightly curved, fleshy, gradually dilated or briefly
inflated at apex, open at mouth or slightly con-
stricted and thickened into fleshy white ring, gla-
brous or pubescent; limb 1.5-8 cm in diameter,
5-lobed, with quincuncial-imbricate estivation,
perpendicular to tube or inclined at an angle,
plane, undulate or crispate at margin; lobes sub-
equal, obovate or broadly rounded, lateral margins
overlapping or free, rarely abruptly deflexed. Sta-
mens 4, didynamous, completely included within
corolla tube or upper pair briefly exserted; fila-
ments 2-10 mm long, inserted in uppermost part
of corolla tube; lower pair posterior, shorter; upper
pair anterior, longer, surpassing the pistil, slender,
terete or somewhat flattened, curved posterially at
apex; anthers 1-3 mm long, all fertile, rarely up-
per pair reduced or rudimentary, medifixed, sim-
ple, 1 -celled, entire or slightly bilobate, oblong or
orbicular-reniform, emarginate, dehiscing by a
single longitudinal slit toward back of tube. Ova-
ry superior, 1.5-3 mm long, sessile, subtended by
short, nectariferous disc, bilocular, with 2 axile
placentae; style filamentous, broader, and curved
backward at apex, equaling the filaments; stigma
1-2 mm long, included between anthers, suben-
tire, shallowly bilobate and subcapitate or briefly
bifid in the form of a forceps with gaping lobes.
Ovules numerous, anatropous. Fruit capsular or
subbaccate, 1-5 cm long, subglobose to ovoid, bi-
locular or unilocular by dissepimental breakdown,
valves entire, exocarp thin, membranaceous or
subcoriaceous, smooth or becoming rugose-retic-
ulate, with corky outgrowths, mesocarp thin co-
riaceous, dry at maturity, or thick, leathery or
fleshy, endocarp thin, cartilaginous, drying brittle,
sparingly dehiscent or indehiscent. Seeds few-nu-
merous, 2-13 mm long, 1-7 mm in diameter, ob-
long, concave, angular, reticulate-pitted, reddish
brown, partially immersed in fleshy placenta or
drying free. Embryo straight or slightly curved,
imbedded in endosperm; cotyledons flattened,
ovate-elliptic. Chromosome number 2n = 22, 24?
The genus Brunfelsia, as here circumscribed,
consists of 46 species of tropical American shrubs
or small trees. The genus is divided into three
sections: Brunfelsia, Guianenses, and Franciscea.
Section Brunfelsia consists of 22 species that are
restricted to the West Indian islands. These spe-
cies are not included in the present revision, but
a brief description of the section and a list of the
names of the species and synonyms are given in
Appendix II. Section Guianenses contains six spe-
cies that are found in South America, being par-
ticularly common in the Amazon basin and east-
ern Brazil. Section Franciscea consists of 18 spe-
cies that are found in tropical areas of South
America, two of which also occur in Central
Artificial Key to the Sections, Species, and Infraspecific Taxa of Brunfelsia
1 . Corolla tube 6-24 times as long as calyx, white turning yellow with age, or reddish. West Indies
1. Corolla tube 1-4 times as long as calyx, purple fading to white or pale violet, or white to greenish
white, not turning yellow. South and Central America
2. Corolla tube open at mouth, not constricted; lobes narrow, deflexed at lateral margins or scarcely
overlapping, white or greenish white Section Guianenses
3. Inflorescence sessile or with a very short peduncle; capsule dry at maturity, thin-walled
4. Pedicels elongate, 13-20 mm long; calyx glabrous 1. B. amazonica
4. Pedicels short, 3-9 mm long; calyx teeth minutely glandular-papillose 4. B. clandestina
3. Inflorescence pedunculate, the penducle occasionally very short but always visible; capsule at
maturity fleshy (or unknown)
5. Inflorescence bracts not ciliolate at margins, glabrous or very occasionally villous
6. Leaves obovate to obovate-elliptic, with 4-5 lateral nerves; calyx 7-10 mm long. Brazil
(Para), Guianas 5. B. guianensis
6. Leaves ovate, with 5-9 lateral nerves; calyx 1 1-20 mm long. Brazil (Rio Tocantins)
2. B. burchellii
5. Inflorescence bracts ciliolate at the margins, otherwise glabrous
7. Petiole 10-15 mm long; leaf blade 9-11 cm wide, with 6-7 lateral nerves; corolla limb
22-25 mm in diameter. NW Colombia, Panama 3. B. chocoensis
1. Petiole 1-4 mm long; leaf blade 4-8 cm wide, with 8-12 lateral nerves; corolla limb
15-22 mm in diameter. E Guyana 6. B. martiana
PLOWMAN: REVISION OF BRUNFELSIA
2. Corolla tube constricted at mouth forming thickened ring; lobes broadly rounded, overlapping at
lateral margins, purple or violet fading to pale violet or white Section Franciscea
8. Calyx distinctly 5-angled, inflated, drying plicate
9. Leaves cuneate to short acuminate at apex; pedicel and calyx glandular or villous
pubescent; calyx teeth ovate to lanceolate, 2-7 mm long
10. Leaves oblong-obovate to elliptic-lanceolate, the margins glabrous; calyx teeth
ovate-lanceolate, 3-7 mm long. S Brazil 12. B. cuneifolia
10. Leaves obovate to elliptic-obovate, the margins villous ciliate; calyx teeth
ovate, 2-4 mm long. Bolivia 8. B. boliviano
9. Leaves blunt to subacute at apex; calyx teeth broadly ovate, 1-4 mm long. E Brazil
20. B. obovata
11. Leaves firmly membranaceous to subcoriaceous; flowers (1)2-5; pedicels 1-8
mm long; calyx tubular, 16-22 mm long; corolla tube l-lVi times as long as
calyx 20a. B. obovata var. obovata
11. Leaves thick coriaceous; flowers (1-2)3-10; pedicels 5-12 mm long; calyx
10-16 mm long; corolla tube 1V2-3 times as long as calyx
20b. B. obovata var. coriacea
8. Calyx terete, scarcely inflated, drying flat
12. Leaves usually 16-31 cm long
13. Flowers solitary on branch tips; calyx greatly accrescent in fruit; capsule
fleshy, 4-5 cm in diameter. Pacific coast of Ecuador and Colombia
18. B. macrocarpa
13. Flowers 3 or more per inflorescence; calyx very slightly or not at all accres-
cent; capsule dry, thin-walled at maturity, less than 2 cm in diameter
14. Corolla tube l-Wi times as long as calyx; capsule completely enclosed
in calyx; calyx 1.5-3.5 cm long. SE Brazil ... 15. B. hydrangeiformis
15. Leaves subspathulate to broadly oblanceolate, reticulately rugose
with sulcate nerves; pedicel 2-4 mm long; calyx 3-6 mm in diam-
eter with yellowish brown pubescence; calyx teeth linear to linear-
lanceolate, 4-10 mm long
15a. B. hydrangeiformis subsp. hydrangeiformis
15. 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 trie homes; calyx teeth ovate to ovate-lanceolate, 2-
4 mm long 15b. B. hydrangeiformis subsp. capitata
14. Corolla tube 2-5 times as long as calyx
16. Corolla lobes deflexed at anthesis; flowers few, 4-7(20); leaves 7-12
cm wide. Amazonian Colombia and Ecuador ... II. B. chiricaspi
16. Corolla lobes spreading at anthesis; flowers (3)5-many; leaves less
than 8 cm wide
17. Leaves subverticillate, with 8-13 lateral nerves; inflorescence
dense, capituliform; corolla tube 25-38 mm long. Peru (Cuzco,
Puno), Bolivia, W Brazil 19. B. mire
17. Leaves scattered, with 5-9 lateral nerves; inflorescence lax, of-
ten short-branched; corolla tube 15-40 mm long. W South
America 14. B. grandiflora
18. Corolla tube 30-45 mm long; limb 35-52 mm across; cap-
sule 1.7-2.2 cm long
14a. B. grandiflora subsp. grandiflora
18. Corolla tube 15-30 mm long; limb 20-40 mm across; cap-
sule 11.6 cm long 14b. B. grandiflora subsp. schultesii
12. Leaves usually 2-15 cm long
19. Calyx with short or long, nonglandular trichomes
36 FIELDIANA: BOTANY
20. Flowers 8-many per inflorescence; pedicels articulated with short, per-
sistent peduncle; trichomes sparse to dense, yellowish to yellowish
brown; corolla tube 17-22 mm long. SE Brazil . . lOa. B. brasiliensis
20. Flowers all 1-3 per inflorescence; pedicels not stalked; trichomes long,
unbranched, pilose; corolla tube 25-32 mm long. S Brazil
22. B. pilosa
19. Calyx glabrous or with sparse glandular hairs
21. Corolla tube 25-40 mm long
22. Flowers solitary on branch tips; leaves thick coriaceous; capsule ru-
gulose, pericarp leathery, 1-4 mm thick. Panama . . 13. B. dwyeri
22. Flowers 1-many per inflorescence; leaves membranaceous to sub-
coriaceous; capsule smooth, pericarp thin- walled, drying brittle
23. Corolla tube 1-2 times as long as calyx; capsule completely
enclosed by calyx
24. Pedicel articulating with short peduncle; peduncle 2-5 mm
long; corolla limb 25-32 mm across; leaves 1.2-2.2 cm
wide. SE Brazil . . lOb. B. brasiliensis subsp. macrocalyx
24. Pedicels sessile or nearly so; corolla limb 40-80 mm across;
leaves 2-6.5 cm wide
25. Leaves oblong to oblong-lanceolate, with 5-11 lateral
nerves; flowers 1-11; pedicel 11-25 mm long; corolla
tube 1-1 Vi times as long as calyx. SE Brazil . . 21. B.
