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Full text of "A revision of the South American species of Brunfelsia (Solanaceae)"

UNIVERSITY OF 

ILLINOIS LIBRARY 

AT URBANA-CHAMPAIGN 

BIOLOGY 

'2001 



5otany 

EW SERIES, NO. 39 



L Revision of the South American 
Ipecies of Brunfelsia (Solanaceae) 

imothy C. Plowmant 

, Knapp and J. R. Press, Editors 



ugust 31, 1998 
ublication 1496 






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FIELDIANA 



Botany 

NEW SERIES, NO. 39 



A Revision of the South American 
Species of Brunfelsia (Solanaceae) 

Timothy C. Plowmant 



Department of Botany 
Field Museum of Natural History 
Roosevelt Road at Lake Shore Drive 
Chicago, Illinois 60605-2496 U.S.A. 



S. Knapp and J. R. Press, Editors 

Department of Botany 
The Natural History Museum 
London SW7 5BD 
England 



Accepted July 2, 1996 
Published August 31, 1998 
Publication 1496 



PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY 



1998 Field Museum of Natural History 

ISSN 0015-0746 
PRINTED IN THE UNITED STATES OF AMERICA 



n,S 
Table of Contents 



FOREWORD 

Warfe Davis vii 

EDITORS' PREFACE 

S. Knapp and J. R. Press xi 

I. MATERIALS AND METHODS 1 

II. TAXONOMIC HISTORY OF THE GENUS 2 

III. GENERIC RELATIONSHIPS 5 

IV. GENERAL MORPHOLOGY 6 

V. CYTOLOGY 16 

VI. HYBRIDIZATION EXPERIMENTS 19 

VII. ECOLOGY AND REPRODUCTIVE 

BIOLOGY 20 

VIII. TRENDS IN SPECIALIZATION 26 

IX. GEOGRAPHICAL DISTRIBUTION AND 

RELATIONSHIPS 29 

X. TAXONOMIC TREATMENT 34 

Brunfelsia Linnaeus 34 

Artificial Key to the Sections, Species, 
and Infraspecific Taxa of 

Brunfelsia 35 

Brunfelsia sect. Guianenses 38 

1. Brunfelsia amazonica 38 

2. Brunfelsia burchellii 40 

3. Brunfelsia chocoensis 40 

4. Brunfelsia clandestina 43 

5. Brunfelsia guianensis 46 

6. Brunfelsia martiana 49 

Brunfelsia sect. Franciscea 51 

1. Brunfelsia australis 53 

8. Brunfelsia boliviano 57 

9. Brunfelsia bonodora 59 

10. Brunfelsia brasiliensis 61 

lOa. Subspecies brasiliensis 62 

lOb. Subspecies macrocalyx 66 

1 1 . Brunfelsia chiricaspi 70 

12. Brunfelsia cuneifolia 72 

13. Brunfelsia dwyeri 74 

14. Brunfelsia grandiflora 77 

14a. Subspecies grandiflora 78 

14b. Subspecies schultesii 80 

15. Brunfelsia hydrangeiformis 84 

15a. Subspecies hydrangeiformis 84 
15b. Subspecies capitata comb, et 

stat. nov. 86 

16. Brunfelsia imatacana 89 

17. Brunfelsia latifolia 91 

18. Brunfelsia macrocarpa 93 

19. Brunfelsia mire 95 

20. Brunfelsia obovata 98 

20a. Variety obovata 99 

20b. Variety coriacea 101 



2 1 . Brunfelsia pauciflora 103 

22. Brunfelsia pilosa 108 

23. Brunfelsia rupestris Ill 

24. Brunfelsia uniflora 113 

Dubious and Excluded Species 119 

Nomina Nuda 119 

XI. EDITORS' ACKNOWLEDGMENTS 119 

XII. PLOWMAN'S ACKNOWLEDGMENTS 120 

XIII. LITERATURE CITED 120 

APPENDIX I: List of Exsiccatae 124 

APPENDIX II: Brunfelsia sect. Brunfelsia 132 

Description 132 

Names of Brunfelsia sect. Brunfelsia 132 

APPENDIX III: Additional References 133 

INDEX . 134 



List of Illustrations 



Timothy Plowman vi 

Brunfelsia hydrangeiformis subsp. hydran- 
geiformis, type xii 

1. Brunfelsia americana 3 

2. Root of Brunfelsia grandiflora subsp. 
schultesii 8 

3 . Floral structure of Brunfelsia 11 

4. Brunfelsia densifolia (Knapp s.n.) from 
Puerto Rico 12 

5. Brunfelsia lactea (Knapp s.n.) from 
Puerto Rico 14 

6. Fruit of Brunfelsia americana 17 

7. Methona themisto, type specimen 24 

8. Caterpillars of Methona confusa on 
Brunfelsia chiricaspi leaf 25 

9. Distribution of section Brunfelsia 31 

10. Distribution of sections Guianenses 

and Franciscea 32 

1 1 . Brunfelsia amazonica 39 

12. Brunfelsia burchellii 41 

13. Brunfelsia chocoensis 42 