25. Leaves oblong-elliptic to oblong-obovate, with 6-7 lat-
eral nerves; flowers 1-2; pedicel 8-9 mm long; corolla
tube 1^2-2 times as long as calyx. E Venezuela
16. B. imatacana
23. Corolla tube 2-5 times as long as calyx; capsule exceeding the
calyx in fruit
26. Flowers 1-3(4) per inflorescence; leaves elliptic-obovate,
apically blunt to acute. S Brazil, Paraguay, Argentina ....
7. B. australis
26. Flowers (3)5-many per inflorescence; leaves oblong-lanceo-
late to lanceolate, apically acuminate. W South America
14. B. grandiflora
21. Corolla tube 15-25 mm long
27. Flowers all solitary. Widespread 24. B. uniflora
27. Flowers >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-
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
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.
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
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
FIG. 1 1 . Brunfelsia amazonica.
PLOWMAN: REVISION OF BRUNFELSIA
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-
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-
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
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.
FIG. 1 2. Brunfelsia burchellii. Reproduced courtesy of the Field Museum of Natural History.
PLOWMAN: REVISION OF BRUNFELSIA
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-
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
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
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
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-
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
BRUNFELSIA clandestiiia Plowman
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
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. 1418'S, 3902'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,
1623'S, 398'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,
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
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
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-
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
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, 55W, 5N, 14 Feb.
1966, van Donselaar 3114 (NY); ad flum. Paloe-
meu et Tapanahoni confl., 320'N, 5527'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, 253'N, 5222'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
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
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-
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
BRUNFELSIA martiana Plowman
FIG. 17. Brunfelsia martiana. Reproduced courtesy of the Botanical Museum of Harvard University.
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
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
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-
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
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
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
FIG. 18. Manaca (Brunfelsia uniflora). From Piso, W., De Medicina Brasiliensis (1648).
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,
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
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.
CHRISTABEL KING del.
PLOWMAN: REVISION OF BRUNFELSIA
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
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
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-
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 35N and 35S (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).
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
BRUNFELSIA boliviana Plowman
FIG. 21. Brunfelsia boliviana. Reproduced courtesy of the Field Museum of Natural History.
ca: mountain above Bartolo, on road from Mon-
teagudo to Sucre, 2000'S, 6445'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, 6332'W, 1912'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,
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,
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
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.
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
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-
10. Brunfelsia brasiliensis (Spreng.) L. B. Sm. &
Downs in Reitz, Fl. Ilustr. Catar., pt. 1, Sola:
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
PLOWMAN: REVISION OF BRUNFELSIA
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.
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
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-
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).
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
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
FIG. 24. Brunfehia brasiliensis subsp. brasiliensis (originally as Franciscea confertiflora). From Pohl, J. E.
Plantarum Brasiliae Icones et Descriptiones (1826).
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. Viosa, 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
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-
taao 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
calyx or up to H/2 times as long; limb 25-32 mm
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, 2227'S,
4439'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. 2346'S, long. 4620'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. &
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
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 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-
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-
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
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
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-
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
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
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
11. Brunfelsia chiricaspi Plowman, Bot. Mus.
Leafl. 23(6): 255, t. 17. 1973. Type: CO-
LOMBIA. Putumayo: Umbria, 054'N,
7610'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
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
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
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
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.
PREFEITURA MUNICIPAL DE CURITIBA
MUSEU BOTANICO MUNICIPAL
i io li -a ZT- A-
icuaa Sta. Clara d
Arbuato 2 B| flor lllas. Interior aata te galria.
PMC Off 004
PLOWMAN: REVISION OF BRUNFELSIA
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.
2706'S, 5245'W, 350-400 m, Smith & Reitz
12541 (GH, R, NY, us); Mun. Irani, Campo de Irani,
ca. 2657'S, 5150'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
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
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
13. Brunfelsia dwyeri D'Arcy, Ann. Missouri
Bot. Gard. 57:259. 1971 ("1970"). Type:
PANAMA. Panama: Cerro Jefe in "Cleusia"
FIG. 27. Brunfelsia dwyeri.
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.
25 January 1970
PLOWMAN: REVISION OF BRUNFELSIA
(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.
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-
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-
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
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-
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
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,
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-
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);
BRUNFELSIA grandiflora D.Don
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
*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. 920' S, long. 69W,
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;
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
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,
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,
348'N, 5153'W, 5 Aug. 1960, Irwin et al. 47359
(MG, NY); lower slopes and base of Mount Tipac,
0-100 m, 336'N, 5119'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
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-
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-
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
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-
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-
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
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
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
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,
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-
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).
PLOWMAN: REVISION OF BRUNFELSIA
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,
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
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
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
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
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).
Del. bl T. PlowiMB Wl - <V & . "JI
, /</; ;
PLOWMAN: REVISION OF BRUNFELSIA
mann 270 (BR), 1824, Riedel 113 (K, LE), 1816-
1821, St. Hilaire B-58 (P). Rio de Janeiro: Pico
do Papagaio, Alto da Boa Vista, Oct. 1928, Brade
s.n. (R), 29 Nov. 1928, Ducke s.n. (RB); Nova Fri-
burgo, Nov. 1842, Clamsen s.n. (F), 6 Dec. 1881,
Glaziou 14173 (c, P); Serra dos Orgaos: Terezo-
polis, 1888, Brunei s.n. (R); Estrada do Fagundes,
9 Dec. 1948, Duarte & Pereira s.n. (GH, RB);
Parque Nacional da Serra dos Orgaos entre km
8.5-9, 11 Jan. 1960, Plaster 56 (R); Picada Sau-
dade, 700 m, Jul. 1915, Lutzelburg 6495 (M); Val
do Rio Beijaflor, Mato entre Abr. 1 e 2, 1400 m,
30 Dec. 1932, Markgraf & Apparicio 10495 (RB),
U.S. Exploring Expedition s.n. (us); caminho para
Pedra do Sino, 2000 m, 2 Nov. 1952, Vidal
11-5435 (R, uc). Petropolis, Morin, 20 Jan. 1883,
Glaziou 14174 (c, F, LE, P, R, us); Quitandinha,
790 m, 29 Dec. 1939, Lutz 1551 (R); Valance, 16
Sep. 1878, Glaziou 9558 (GH); Cantagallo, 1859,
T. Peckolt 71 (BR); Serra de Estrella, 11 Dec.
1823, Riedel s.n. (LE), 1844, Weddell 818 (NY);
Itatiaia, lote 90, 8 Jan. 1947, Aparicio & Edmundo
868 (RB); Monte Serrat, 2 Jan. 1929, Campos Por-
to 1879 (B, RB); without locality, 1838, Miers s.n.
(F, P), 1844, Widgren s.n. (LD, s, us); *Serra dos
Orgaos, Gardner 563 (BM, F, G, K, NY, OXF, P, us,
w); *Municipio Santa Maria Madalena; Santa Ma-
ria Madalena, Pedra Dubois, Plowman & de Lima,
H.C., 12901 (F). Sao Paulo: Alto da Serra, Jan.
1904, Edwall 15183 (SP); Serra da Bocaina, Lutz
Boc 14 (R). Without state: Caminho de Lucina-
dos, 24 Aug. 1879, Franklin & Glaziou 5319 (R);
Glaziou 4909 (c), 8471 (c), Lobb s.n. (K), Dec.
1837, Miers s.n. (NY, us), Sellow 719 (G), Sellow
s.n. (BM, F, LE, M), Widgren 596 (s).
Brunfelsia hydrangeiformis is a beautiful shrub
of southeastern Brazil. It grows in moist, shady
primary forests between 700 and 2000 m eleva-
tion. It is now a rare plant and can be found only
in certain protected natural areas, such as the Ser-
ra da Estrella and Serra dos Orgaos in the State
of Rio de Janeiro and perhaps elsewhere where
patches of old forests have survived.