14. Brunfelsia clandestina 44 

15. Distribution of species in section 
Guianenses 46 

16. Brunfelsia guianensis 47 

17. Brunfelsia martiana 50 

18. Manaca (Brunfelsia uniflora) 52 

19. Brunfelsia australis 55 

20. Distribution of Brunfelsia australis, 

B. boliviana, and B. bonodora 56 

21. Brunfelsia boliviana 58 

22. Brunfelsia bonodora, illustration of 
lectotype 60 



in 



23. Distribution of Brunfelsia brasiliensis 

and B. chiricaspi 63 

24. Brunfelsia brasiliensis subsp. bras- 
iliensis 64 

25. Brunfelsia chiricaspi 71 

26. Brunfelsia cuneifolia 73 

27. Brunfelsia dwyeri 75 

28. Distribution of Brunfelsia cuneifolia, 

B. dwyeri, and B. grandiflora 77 

29. Brunfelsia grandiflora subsp. grandiflo- 
ra and subsp. schultesii 79 

30. Brunfelsia hydrangeiformis subsp. hy- 
drangeiformis, illustration of type 85 

3 1 . Isotype of Brunfelsia hydrangeiformis 
subsp. capitata stat. nov 87 

32. Brunfelsia imatacana 89 

33. Distribution of Brunfelsia hydrangei- 
formis, B. imatacana, and B. mire 90 

34. Brunfelsia latifolia, illustration of 

type 92 

35. Brunfelsia macrocarpa 94 

36. Brunfelsia mire 96 

37. Distribution of Brunfelsia latifolia, B. 
macrocarpa, B. obovata, and B. 

pilosa 99 

38. Brunfelsia obovata var. obovata 100 



39. Brunfelsia obovata var. coriacea 102 

40. Brunfelsia pauciflora 104 

41. Distribution of Brunfelsia pauciflora 

and B. rupestris 105 

42. Brunfelsia pauciflora cv. "Lindeniana" 109 

43. Brunfelsia pilosa 110 

44. Brunfelsia rupestris 112 

45. Brunfelsia uniflora, illustration of type 114 

46. Distribution of Brunfelsia uniflora 116 



List of Tables 



1 . Historical classification of Brunfelsia .... 6 

2. Chromosome counts in Brunfelsia 18 

3. Chromosome numbers and size in Brun- 
felsia 19 

4. Pollinators of Brunfelsia grandiflora 23 

5. Brunfelsia species recorded as larval food 
plants for species of Methona 26 

6. Trends in specialization of Brunfelsia ... 30 

7. Flowering and fruiting times of Brunfel- 
sia uniflora 117 



IV 




Tim Plowman 



Foreword 



The idea for this monograph emerged from 
somewhat unusual circumstances. On March 29, 
1942, ethnobotanist and plant explorer Richard 
Evans Schultes arrived at the Kofan Indian village 
of Conejo, on the banks of the Rio Sucumbios in 
the remote rain forests of southern Colombia. In 
the midst of his first sojourn in the Amazon, with 
the goal of surveying medicinal plants and arrow 
poisons, Schultes found himself immersed in a 
world of phytochemical wizardry unlike anything 
he had ever known. Psychoactive and toxic plants 
touched every aspect of the lives of the Kofan. 
The most important and sacred of their plants was 
the hallucinogen Banisteriopsis caapi, or yage, 
the "vine of the soul." When Schultes inquired 
how often the people imbibed yage, his infor- 
mant's response suggested that the question had 
no meaning. During illness, of course, and in the 
wake of a death. In times of need or hardship. At 
certain passages in life. When a young body of 
six has his initial haircut, or when he kills for the 
first time. And naturally, the shaman suggested, a 
youth will drink yage at puberty, when his nose 
and ears are pierced and he obtains the right to 
wear the tail feathers of the macaw. As a young 
man, he may drink it at his leisure to improve his 
hunting technique or simply to flaunt his physical 
prowess. The message that Schultes received was 
that the Kofan took yage whenever they felt like 
it. At least once a week, and no doubt on any 
occasion that warranted it, such as the eve of his 
own departure from Conejo a fortnight later. With 
the people dancing, the men facing the women in 
long lines moving forward and then back, turning 
from side to side, the dancers stamping the ground 
lightly in time with the drums, this solitary stu- 
dent of plants, a man who in time would inspire 
a generation of ethnobotanists, took yage with, as 
he would recall 50 years later, "the whole damn 
village." 

The next morning, his face still painted and his 
mind swirling with the sounds of chants, Schultes 
paddled up the Sucumbios in a dugout canoe. His 
head throbbed. Unbeknownst to him, the prepa- 
ration had contained bark of tsontinb"k'o, the cold 
fever tree, together with su-tim-ba-che, a root said 
by the Kofan to cause a "drunk worse than yage." 
Indeed, it had. Despite his discomfort, Schultes 
made a note to investigate the admixtures at 
length. Thirty years later he did, by dispatching 
his protege, Timothy Plowman, to the Putumayo. 



After 15 months in the field in South and Central 
America and the Caribbean, Tim Plowman 
emerged with a definitive study, a monographic 
revision of the genus Brunfelsia, including de- 
scriptions of several new species and varieties. 
The plant that Schultes drank at Conejo and that 
gave him such a headache is now known as Brun- 
felsia grandiflora subsp. schultesii. 

Tim Plowman approached botanical research 
with the intensity and rigor of a scientist, the cour- 
age of his mentor, and the flair of a poet. His first 
Colombian excursion set the pattern of his career. 
Arriving in Bogota, he traveled south to the valley 
of Sibundoy, where he contacted Pedro Juajibioy, 
a Kamsa Indian healer who as a boy guided 
Schultes into the upper Putumayo. Tim and Pedro 
retraced that journey to reach a Kofan settlement 
on the Rio Guamues, also visited by Schultes in 
1942. The Kofan recognized B. grandiflora as 
tsontinba"k"d. Another of Schultes's students, 
Homer Pinkley, who spent a year among the tribe 
in 1965, had written that their shaman occasion- 
ally ingested the plant in order to diagnose dis- 
ease. This observation supported other reports 
dating back to the 19th century that suggested the 
plant might be hallucinogenic. Plowman wanted 
to know. At Santa Rosa on the Rio Guamues he 
found B. grandiflora commonly cultivated in 
house yards, but he also met an old shaman who 
brought him from the forest a rare but related 
plant named for the tapir because of its inordinate 
strength as a drug. Plowman immediately recog- 
nized the plant as a new species, which he later 
named Brunfelsia chiricaspi after the Quechua 
word meaning cold tree. He asked the old man to 
prepare it, but the shaman refused. He described 
the plant as a dangerous messenger of the forest 
and disavowed any knowledge of taking it for vi- 
sions. Tim persisted. Eventually the shaman 
agreed, although reluctantly and only on the con- 
dition that Pedro also drink the preparation. 

The drug, an extract from the bark, was murky 
brown and bitter to swallow. Tim felt the effects 
within ten minutes: a tingling sensation such as 
one feels when the blood rushes back to a limb 
that has fallen asleep. Only in this case the sen- 
sation grew to a maddening intensity, spreading 
from his lips and fingertips toward the center of 
his body, progressing up his spine to the base of 
his skull in waves of cold that flooded over his 
consciousness. His breathing collapsed. Dizzy 



vn 



with vertigo, he lost all muscular control and fell 
to the mud floor of the shaman's hut. In horror he 
realized that he was frothing at the mouth. An 
hour passed. Paralyzed and tormented by an ex- 
cruciating pain in his stomach, he remained only 
vaguely aware of where he was on the earth, 
face to face with three snarling dogs fighting over 
the vomit that had spread in a pool around his 
head. 