This brunfelsia shows considerable morpholog-
ical variation, and two subspecies are readily dis-
tinguished. The typical subspecies, hydrangeifor-
mis, as described by Pohl (Fig. 30) has a long,
tubular calyx that is hirsute or rarely glandular
pubescent in varying degrees. The leaves of this
species are usually broad, often subspathulate, and
deeply sulcately veined, giving a rough, reticulate
pattern to the surface. The nerves and principal
veinlets are sunken in shallow furrows on the up-
A plant related to B. hydrangeiformis was de-
scribed as Franciscea macrophylla Cham, and
Schltdl. (Chamisso and Schlechtendal, 1827) and
later transferred to B. macrophylla by Bentham
(1846). The type, collected by Sellow, was de-
stroyed in Berlin, but a photograph and fragment
are preserved at the Field Museum of Natural His-
tory in Chicago. This plant differs from B. hy-
drangeiformis in having a more lax inflorescence
in which the pedicels are elongated and articulat-
ed with a peduncle, similar to that of B. bras-
iliensis. The entire inflorescence and calyx bear a
ferruginous pubescence of sessile glandular hairs,
almost scurfy in appearance. Only one modern
collection (Williams & Assis 7928) from Minas
Gerais matches the type of this plant, which ap-
pears to be a rare aberrant form of subsp. hydran-
Brunfelsia hydrangeiformis subsp. capitata,
originally published by Bentham as B. capitata,
is a well-defined subspecies differing from sub-
species hydrangeiformis in having narrower, ob-
lanceolate leaves; longer pedicels (4-8 vs. 2-4
mm); and a glabrous or sparsely pubescent calyx
that tends to be tubular-campanulate and broader
above than at the middle. In addition, the calyx
teeth in subspecies capitata are usually ovate in-
stead of linear-lanceolate and shorter (2-4 vs. 4-
10 mm). The leaves are often smooth and lack the
rugose furrowing of the nerves.
Intermediates between the two subspecies are
not uncommon (cf. Gardner 5065) and indicate
that hybridization probably occurs between them.
The subspecies of B. hydrangeiformis are not as
well-defined, ecologically or geographically, as
those in other species of Brunfelsia. Subspecies
hydrangeiformis occurs further inland in Minas
Gerais and extends further north than subspecies
capitata. However, field studies of these plants
may reveal ecological differences between them,
such as a difference in the time of flowering. Sub-
species hydrangeiformis flowers from August to
November; subspecies capitata from October to
January, as indicated from herbarium labels.
Brunfelsia hydrangeiformis is most closely re-
lated to B. mire of Bolivia, a plant that is known
from several heterogeneous populations, some of
which are very similar to B. hydrangeiformis.
Their differences are discussed under B. mire.
Brunfelsia hydrangeiformis is perhaps the
showiest of all the South American brunfelsias
and yet has received little attention from horti-
FIG. 32. Brunfelsia imatacana. Reproduced courtesy of the Field Museum of Natural History.
culturalists. After a brief period of cultivation in
European conservatories, these plants were ne-
glected, possibly because of difficulties in propa-
gation. As indicated by its specific epithet, B. hy-
drangeiformis bears a superficial resemblance to
16. Brunfelsia imatacana Plowman, Fieldiana,
Bot. n.s. 8: 4, 1981. Type. VENEZUELA.
Bolivar: About 20 km NE of Guasipati on
road to El Miamo along side road just before
Rio Cabeza Mala, alt. 200 m, open woodland
with white sandy soil. Treelet 4-5 m tall,
flowers violet fading pure white with age,
odor of jasmine. N. v. Jazmm del monte,
Juan de la Calle. Used for banos by healers.
25 Sept. 1968. T. Plowman 1918 (holotype,
F- 18 13259; isotypes, F- 1746565, GH, K, NY, p,
uc, us, YEN). Figure 32.
Shrub or treelet 3-5 m tall. Bark reddish or
grayish brown, longitudinally and transversely
cracked, shedding in thin flakes. Branchlets about
2 mm in diameter, pubescent or glabrate, grayish
brown, longitudinally cracked. Leaves scattered
along branchlets, 65-150 mm long, 20-65 mm
wide, oblong-elliptic or elliptic-obovate, short to
long acuminate, the apex itself acute to blunt, cu-
neate to narrowed at base, glabrous or sparsely
glandular-pubescent beneath, especially at midrib,
dull dark green above, dull paler green beneath,
firmly chartaceous to subcoriaceous, the lateral
nerves 6-10, spreading, straight to somewhat
arching, indistinct; petiole short, 5-10 mm long,
1.5 mm in diameter, pubescent. Inflorescence ter-
minal, 1- or 2-flowered, subsessile. Bracts 1-3
mm long, linear-lanceolate or squamiform, pubes-
cent, caducous. Flowers light violet with a white
spot at orifice, fading to pure white with age, fra-
grant with odor resembling jasmine. Pedicel 8-9
mm long, 1 mm in diameter, glabrous, becoming
thickened, warty in fruit. Calyx 15-21 mm long,
7-10 mm in diameter, tubular, terete, glabrous or
with scattered glandular hairs, teeth 4-5 mm long,
subequal, ovate, acuminate; calyx in fruit 18-25
mm long, campanulate, thickly coriaceous, dotted
PLOWMAN: REVISION OF BRUNFELSIA
FIG. 33. Distribution of Brunfelsia hydrangeiformis (solid triangle), B. imatacana (solid square), and B. mire
with lenticels. Corolla tube 30-38 mm long, 1-2
mm in diameter, twice as long as calyx, straight,
with scattered glandular hairs, orifice 3 mm
across, lobes 13-23 mm long, subequal, rounded.
Stamens inserted in upper part of corolla tube;
filaments 1 mm wide, strap-shaped; the longer an-
terior pair 5 mm long, suberect, included in mouth
of corolla tube, the shorter posterior pair 3 mm
long; anthers orbicular-reniform, upper pair 1.5
mm across, lower pair 1.8 mm across. Ovary 1.8
mm long, 2 mm in diameter, broadly ovoid, with
7-8 ovules per locule; style 22-23 mm long; stig-
ma 1.5-2 mm long, briefly bifid. Capsule dry at
maturity and completely enclosed by coriaceous
calyx, 12-17 mm long, 14-16 mm in diameter,
ovoid to subglobose, smooth, light green, pericarp
0.5 mm thick, crustaceous. Seeds 6-10, 6-9 mm
long, 2-5 mm in diameter, oblong-ellipsoid,
somewhat angular, reddish brown, reticulate-pit-
DISTRIBUTION Restricted to the region of the
Serrania de Imataca and Serrania de Nuria in the
northeastern part of the state of Bolivar, Venezue-
la. See Figure 33.
SPECIMENS EXAMINED VENEZUELA. Boli-
var: Reserva Forestal de Imataca, along logging
road to Campamento Paraiso, 28 km NE of Upata,
ca. 100 m, 26 Sep. 1968, Plowman 1919 (ECON,
F, GH, K, s, us, YEN); low, flat woodland E of Rio
Cabeza Mala, 15 km NE of Guasipati, on road to
Miamo, 2 Jun. 1960, Steyermark 86244 (us, YEN);
Altiplanicie del Nuria, 5 km from Hato de Nuria,
E of Miamo, 400 m, 10 Jan. 1961, Steyermark
88290 (NY, YEN); more or less level forest along
pica 105, 40 km S of Tumeremo, E of highway
between Tumeremo and El Dorado, 29 km N of
El Dorado, 220 m, 23 Jul. 1960, Steyermark
86574 (NY, us, YEN); Reserva Forestal "La Para-
gua," margenes del Rio Asa, Jun. 1970, Blanco
826 (F, YEN); El Dorado, cultivated, Aug. 1957,
Trujillo 3488 (MY).
Previously, I considered B. imatacana to be a
subspecies of the related species B. pauciflora of
southeastern Brazil (Plowman, 1973, 1979). Al-
though the two species are superficially similar,
B. imatacana can be readily distinguished by the
leaf shape; the number of lateral nerves; fragrant
flowers; the shorter pedicel, calyx, and capsule;
and the smaller number and larger size of the
seeds. In addition, the two species are widely dis-
junct by a distance of 3,800 km.
Brunfelsia imatacana is endemic in the north-
easternmost part of Bolivar state in Venezuela, in
the range of low hills known as the Serrania de
Imataca and Serrania de Nuria. This area occupies
the northern rim of the Venezuelan section of the
Guayana Shield. This region has recently been
recognized by Steyermark as a distinct phytoge-
ographic unit of the flora of Venezuela. Most of
the flora found here represents species of the
Guianas at their western limits as well as various
eastern Amazonian species at their northeastern
limits of dispersal (Steyermark, 1968, 1979).
Steyermark (1968) lists 278 plant species known
only from the Serrania de Imataca and another
514 species that are restricted to the even more
limited area of the Altiplanicie de Nuria. The Im-
ataca region has been documented as a major cen-
ter of both plant and animal species endemism
and has been proposed as a Pleistocene forest ref-
uge area (Prance, 1973; Brown, 1979; Steyer-
During the collection of the type material, local
guides informed me that the foliage of B. imata-
cana was used by healers (brujos) for herbal
baths. Although this use of the plant was not ob-
served, bulk samples of roots, stems, and leaves
were collected for chemical analysis. The freshly
air-dried leaves and stems were sent to Dr. John
Leary of the Massachusetts College of Pharmacy,
Boston, for testing. Dr. Leary (pers. comm., 1973)
reported that extracts of this material yielded al-
kaloid-positive precipitates based on six different
precipitating agents. In addition, he obtained a
positive test for phytosteroids but negative tests
for flavonoids, tannins, and saponins. Dried root,
stem, and leaf material of this same sample were
also sent to Mr. Jan-Erik Lindgren of the Karolin-
ska Institutet, Stockholm, Sweden. Mr. Lindgren
(pers. comm., 1973) prepared methanol extracts
of the samples, which were examined by thin-lay-
er chromatography. He also performed a complete
alkaloid extraction on the material, followed by
gas and thin-layer chromatography on the ex-
tracts. Mr. Lindgren was unable to detect the pres-
ence of any alkaloids in the samples. It remains
uncertain what, if any, active constituents may be
present in this species. However, isolation and
identification of the active constituents in other
brunfelsias with demonstrated pharmacological
activity have proved to be similarly difficult
(Plowman, 1977; Lloyd et al., 1985).