The shaman, noticing his plight, did what sha- 
mans normally do under such circumstances: He 
went to bed. Desperate to escape the sensation, 
half-blinded by the drug, and incapable of walk- 
ing, Tim and Pedro stumbled and crawled through 
the forest for two hours until finally, toward dawn, 
they reached the village of San Antonio, where 
they were staying in an abandoned jail. As light 
came to the forest, they crawled into their ham- 
mocks, where they remained, motionless, for two 
days. Pedro Juajibioy, whose experience as a folk 
healer had taken him on a thousand flights of the 
spirit, summed up the experience succinctly: "The 
world was spinning around me like a great blue 
wheel. I felt that I was going to die." 

This story reveals something of the character 
of this remarkable young scholar, a man of ad- 
venture, grace, generosity, kindness, modesty, and 
honor, whose academic achievements in time 
might well have surpassed even those of his fa- 
mous mentor, Richard Evans Schultes, the "Fa- 
ther of Ethnobotany." Sadly, Tim's life was cut 
short by cruel circumstances. His untimely death 
at the age of 45 from AIDS on January 7, 1989, 
severed a career of immense promise. Already far 
on the way as one of the most discerning, original, 
and effective naturalists of the century, Tim was 
a gentleman, a friend of everyone, an understand- 
ing and devoted teacher, an intellectual of extraor- 
dinary depth, and a tireless and demanding re- 
searcher happy to share his experience and coun- 
sel with whomever sought his advice. 

Tim Plowman's interest in and love of plants 
developed when he was a child growing up in the 
temperate woodlands surrounding Harrisburg, 
Pennsylvania. An avid collector even as a boy, his 
passion for plants grew into the central metaphor 
of his life. After attending college at Cornell Uni- 
versity, he went as a graduate student to the Bo- 
tanical Museum of Harvard University. Such was 
his promise that even before enrolling in graduate 
school, Tim was dispatched by Schultes to the 
Amazon on an expedition that would define the 
course of his professional life. In the fall of 1966 
Tim returned from Brazil flush with excitement 



and fully committed to spending the rest of his 
life in pursuit of the mysteries of the tropical rain 
forest. 

His dissertation, with its thorough bibliographic 
research, precise and deliberate taxonomic and 
ethnobotanical insights, elegant prose, and exqui- 
site line drawings, set the standard of excellence 
by which he would operate throughout his pro- 
fessional career. By the time his Ph.D. was offi- 
cially conferred, in 1974, Tim was already deeply 
involved in the project for which he will always 
be remembered, a 15 -year effort to decipher the 
complex taxonomy of Erythroxylum and to study 
the ethnobotany of coca, the sacred leaf of the 
Andes and the notorious source of cocaine. Of 
Tim's 80 published scientific papers, 46 are relat- 
ed to his work on Erythroxylum, and his position 
as the world's authority on the genus enabled him 
to speak eloquently and powerfully in defense of 
the traditional use of coca by beleaguered indig- 
enous peoples of the Andes and northwest Ama- 
zon. 

In 1978 Tim left Harvard for the Field Museum 
of Natural History, where he became tenured in 
1983 and was appointed curator in 1988. If Tim 
grew up at the Botanical Museum at Harvard, he 
came into his own at the Field Museum, and his 
years there were both the happiest and most pro- 
ductive of his remarkable career. His interdisci- 
plinary interests in systematics, ethnobotany, and 
ethnopharmacology led him to interact with an in- 
creasingly diverse group of scholars that included 
not only fellow botanists but also archaeologists, 
phytochemists, ethnographers, and pharmacolo- 
gists. In addition to carrying out an active scien- 
tific research program as co-principal investigator 
of the National Science Foundations "Projeto Flo- 
ra Amazonica," he served on the editorial boards 
of numerous journals, including Flora Neotropi- 
cal Monographs, Advances in Economic Botany, 
Journal of Psychoactive Drugs, and Journal of 
Ethnopharmacology. Between 1984 and 1988 he 
was co-editor-in-chief of the Journal of Ethno- 
pharmacology and the scientific editor of Fieldi- 
ana. He was vice president of the Beneficial Plant 
Research Association, a fellow of the Linnean So- 
ciety, and a member of many professional soci- 
eties, including the American Society of Plant 
Taxonomists, the Society of Economic Botany, 
the Council of Biology Editors, the Society of 
Ethnobiology, and the New England Botanical 
Club. As chairman of the botany department of 
the Field Museum of Natural History (1986- 
1988), Tim secured a substantial increase in Na- 