17. Brunfelsia latifolia (Pohl) Benth. in DC.,
Prodr., 10:199. 1846. Figure 34.
Franciscea latifolia Pohl, PI. Bras. Ic. 1: 3, t. 1. 1826.
Type: BRAZIL. Rio de Janeiro: Tijuca, habitat
in umbrosis, inter frutices, Schott 6161 (holo-
Brunfelsia maritima Benth. in DC. Prodr., 10: 200.
1846. Type: BRAZIL. Rio de Janeiro: pr. Rio-
Janeiro, habitat in maritimis Taipu (holotype, c-
DC, non vidi; photograph of holotype, c; iso-
types, G-DC, non vidi).
Franciscea maritima (Benth.) Miers, Ann. Mag. Nat.
Hist, sen 2, 5: 250. 1850.
Low shrub 0.2-1.0 m tall, much branched from
near base. Branches often somewhat zigzag,
leafy. Bark thin, light brown to grayish. Branch-
lets light brown, with internodes about 1 cm long.
Leaves subsessile, variable in form, blade 4-9 cm
long, 2.2-5.5 cm wide, elliptic to oblong, ovate
or obovate, apically acute, blunt or rounded,
sometimes with a short acumen, or emarginate,
basally obtuse to cuneate, rarely truncate, gla-
brous, at times minutely pubescent beneath at
midrib, dull or shiny above, dark green, dull and
pale green beneath, firmly membranaceous to co-
riaceous, lateral nerves 5-8, mostly straight; pet-
iole 1-4 mm long, short, erect, glabrous, dark pur-
ple. Inflorescence terminal, compact, the axis 3-
7 mm long, glabrous. Flowers 3-20, pale violet
with prominent white thickening at mouth, fading
to white, scentless. Bracts 1-2 per flower, 1-10
mm long, lanceolate, ciliolate, caducous. Pedicels
6-8 mm long, slender, glabrous or sparsely glan-
dular, light green, becoming thicker in fruit. Ca-
lyx 9-13 mm long, tubular-campanulate, obovoid
in bud, glabrous or sparsely glandular, light green,
smooth, teeth 2-3 mm long; erect to recurved,
triangular to lanceolate, acute at apex, calyx in
fruit coriaceous, striately veined, enclosing fruit.
PLOWMAN: REVISION OF BRUNFELSIA
Corolla tube 15-20 mm long, 2 mm in diameter,
twice as long as calyx, sparsely glandular, pale
violet near base, white toward apex; limb 20-30
mm in diameter, spreading, plane, white beneath,
pale violet above fading to white, thickening at
mouth rounded or quadrangular, white, lobes
slightly unequal, the uppermost smaller, 5 mm
long, subtruncate at apex, the others 8 mm long,
rounded at apex. Stamens included in upper part
of corolla tube; filaments slender, cylindric, white,
upper pair 2 mm long, lower pair 1 mm long;
anthers 1 mm long, orbicular-reniform, brownish.
Ovary 2 mm long, ovoid, pale green; style slen-
der, white, curved at apex; stigma briefly bifid,
lobes unequal, light green. Capsule enclosed by
persistent calyx, 11-13 mm long, 8-10 mm in di-
ameter, ovoid, apiculate at apex, smooth, gla-
brous, dark green, pericarp thin-walled, dry at ma-
turity, crustaceous, sparingly dehiscent. Seeds
10-12, 5-6 mm long, 3 mm in diameter, oblong-
ellipsoid, angular, dark brown, reticulate-pitted.
Embryo 4 mm long, nearly straight; cotyledons
1.5 mm long, ovate.
DISTRIBUTION Brazil (Rio de Janeiro). See
SPECIMENS EXAMINED BRAZIL. Rio de Ja-
neiro: Recreio dos Bandeirantes, 22 Oct. 1938,
Alston & Lutz 138 (BM); Pedra de Itauna, BR-6,
J.U. Santos 5790 (us); Jacarepagua, Pedra de Itau-
na, 24 Mar. 1971, Araujo & Kennedy s.n. (ECON);
restinga de Jacarepagua do lado sudoeste da Pedra
de Itauna, 6 Sep. 1969, Sucre 5314 (GH, RB), 7
Sep. 1969, Sucre 5348 (GH, RB); Copacabana,
Praia Grande, 1834, Luschnath s.n. (BR). Lagao
de Piratininga, 28 Dec. 1871, Glaziou 5970 (c);
restinga de Piratininga, 8 Nov. 1922, J.G. Kuhl-
mann s.n. (RB); Barra de Marica at the restinga, 7
Nov. 1972, Rizzini s.n. (GH); prope Taipu, Apr.
1833, Riedel 1308 (c, LE); *restinga de Jacarepa-
gua, Pedra de Itauna, Sucre et al. 6428 (F), *Vidal
311 (F). Without state: Mikan s.n. (F, K, w), Rie-
del s.n. (GH).
Brunfelsia latifolia has long been misinterpret-
ed by taxonomists since Bentham's treatment of
the genus in 1846. Examination of the type
(Schott 6161), which is preserved in Vienna, and
comparison with contemporary collections make
it clear that B. latifolia is both morphologically
and ecologically distinct from any other species.
Brunfelsia maritima, a species often confused
with B. grandiflora, is clearly a synonym of B.
Brunfelsia latifolia is a diminutive shrub en-
demic to the state of Rio de Janeiro in Brazil, in
the environs of the city of Rio de Janeiro. A rare
plant, this species grows only in sandy restingas,
a maritime formation of low shrubs occurring
along the sea coast. It flowers from October to
April. Brunfelsia latifolia is distinguished by its
low habit; the rather small, elliptic leaves; and
small, pale violet flowers. It seems to be most
closely related to B. bonodora, an equally rare,
erect-growing shrub of the mountains near Rio de
Janeiro. This species has narrower, lanceolate
leaves and larger flowers than B. latifolia. Some
specimens (Mikan s.n.) are somewhat intermedi-
ate between these two species and indicate a close
relationship and possibly hybridization. However,
this remains to be determined when additional
material is available.
In the horticultural trade, the name B. latifolia
is frequently misapplied to at least two species,
B. australis and B. grandiflora. The confusion be-
tween B. australis and B. latifolia stems from
1841, when Hooker published an illustration of B.
australis that he misidentified as B. latifolia. In
reality, B. latifolia has never been cultivated as an
18. Brunfelsia macrocarpa Plowman, Bot. Mus.
Leafl. 23(6):251, t. 16. 1973. Type: ECUA-
DOR. Esmeraldas: Rio Hoja Blanca con Rio
Hualpi, bosque humedo primario, 75 m, arbol
pequena del sotobosque, de 7 m, 7 cm d.a.p.
Corteza lisa, gris, corola morada, fruto co-
mestible, guayabilla, 14 Sep. 1965, Little &
Dixon 21064 (holotype, GH; isotype, us
2639557). Figure 35.
Shrub to small tree 2-7 m tall. Branches strag-
gling, bare, with smooth grayish bark. Branchlets
somewhat crooked, terete, glabrous, yellowish
brown. Leaves sparse toward tips of the branch-
lets, blade 14-25 cm long, 6.5-11.5 cm wide,
broadly elliptic, occasionally oblong, abruptly
acuminate at apex, the acumen often subfalcate,
FIG. 34. Brunfelsia latifolia (originally as Franciscea latifolia). From Pohl, J. E., Plantarum Brasiliae Icones et
PLOWMAN: REVISION OF BRUNFELSIA
BRUNFELSIA macrocarpa Plowman
FIG. 35. Brunfelsia macrocarpa. Reproduced courtesy of the Botanical Museum of Harvard University.
blunt or abruptly cuneate at base, glabrous, dark
green above, yellowish green beneath, charta-
ceous, lateral nerves 4-6, spreading; petiole 6-17
mm long, glabrous. Inflorescence terminal at tips
of branchlets, 1 -flowered, bracteate. Flowers fra-
grant, showy, violet turning white with age.
Bracts few, 2-3 mm long, lanceolate, concave,
puberulent. Pedicel 8-10 mm long, erect, thick-
ened at apex, glabrous. Calyx 20-22 mm long,
6-8 mm in diameter, tubular, ovate in bud, weakly
inflated, more or less punctate, firmly membra-
naceous, teeth 4-8 mm long, subequal, ovate-lan-
ceolate, acute to acuminate at apex; calyx in fruit
greatly accrescent, 4-7.5 cm long, to 4 mm thick,
coriaceous, somewhat shiny, with punctate lenti-
cels. Corolla tube 40-45 mm long, 3-5 mm in
diameter, twice as long as calyx, angled in cross
section, glabrous; limb 50-55 mm in diameter,
spreading, the lobes 18-22 mm long, rounded,
subequal, the uppermost a little larger, overlap-
ping at lateral margins. Stamens included; fila-
ments subligulate, the upper pair 2 mm long, the
lower 3 mm long; anthers 2 mm in diameter, or-
bicular-reniform. Ovary 3 mm long, conical; style
36-38 mm long, slender, curved and thickened at
apex; stigma 2 mm long, briefly bifid, gaping, the
upper lobe larger. Fruit subbaccate, enclosed by
accrescent calyx, 4-5 cm long, 4-5 cm in diam-
eter, broadly ovoid or subglobose, smooth, yel-
low-ochre, pericarp 6-8 mm thick, exocarp leath-
ery, mesocarp thick, fleshy, endocarp thin, carti-
laginous, conspicuously veined. Seeds 10-20,
10-13 mm long, 5-7 mm in diameter, dark
reddish brown, reticulate-pitted. Embryo 10-11
mm long, straight; cotyledons 4-5 mm long,
DISTRIBUTION Colombia (Narino), Ecuador
(Esmeraldas). See Figure 37.
SPECIMENS EXAMINED COLOMBIA. Narino:
Isla Gorgona, 11-14 Jan. 1837, Barclay 911 (BM);
Costado Oriental, 21 Jun. 1950, Fernandez 365
(us), 7 Sep. 1924, Longfield 367 (K).
Known from only a few collections, this inter-
esting treelet occurs in the Island of Gorgona,
which lies just off the Pacific coast of southern
Colombia, and in the coastal forests of northern
Ecuador, where it grows as a small tree in the
understory of primary forests. The large, showy
flowers are said to be fragrant with the odor of
honeysuckle (Longfield 367). The fleshy yellow
fruit is edible and bears the common name guay-
abilla, or "little guava," which it resembles.
Brunfelsia macrocarpa is a very distinctive
species in having a large, fleshy, accrescent calyx
that encloses the fleshy yellow fruit. It most close-
ly resembles B. dwyeri of Panama in having large,
solitary, purple flowers that fade to white and a
thick-walled fruit. However, the fleshy fruit and
leaves are also very similar to those of B. cho-
coensis, which is placed in a different section.
This resemblance may be due to convergent evo-
lution occurring in these species, which have been
isolated for a long period in the lowland Choco
forest under the pressures of a similar and uniform
environment. No other species of section Fran-
ciscea has such a large and fleshy fruit, and B.
macrocarpa is the only known species of Brun-
felsia with a large, accrescent calyx.
19. Brunfelsia mire Monach., Phytologia 4(5):
342. 1953. Type: BOLIVIA: La Paz: Mapiri
Region, San Carlos, 850 m, baumchen 1 m,
bliiten violett, 25 Jan. 1927, Buchtien 1298
(holotype, NY; isotypes K, us 1399371 and
1399372). Figure 36.
Shrub 0.2-1.0 m tall, rarely to 2 m, mostly
unbranched. Stem to 8 mm in diameter, naked,
glabrous. Bark thin, light to dark brown, rugu-
lose. Leaves crowded toward apex of stem, sub-
verticillate, to about 6 per whorl, short petiolate
to subsessile, blade 12-28 cm long, 4-8 cm wide,
oblanceolate to oblong, rarely ovate-oblong, acute
to acuminate, often subfalcate at apex, cuneate to
blunt at base, glabrous, dull, dark green above,
pale green beneath, subcoriaceous to coriaceous,
smooth, lateral nerves 8-13, straight or somewhat
arching, spreading, rarely divaricating, veinlets
conspicuously reticulate with broad areolae; peti-
ole stout, 2-12 mm long, roughish. Inflorescence
terminal, arising from upper crown of leaves, ses-
sile, capituliform to corymbiform, usually dense,
the axis to 10 mm long. Flowers 8-30, showy,
violet fading white with age. Bracts 2-4 mm
long, lanceolate, ciliolate, caducous. Pedicel 3-10
mm long, slender, glabrous. Calyx 10-18 mm
long, 4-8 mm in diameter, tubular to tubular-cam-
panulate, glabrous, rarely sparsely glandular-pu-
bescent, reticulate-veined, teeth 3-7 mm long,
ovate to ovate-lanceolate, acuminate; calyx in
fruit subcoriaceous, the teeth becoming deeply
cut, dotted with lenticels. Corolla tube 25-38 mm
long, 2-3 mm in diameter, 2-3 times as long as
calyx, slender, glabrous or with scattered glandu-
lar pubescence; limb 25-50 mm across, spreading,
lobes to 1 5 mm long, broad, rounded, overlapping
PLOWMAN: REVISION OF BRUNFELSIA
FIG. 36. Brunfelsia mire. Reproduced courtesy of the Botanical Museum of Harvard University.
at lateral margins. Stamens included in upper part
of corolla tube; filaments slender, upper pair 4 mm
long, lower pair 3 mm long; anthers ca. 1.5 mm
long, reniform-hippocrepiform. Ovary 2 mm
long, ovoid; style filamentous, equaling the fila-
ments; stigma briefly bifid, gaping, lobes ca. 1
mm long, subequal. Fruit 13-20 mm long, 12-18
mm in diameter, subglobose to ovoid, apiculate at
apex, smooth, pericarp 0.5 mm thick, cartilagi-
nous, dry at maturity, tardily dehiscent. Seeds 10-
15, 5-6 mm long, 3 mm in diameter, oblong-el-
lipsoid, angular, dark brown, reticulate-pitted.
DISTRIBUTION Peru (Cuzco, Puno), Brazil
(Acre, Mato Grosso, Para, Rondonia), Bolivia
(Beni, Cochabamba, La Paz, Santa Cruz). See
SPECIMENS EXAMINED PERU. Cuzco: Rio
Marcopata, 20 km above Quince Mil, 780 m, 15
Jan. 1973, Madison 986 (GH); Prov. Convencion:
Echarate, 900 m, 2 Feb. 1939, Stork, Morton &
Vargas 10461 (G, K, uc); Sahuayaco 920 m, 17
Jan. 1947, Vargas 6311 (us). Prov. Paucartambo:
Pilcopata, Villa Carmen, 720 m, 17 Nov. 1964,
Vargas 15767 (us); *Prov. La Convencion; Sa-
huayaco, Rio Challpimayo near Pacchar, Plow-
man & Davis 4852 (COL, F, GH, K, IAN, MO, p, u,
us, USM); *Prov. Paucartambo; near Pilcopata,
Plowman & Davis 5000 (F, GH, K); *Prov. Pau-
cartambo, Plowman & Davis 5036 GH, K). Puno:
*near Puno, Soukup 446 (F). BRAZIL. Acre: Rio
Acre, Porvir, 14 Nov. 1923, J. G. Kuhlmann 809
(RB); Rio Acre, Monte Mo, Dec. 1911, Ule 9750
(G, K, MG). Mato Grosso: Arinos, caminho do
Porto Velho, Nov. 1914, J. G. Kuhlmann 1326 (R),
J. G. Kuhlmann 1327-k (SP, us); Utiarity (Papa-
gaio), Apr. 1918, .7. G. Kuhlmann 2287 (R, RB);
Mato proximo do Rio Sacre, 17 Nov. 1944, A. S.
Lima 7859 (SP); *Municipio de Colider; estrada
Santarem-Cuiaba, 30 km de Guarantan, Serra do
Cachimbo, /. L. Amaral et al. 803 (F); *Municipio
de Colider; estrada Santarem-Cuiaba, Serra do
Cachimbo a 30 km da cidade de Guaranta, M. N.
Silva et al. 17 (F). Para: *Alto Tapajos, Rio Cu-
ruru, Anderson 11129 (NY); *Municipio de Cas-
tanhal; Colonia 3 de Outubre, Guedes 265 (IAN);
*Belem; IPEAN, Pires & Silva 11811 (IAN). Ron-
donia: Rio Madeira, Porto Velho, 10 Jan. 1930,
Ducke s.n. (RB); 8 km NE of Porto Velho, 7 Nov.
1968, Prance et al. 8265 (wis); Machado River
region, source of Jatuaraha River, Angostura, Dec.
1931, Krukoff 1532 (NY); *proximo de Vilhena,
Vieira et al. 641 (F). BOLIVIA. Beni: *Prov.
Ballivian; Rio Chimane, environs Fatima, Davis
& Marshall 1106 (F). Cochabamba: *Prov. Cha-
pare; N of Villa Tunari, Besse et al. 513 (SEL). La
Paz: Uchimachi-Coroico, 20 Jul. 1894, Bang
2352 (BM, F, G, GH, K, LE, M, MICH, MO, NY, PH, us,
w); Hacienda Simaco sobre el camino a Tipuani,
1400 m, Feb. 1920, Buchtien 5640 (GH, us); Ha-
cienda Casano, 1,400 m, 27 Jan. 1923, Buchtien
7615 (GH, uc); near Yungas, 4000 ft, 1885, Rusby
1030 (F, NY, PH, us); Mapiri, 2,500 ft, May 1886,
Rusby 2611 (NY); Huachi, head of Beni River,
3000 ft, Aug. 1921, White 1072 (GH, NY); Beni
River, Jul. 1886, Rusby 862 (NY); *Prov. Larecaja;
Mapiri, Beck 4937 (F); *Apolo, Salinas- Yanaloma
trail, Boeke 1477 (F); *Prov. Larecaja, road from
Caranavi to Guanay, beyond Alcoche, Plowman
& Davis 5172 (F, GH, K, p, u, USM); *1.8 km SW
of Yolosa on road to Chuspipata, Solomon et al.