Vlll 



tional Science Foundation funding for the herbar- 
ium and developed a new facility for the curation 
of economic collections. His enthusiasm, spirit of 
cooperation, professional rigor, and passionate 
commitment to botany proved infectious, and un- 
der his leadership, morale within the botany de- 
partment soared. 

Credentials alone, however, present but a shad- 
ow of the man who affected so many lives in such 
profound ways. For Tim, life was but a vehicle 
for seeking understanding and for expressing free- 
dom. If there is a word to describe Timothy Plow- 
man it would be freedom, and he lived with the 
conviction that every person had the right to pur- 
sue his or her own path unshackled by the burdens 
of social convention. Equally at ease in the tran- 
quil world of plants and the society of people, 
Tim was charismatic, and those privileged to have 
spent time with him often developed a respect that 
bordered on reverence, for he was a true renais- 
sance scholar, a man out of time, whose breadth 
of interests and passions went far beyond the 
boundaries of his beloved field of botany. 

But it is as a botanist and intrepid plant explorer 
that Tim will be best remembered. He spent more 
than five years of his life in the most remote and 
inhospitable regions of the Andes and Amazon, 
making more than 15,000 collections of unsur- 
passed quality. He always considered his time in 
the field a privilege, and he never failed to re- 
member his fellow botanists toiling away in the 
less romantic confines of the herbaria. Tim 
seemed to have a Rolodex in his head that re- 
corded the name of every specialist in every 
group of plants, and he was constantly on the 
lookout for specimens that might prove useful to 
a distant colleague. He collected everything. His 
voucher specimens were not only complete but 
aesthetically beautiful, and whenever possible he 
augmented them with invaluable collections of 
live material. Living plants, many new to science 
and collected first by Tim, may be found in bo- 
tanical gardens throughout the world. 



In the rain forests of the Amazon Tim felt the 
fullness of life. He marveled at the thousand 
themes, the infinitude of form, shape, and texture 
that so clearly mocked the terminology of tem- 
perate botany. He always traveled in the forest as 
a student, and his commitment to ethnobotany 
grew in part from his direct experience with the 
indigenous peoples who understood the plants in 
ways that he believed he could only hope to em- 
ulate. To be in the forest, he said, was to be in 
Eden, and to say the names of the plants was to 
recite the names of the gods. He believed that all 
forms of life were manifestations of the sacred. 
Hence, for Tim, biological and cultural diversity 
represented far more than the foundation of sta- 
bility; they were articles of faith, fundamental 
truths that indicated the way things were supposed 
to be. 

Tim had a special affinity for Indians, and his 
uncanny ability to gain their trust and confidence 
was one measure of the deep respect he had for 
their way of life. He empathized with their world 
view, which defined man as but one element in- 
extricably linked to the whole of creation. It was 
this unique cosmological perspective, he believed, 
that enabled the Indians to comprehend implicitly 
the intricate ecological balance of the forest he 
loved so dearly. Tim viewed with pain, dismay, 
and increasing anger the other world view, in 
which man stands apart, that now threatens the 
forest with devastation. It was one of his fondest 
hopes that the lessons of ethnobotany might ulti- 
mately facilitate a dialogue between these two 
world views such that folk wisdom might temper 
and guide the inevitable development processes 
that today ride roughshod over much of Earth. 
The many of us who loved him as a brother and 
respected him as a colleague can do no better ser- 
vice to his memory than continue our own strug- 
gles to make his dream a reality. 