12022 (F); *14.3 km SW (above) Yolosa on road
to Chuspipata, Solomon et al. 12071 (F). Santa
Cruz: Lagunillas, Cordillera of Tucahuasi, 900 m,
Aug. 1934, Cardenas 2813 (F); Prov. Gutierrez,
Fraile Buenavista, 450 m, 3 Jan. 1926, Steinbach
7404 (GH, NY, s).
Brunfelsia mire is a small, unbranched shrub
rarely attaining more than 1 m in height. It grows
in humid, often swampy forests in deep shade,
mostly between 400 and 1400 m elevation. Its
area extends along the eastern flank of the Cor-
dillera from central Bolivia to Peru and north and
east into parts of Brazil. The flowers are violet
and fade to white with age; they appear between
November and February.
Still known from relatively few collections, B.
mire shows considerable variation in the size and
shape of the leaves and calyx. Populations from
central Bolivia, exemplified by the type collec-
tion, have typically large, oblanceolate leaves
with divaricating lateral nerves and a relatively
short, tubular-campanulate calyx. Populations
from Mato Grosso (/. G. Kuhlmann 1326) and
southern Peru (Stork et al. 10461) have ovate-lan-
ceolate leaves with arching, diverging lateral
nerves. Still other collections from the state of
Acre in Brazil (Ule 9750, Kuhlmann 809) and ad-
jacent Peru (Madison 986) have oblong to oblan-
ceolate leaves with spreading nerves and a longer,
narrowly tubular calyx. These last populations
strongly resemble B. hydrangeiformis of south-
eastern Brazil, a closely related species. The full
range and nature of variation within B. mire is still
incompletely known because of the paucity of
herbarium material and lack of field studies. At
this time, however, it seems best considered as a
PLOWMAN: REVISION OF BRUNFELSIA
polymorphic species exhibiting several different
phenotypes in different parts of its range.
The affinities of B. mire lie with B. chiricaspi
of Colombia and B. hydrangeiformis of south-
eastern Brazil, especially B. hydrangeiformis
subsp. capitata. The similarity between the latter
species and B. mire is striking in certain popula-
tions mentioned above and suggests that these
plants may have had a more or less continuous
distribution at some time in the past. This pattern
of distribution is also seen in B. uniflora and B.
bonodora-B. grandiflora. Subsequent differentia-
tion of populations of B. mire in western South
America has given rise to the heterogeneous com-
plex that we see today. Introgressive hybridization
may have occurred with B. grandiflora, the flow-
ers of which closely resemble some populations
of B. mire.
Brunfelsia mire may be distinguished from B.
hydrangeiformis subsp. capitata by the smoother
leaves, by the smaller calyx, and by the corolla
tube, which is exserted at least twice the length
of the calyx.
Brunfelsia mire also shows affinities with B.
chiricaspi, which also has large oblong-obovate
leaves and a corymbiform inflorescence. Brunfel-
sia mire may be distinguished primarily by the
longer corolla tube (25-38 vs. 22-25 mm), by the
spreading corolla lobes, and by usually having
many flowers per inflorescence.
The specific epithet of B. mire is taken from
the native name of the plant. It is used as a strong
medicine by the Indians of central Bolivia to ex-
pel cutaneous parasites and has been the object of
some pharmacological-pharmacognostic investi-
gation [see Plowman, 1977 Eds.].
20. Brunfelsia obovata Benth. in DC., Prodr., 10:
199. 1846. Type: BRAZIL: Minas Gerais,
near Parahybuna, 3 ft high in a marsh in wa-
ter 2 ft deep, 1841, Gardner 5064 (holotype,
K; isotypes, BM, CGE, F, G, GH, K, NY, us
Franciscea obovata (Benth.) Miers, Ann. Mag. Nat.
Hist. Sen 2, 5: 250. 1850.
Shrub to 1 m tall, much branched. Branches
spreading, naked, glabrous. Bark light brown.
Branchlets short, slender, sparsely pubescent to
villous, yellowish brown. Leaves mostly toward
the tips of branchlets, rarely scattered, blade 2-
9 cm long, 1-3.5 cm wide, obovate to oblong-
obovate, blunt to rounded at apex, at times sub-
acute or with short acumen, somewhat revolute
at margin, cuneate to narrowed at base, upper
surface usually glabrous, smooth, dull, dark
green, lower surface rugulose (in siccitate), pu-
bescent-villous, especially at midrib, yellowish
green, firmly membranaceous to thickly coria-
ceous, lateral nerves often obscure, 4-7, straight
or somewhat curved; petiole 2-5 mm long, more
or less villous, rarely glabrous. Inflorescence
terminal on current year's branchlets, sessile.
Flowers 1-10, showy, bluish to lilac, fading to
white with age. Bracts 2-10 mm long, small,
linear-lanceolate, sometimes concave, villous,
ciliolate at margin, caducous. Pedicel 2-9 mm
long, slender, thickened toward apex, angular-
grooved, glabrous, rarely glandular. Calyx 10-
22 mm long, 6-10 mm in diameter, tubular to
tubular-campanulate, oblong-ovoid in bud,
strongly inflated, 5-angled at sinuses, concave
between sinuses at lobes, glabrous, smooth, light
green, firmly membranaceous, sometimes stria-
tely nerved, teeth short, 1-4 mm long, broadly
ovate, rarely lanceolate-ovate, blunt to subacute
at apex; calyx in fruit enlarging somewhat to in-
clude capsule, nitid, coriaceous, prominently
striate-nerved. Corolla tube 20-30 mm long,
1.5-2 mm in diameter, 1-3 times as long as ca-
lyx, glandular, especially toward apex; limb 25-
45 mm long, spreading, lobes 9-16 mm long,
broadly rounded. Stamens inserted in upper part
of corolla tube; upper pair of filaments suberect,
3 mm long, anthers briefly exserted from tube;
lower pair included, 2 mm long; anthers 1.5 mm
long, reniform. Ovary 2 mm long, conical-
ovoid; style 18-23 mm long, slender; stigma 1
mm long, briefly bifid, gaping. Capsule nearly
enclosed by persistent calyx, 8-10 mm long, 8-
12 mm in diameter, subglobose, apiculate at
apex, rugose (in siccitate), greenish brown, peri-
carp coriaceous, 0.5-1 mm thick, sparingly de-
hiscent. Seeds 4-8, 4 mm long, 2.5 mm in di-
ameter, oblong, somewhat narrowed at ends, an-
gular, dark, reddish brown, reticulate-pitted. Em-
DISTRIBUTION Brazil. See Figure 37.
Key to the Varieties of Brunfelsia obovata
1 . Leaves 4-9 cm long, firmly membranaceous to
subcoriaceous; flowers (1)2-5; pedicels 1-8
mm long; calyx 16-22 mm long, tubular-in-
FIG. 37. Distribution of Brunfelsia latifolia (solid square), B. macrocarpa (solid triangle), B. obovata (open circle),
and B. pilosa (solid circle).
flated; corolla tube 1-1V2 times as long as calyx
20a. Brunfelsia obovata var. obovata
1. Leaves 2.5-7.5 cm long, coriaceous, stiff;
flowers 3-10 (rarely reduced to 1-2); pedicels
5-12 mm long; calyx tubular-campanulate; co-
rolla tube 1^-3 times as long as calyx
20b. Brunfelsia obovata var. coriacea
20a. Brunfelsia obovata var. obovata. Figure
Leaves blunt to rounded at apex, or short acu-
minate, 4-9 cm long, 1.8-3.5 cm wide, firmly
membranaceous to subcoriaceous, with 5-7 lateral
nerves. Flowers (1)2-5. Pedicels 1-8 mm long.
Calyx 16-22 mm long, tubular. Corolla tube 20-
25 mm long, 1-1 J /2 times as long as calyx.
DISTRIBUTION Brazil (Goias, Minas Gerais,
SPECIMENS EXAMINED BRAZIL. Goias:
*Municipio Alto Paraiso; Chapada dos Veadeiros,
Hatschbach & Kummrow 37257 (GH). Minas
Gerais: Dist. de Carangola, Alto da Serra da Yra-
ma, 19 Apr. 1935, /. G. Kuhlmann 141 (RB); Ca-
manducaia, margem do Rio do Sellado, 1800 m,
14 Nov. 1961, Handro 984 (SP); Sapucai-Mirim,
Serra da Mantiqueira, 1400 m, 6 Nov. 1953, M.
Kuhlmann 2926 (SP); without locality, 1816-1821,
St. Hilaire B-2051 (P). Sao Paulo: Sao Paulo,
1835, Lund 755 (c, G-DC, non vidi; photographs,
F, PH); in silvis paludosis inundatisque pr. Sao
Jose, pr. Sao Paulo, Oct.-Nov. 1833, Riedel 1467
(GH, K, LE); Est. Campo Grande (Sao Paulo Rail-
way), 17 Nov. 1893, Edwall 1981 p.p. (c, SP).
PLOWMAN: REVISION OF BRUNFELSIA
20b. Brunfelsia obovata var. coriacea J. A.
Schmidt in Mart., Fl Bras. 8(1): 259. 1864.
Type: BRAZIL: Piaui, in silivis ad fl. Gur-
guea, May 1819, Martius s.n. (holotype, M). 14
Brunfelsia silvicola Taub. in Bot. Jahrb. Syst. 21: 450.
1896. Type: BRAZIL: Goias, in silvarum mar-
ginibus in ditione fluminis superioris Maranhao,
Sep., without year, Vie 3019 (type destroyed at
Berlin). Representative specimen and possible
topotype, same locality, Sep. 1892, Vie 77 (RB-
621, P, R-66267).
Leaves 2.5-7.5 cm long, 1-3.3 cm wide, blunt
to subacute at apex, coriaceous, with 4-6 lateral
nerves. Flowers 3-10, rarely 1-2. Pedicels 5-12
mm long. Calyx 10-16 mm long, tubular-cam-
panulate. Corolla tube 20-30 mm long, 1^-3
times as long as calyx.
DISTRIBUTION Brazil (Bahia, Distrito Federal,
SPECIMENS EXAMINED BRAZIL. Bahia: Jato-
ba (Chapada), 1913, Lutzelburg 4019 (M); *Cha-
padao Occidental da Bahia, 12 km N of Corren-
tina on road to Inhaumas, Harley 21898 (F). Dis-
trito Federal: cerrado perto Corrego Guara, 5
Mar. 1962, Castellanos 23294 (R); Brasilia, Fun-
dacao Zoobotanica, 8 Sep. 1961, Heringer 8662
(SP); Sobradinho, 7 Sep. 1962, Heringer 8993/
1197 (SP, us), 21 Mar. 1963, Heringer 9096/1290
(us); Corrego Samambaia, near Taguatinga, 1100
m, 10 Sep. 1965, Irwin et al. 8152 (wis); road
Brasilia to Taguatinga, 12 Sep. 1964, Prance &
Silva 59030 (wis); ca. 5 km S of Brasilia, on road
to Belo Horizonte (Rio Gama), 1175 m, 21 Sep.
1965, Irwin, Souza & Reis dos Santos 8514 (wis);
*Brasilia, Duarte & Santos 102 (F, NY); *Nucelo
Bandeirante, Heringer 10556 (F); *Bacia do Rio
Sao Bartolomeu, Heringer et al. 4281 (F); *ca. 5
km SE of Brasilia on road to Belo Horizonte, Ir-
14 In this thesis (Plowman, 1973) and in the manu-
script we received, this taxon was cited as subspecies
coriacea stat. nov. However, in the herbarium at F, all
later aquired collections were annotated by Plowman in
the mid-1980s as variety coriacea J. A. Schmidt, indi-
cating to us that he had changed his mind as to the status
of the taxon. We have chosen to use the status of variety
as this was apparently a more recent decision of Plow-
win et al. 13126 (F); *Brasilia; Corrego de Ban-
anal, Plowman 10020 (F). Goias: entre cabeceira
do Rio das Pedras et le campement du Corto, 2
Sep. 1893, Glaziou 21920 (BR, K, LE, P, R); *Cha-
pada dos Veadeiros, Anderson 6751 (F, MO);
*Cristalina, Hatschbach 43843 (F); *Formosa,
Heringer 10719 (us).
Brunfelsia obovata was described by Bentham
in 1846 in de Candolle's Prodromus, citing a
specimen collected by George Gardner. In the
same work, however, he cited Lund 755, which
also belongs to B. obovata, as a syntype of B.
Brunfelsia obovata is a distinct species readily
recognized by the obovate, apically blunt leaves
and by the inflated, angular calyx. It is closely
allied with B. cuneifolia of southern Brazil and
has been confused with this species. Brunfelsia
cuneifolia differs in having the leaves cuneate to
acuminate at the apex; longer, lanceolate calyx
lobes; and a well-developed indument.
From an ecological standpoint, B. obovata is
somewhat specialized. It is almost always found
in aquatic or semi-aquatic environments, often
standing in water. It occurs in brejos (swampy
secondary forest), along the flooded margins of
rivers, and in humid capoeiras, from about 300 to
1800 m elevation.
Two varieties are distinguished within B. obov-
ata. The typical variety occurs in Minas Gerais
and Sao Paulo. Variety coriacea grows in the in-
terior states of Goias and Piuai and is distin-
guished by smaller, more coriaceous leaves; more
flowers (up to 10) per inflorescence; a shorter ca-
lyx (10-16 vs. 16-22 mm); and the corolla tube
relatively longer than the calyx (1^-3 vs. 1-1 ^
times as long). In addition, the calyx of var. cor-
iacea is more coriaceous and more strongly an-
gled than in var. obovata.
Few collections of B. obovata are known, and
only one (St. Hilaire B-2051) shows characters
somewhat intermediate between the two varieties,
indicating possible hybridization between them.
Variety obovata flowers in October and Novem-
ber. Its fruits are as yet unknown. One common
name, broma, has been reported for the species
Brunfelsia obovata var. coriacea corresponds
FIG. 38. Brunfelsia obovata var. obovata. From Martius, Flora Brasiliensis (1846).
PLOWMAN: REVISION OF BRUNFELSIA
THE NEW YORK BOTANICAL GARDEN
Plants of the Planalto do Brasil
< Isin nbov.i*,a ssn. c -i-iaca
.dot. w<*n la
sri >;^HtL s S-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
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
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
Brunfelsia calycina var. macrantha (Lem.) L. H. Bai-
ley & Raffill, Standard Cyclop. Hort. 1: 581.
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
pauci flora (Cham.K Schlecht] Benth.
FIG. 40. Brunfelsia pauciflora. Reproduced courtesy of the Linnean Society, London.
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,
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
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
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
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
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
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-
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. 2632'S,
5344'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
Revision of lirunl.-1-i.i (SoUnncne)
Brunfelsia pauciflora (Cham. & Schlecht.) Benth. in
DC. Prodr. JO: 199. 1846.
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
BRUNFELSIA pilosa Bowman
FIG. 43. Brunfelsia pilosa. Reproduced courtesy of the Botanical Museum of Harvard University.
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. Caador, 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. 2640'S, 5334'W,
700-750 m, Smith & Reitz 12757 (NY, R, us);
Mun. Abelardo Luz, N bank of Rio Chapeco at
Abelardo Luz, ca. 2635'S, 5220'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-
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
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
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
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
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,
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
24. Brunfelsia uniflora (Pohl) D. Don, Edin-
burgh New Philos. J. 86. July. 1829. Figure
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
 (holotype, w; isotype, PR).
Franciscea hopeana Hook., Bot. Mag. 55: t. 2829.
1828. Type: Hooker, Bot. Mag. 55: t. 2829.
Brunfelsia hopeana (Hook.) Benth. in DC., Prodr. 10:
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.
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
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-
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
PLOWMAN: REVISION OF BRUNFELSIA
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 Poo 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);
TABLE 7. Flowering and fruiting times of Brunfelsia uniflora in different parts of its geographical range.
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
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
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
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
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-
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
a number of common names that, along with an
account of the uses of the drug, are discussed in
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
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,
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.
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.
Brunfelsia multiflora Regel, Cat. PI. Hort. Aksa-
kov. 21. 1860.
Brunfelsia schomburgkiana Klotsch ex Schom-
burgk, Versuch. Fauna Fl. Br. Guy. 1155.
Brunfelsia sieberi Regel., Cat. PI. Hort. Aksakov.
Brunfelsia spinosa Jacq., Enum. Syst. PI. 14.
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.
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
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-
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
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-
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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-
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
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),
Assis 102 (brasiliensis subsp. brasiliensis), 137
(brasiliensis subsp. brasiliensis), 142 (bras-
iliensis subsp. brasiliensis), 181 (brasiliensis
Atamp. (?) 6560 (brasiliensis subsp. brasiliensis)
Bailey & Bailey 758 (brasiliensis subsp. bras-
Baker 61 (grandiflora subsp. grandiflora)
Balansa 2240 (australis)
Balgooy 1592 (australis)
Bang 1398 (grandiflora subsp. schultesii), 2352
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
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
Blanco 826 (imatacana)
Boeke 1224 (grandiflora subsp. grandiflora),
Boer 1047 (guianensis)
Bonpland 528 (australis)
Bornmuller 219 (australis)
Bowie & Cunningham 100 (hydrangeiformis
subsp. hydrangeiformis), 228 (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
(pauciflord), Oct. 1928 (hydrangeiformis subsp.
Brade & Duarte 20119 (pauciflord)
Brade & Tamandare 6576 (brasiliensis subsp.
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
British Guiana Forest Department 4422 (marti-
Brooke 5653 (boliviano)
Brunei sp. nov. 1888 (hydrangeiformis subsp.
Buchtien 1299 (grandiflora subsp. schultesii),
1300 (grandiflora subsp. schultesii), 2029
(grandiflora subsp. schultesii), 5640 (mire),
Bunting et al. 10351 (hydrangeiformis subsp. hy-
Burchell 1372 (brasiliensis subsp. brasiliensis),
8494 (burchellii), 8653 (burchellii), 8752 (bur-
Buttura 157 (australis)
Camargo 2123 (cuneifolia), 3 Oct. 1957 (cunei-
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-
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-
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-
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
Davidse et al. 11611 (brasiliensis)
Davidson 3846 (uniflora)
Davis & Marshall 1106 (mire)
Davis & Yost 958 (grandiflora subsp. schultesii)
Dawe 241 (grandiflora subsp. schultesii), Mar.
Dobereiner & Tokarnia 732 (cuneifolia), 841
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-
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-
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),
Ducke & Kuhlmann s.n. (brasiliensis subsp. bras-
Duke 11351 (chocoensis), 13331 (chocoensis),
16047 (chiricaspi), 5300 (chocoensis), 9429
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
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-
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-
Forestry Department of British Guiana 7495 (gui-
Franklin & Glaziou 5319 (hydrangeiformis subsp.
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),
Garcia Barriga 4586 (grandiflora subsp. schul-
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),
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.
Goes & Constantino 543 (brasiliensis subsp.
brasiliensis), 883 (brasiliensis subsp. bras-
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
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)
Marling et al. 7021 (chiricaspi), 7151 (chiricas-
pi), 7474 (grandiflora subsp. schultesii), 7700
(grandiflora subsp. schultesii), 7772 (chiricas-
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
Hatschbach & Cervi 46844 (pauciflora)
Hatschbach & Guimaraes 15286 (pilosa), 25473
(pilosa), 30302 (cuneifolia), 45004 (brasiliensis
Hatschbach & Kummrow 37257 (obovata var.
obovata), 45569 (brasiliensis subsp. bras-
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
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
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-
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),
PLOWMAN: REVISION OF BRUNFELSIA
Krapovickas & Schinini 31476 (boliviano), 35155
(grandiflora subsp. schultesii)
Krapovickas et al. 26092 (australis)
Krukoff 1532 (mire), 5284 (grandiflora subsp.
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
Kummrow 1575 (pauciflora), 2420 (pauciflora),
2432 (cuneifolia), 94 (pilosa), 970 (brasiliensis
Lalandes s.n. (brasiliensis subsp. brasiliensis)
Landrum et al. 4150 (brasiliensis subsp. bras-
Langsdorff 43 (hydrangeiformis subsp. hydran-
geiformis), s.n. (brasiliensis subsp. bras-
iliensis), s.n. (hydrangeiformis subsp. hydran-
Lanjouw 345 (guianensis)
Lanjouw & Lindeman 2137 (guianensis), 2564
(guianensis), 2888 (guianensis)
Leite 3660 (pauciflora), 4188 (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.
Lindberg 203 (uniflora)
Lindeman 5889 (guianensis), 6069 (guianensis),
Lindeman & Haas 1174 (brasiliensis subsp. mac-
rocalyx), 3248 (pilosa), 5157 (brasiliensis
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),
Lofgren & Cowan 1660 (pauciflora)
Lofgren & Edwall 1435 (uniflora), 2705 (pauci-
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.
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-
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
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-
Magalhaes 1643 (hydrangeiformis subsp. hydran-
Maguire 23894 (guianensis)
Markgraf 3495 (brasiliensis subsp. brasiliensis)
Markgraf & Apparicio 10495 (hydrangeiformis
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)
Mattos Silva et al. 357 (clandestina)
Mattos, J., & Mattos, N., 14284 (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-
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.
Meyer 12621 (australis), 166 (australis), 3076
Miers 4495 (brasiliensis subsp. brasiliensis),
4721 (uniflora), Dec. 1837 (hydrangeiformis
subsp. capitata), Feb. 1838 (brasiliensis subsp.
macrocalyx), s.n. 1838 (hydrangeiformis subsp.
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
Mori 6828 (chocoensis)
Mori & Bolten 8394 (guianensis), 8575 (guianen-
Mori & Boom 14726 (guianensis), 14834 (gui-
Mori et al. 12082 (clandestina), 8767 (guianen-
sis), 9775 (clandestina)
Moura s.n., 1887 (hydrangeiformis subsp. hy-
Mosen 673 (uniflora)
Mueller 435 (pauciflora)
Muniez 99 (australis)
Neill 6546 (grandiflora subsp. grandiflora)
Neill & Palacios 7101 (grandiflora subsp. gran-
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-
Pabst & Pereira 6023 (uniflora)
Paiva & Stehmann s.n. (australis)
Palacios et al. 3467 (brasiliensis subsp. bras-
Palmer 155 (australis)
Pearce 290 (grandiflora subsp. grandiflora), 295
(grandiflora subsp. grandiflora), Mar. 1868
(grandiflora subsp. grandiflora)
Peckolt, O., Aug. 1934 (hydrangeiformis subsp.
Peckolt, T., 71 (hydrangeiformis subsp. capitata)
Pedersen 418 (australis), 418a (australis)
Pereira 6066 (pauciflora), 9241 (hydrangeiformis
subsp. hydrangeiformis), 9451 (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
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.
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.
Poeppig 2206 (grandiflora subsp. schultesii)
Pohl 263 (brasiliensis subsp. brasiliensis), 264
(uniflord), 265 (brasiliensis subsp. bras-
iliensis), 266 (brasiliensis subsp. brasiliensis),
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
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-
Reiss 99 (pilosa)
Reitz 2743 (pauciflord), 2768 (pauciflord), 3985
(brasiliensis subsp. macrocalyx), 4375 (bras-
iliensis subsp. macrocalyx), C-171 (pilosa), C-
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-
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-
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-
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-
Santos Lima Aug. 1940 (pauciflord)
Santos-Lima 200 (hydrangeiformis subsp. hydran-
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),
Schultes & Cabrera 19115 (grandiflora subsp.
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)
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),
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
Souza et al. 521 (brasiliensis subsp. brasiliensis)
Spruce 1495 (amazonica), 3973 (grandiflora
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-
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-
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.
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
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-
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
Vauthier 42 (brasiliensis subsp. brasiliensis), 541
(brasiliensis subsp. brasiliensis)
Venturi 4408 (australis)
Vert 6152 (pauciflora)
Vianna et al. 1574 (brasiliensis subsp. bras-
Vidal 311 (latifolia), 11-5435 (hydrangeiformis
subsp. capitata), 11-5784 (brasiliensis subsp.
brasiliensis), s.n. (brasiliensis subsp. bras-
Vieira et al. 641 (mire)
PLOWMAN: REVISION OF BRUNFELSIA
Vincent s.n. (australis), s.n. (brasiliensis subsp.
Vogel 677 (brasiliensis subsp. macrocalyx)
Warming 103 (brasiliensis subsp. brasiliensis)
Wawra & Maly 426 (brasiliensis subsp. bras-
Weberbauer 6151 (grandiflora subsp. grandiflo-
Weddell 1648 (brasiliensis subsp. brasiliensis),
3621 (boliviano), 818 (hydrangeiformis subsp.
capitatd), 878 (hydrangeiformis subsp. hydran-
geiformis), s.n. 1844 (uniflord), s.n. 1858 (un-
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.
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-
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.
Wullschlagel 1505 (guianensis)
Ybarrola 2963 (australis), 3270 (australis), 3430
Zarucchi & Andrade 2316 (grandiflora subsp.
Zaruma et al. 56 (chiricaspi)
Zernys s.n. (brasiliensis subsp. macrocalyx)
Appendix II: Brunfelsia sect.
Brunfelsia L. sect. Brunfelsia. Type species: B.
Brunfelsia L., Sp. PL 191. 1753. Type species: B.
Brunfelsia L. sect. Eubrunfelsia Griseb, Fl. Brit.
W. Ind. 432. 1861. Type species: B. americana
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-
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
Brunfelsia grisebachii Amshoff, Contrib. Ocas.
17 This Caribbean group of brunfelsias is being revised
at present by Victor Fuentes Fiallo of INIFAT, Havana,
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.
Brunfelsia maliformis Urb., Symb. Ant. 3: 372.
1903. (B. fawcettii Urb, Symb. Ant. 3: 371.
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.
Brunfelsia pluriflora Urb., Symb. Ant. 9: 252.
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.
Brunfelsia undulata Sw., Nov. Gen. Spec. PL 90.
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-
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,
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.
TANNER, E. V. J., AND V. KAPOS. 1982. Leaf structure of
Jamaican upper montane rain-forest trees. Biotropica,
THYR, B. D., M. J. SAMUEL, AND P. G. BROWN. 1975.
New solanaceous host records for Corynebacterium
michiganense. Plant Disease Reporter, 59(7): 595-
TOKARNIA, C. H., A. GAVA, L. STOLF, AND P. V. PEIXOTO.
1991. Experimental poisoning in cattle by Brunfelsia
pauciflora (Solanaceae). Pesquisa Veterinaria Brasi-
TRIGO, M. M. 1992. Contribution to the study of pollen
in ornamental species. Solanaceae, Convolvulaceae
and Hydrophyllaceae. Acta Botanica Malacitana, 17:
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
Synonyms and non-English common names are italicized. New names or combinations are in bold-
face. Page numbers for illustrations are also in boldface.
Banisteriopsis caapi vii
Besleria bonodora 59, 60
Besleria inodora 103
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
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
Conejo, village vii
edible fruits 43, 94
erva de macaco 69
evolutionary trends 26-30
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
Guianenses, section 38
hybridization 19-20, 83, 88, 98
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,
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
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
Schultes, R. E. vii
Schultesianthus megalandrus 76
section Franciscea 51
section Guianenses 38
specialization, trends 26-30
thunderstorm flower 74
UNIVERSITY OF ILLINOIS-URBANA