Wade Davis 



IX 



Editors' Preface 



Tim Plowman not only was a "botanist's bot- 
anist," he also inspired a whole generation of 
young taxonomists with his enthusiasm for plants 
in both the field and herbarium. 

The study of the systematics of the South 
American species of Brunfelsia was the subject of 
his Ph.D. research while he was at Harvard Uni- 
versity. After receiving his Ph.D. in 1974, Tim 
worked mainly on Erythroxylum, but Brunfelsia 
remained an area of special interest for him. He 
was preparing a major revision of the group, using 
his dissertation as a basis and backing up with 
further study and extensive field work, when he 
died in 1989. 

In 1994 Tim's colleagues at the Field Museum 
of Natural History in Chicago sent the manu- 
script, together with various notes and archival 
material, to one of us (S.K.) to assess whether the 
work could be brought to completion. We are very 
pleased to have been able to do this, both because 
it will make available this extensive study of an 
interesting group of Solanaceae and because it 
will serve as a tribute to the work of a well-liked 
and respected colleague. 

In taking the working manuscript and bringing 
it to publication, we tried throughout to keep in 
mind Tim Plowman's concept of the group and 
changed his work as little as possible. Most of the 
manuscript remains essentially unaltered, but 
where necessary we rearranged some sections and 
augmented others in line with Tim's own publi- 
cations, notes, and records. The only major de- 
parture from this approach is where we felt a need 
to update the manuscript in the light of works 
published since Dr. Plowman's death. Such up- 
dates usually take the form of explanatory notes 
in the text that are clearly indicated as being ed- 
itors' contributions. 



To this end, the key was revised to incorporate 
additional species published (by Plowman) since 
the original manuscript was written. The order of 
specimen citations is Plowman's. In some places 
reasoning is not clear, but we chose not to change 
the order. Specimens not cited in the original work 
but seen by Dr. Plowman and recorded in his 
notes and in the herbarium at the Field Museum 
were added; they are marked with an asterisk and 
are inserted in order of collector at the end of each 
state or province. These later records are included 
in the distribution maps. Infraspecific taxa are not 
indicated on the maps because Plowman's later 
annotations were only to specific rank and could 
not be integrated with distributions of infraspecif- 
ic taxa. Additional references, some of them 
Plowman's own subsequent publications, were in- 
serted; like the additional specimens, these are 
marked in the bibliography with an asterisk. 

Several new combinations remained unpub- 
lished at the time of Tim's death and are validated 
here. The authority for these combinations should 
be simply Plowman, as they represent Tim's as- 
sessment of the status and rank of the taxa and 
should be attributed solely to him. 

The additional bibliography in Appendix II lists 
references to publications that postdate the origi- 
nal manuscript and that we believe will provide a 
better understanding of the current state of knowl- 
edge in the genus. 

This revision of Brunfelsia is the product of 
Tim Plowman's research, but we accept sole re- 
sponsibility for any errors of interpretation or pre- 
sentation. 



S. Knapp 
J. R. Press 



XI 



Opposite: Brunfelsia hydrangeiformis subsp. hydrangeiformis (originally as Fran- 
ciscea hydrangeiformis), illustration of type. Illustration reproduced from J. E. 
Pohl, Plantarum Brasiliae Icones et Descriptiones (1826). 



A Revision of the South American 
Species of Brunfelsia (Solanaceae) 

Timothy C. Plowmanf 

S. Knapp and J. R. Press, Editors 



I. Materials and Methods 

Beginning in the fall of 1968, I began studying 
the genus Brunfelsia L. in the field, both in South 
America and in the West Indies. During trips 
varying in duration from a few weeks to several 
months, I studied many of the species of Brun- 
felsia in their natural habitats, giving special at- 
tention to population variability and ecological 
conditions. At this time I collected extensive ma- 
terial for morphological, cytological, and chemi- 
cal studies, including numerous herbarium speci- 
mens and living material in the form of seeds and 
cuttings for propagation in the greenhouse. I also 
took this opportunity to investigate the medicinal 
properties and uses of the plants among native 
people who still possess an intimate knowledge of 
plant medicines. I learned about these aspects of 
the plants not only by inquiring among native 
healers but also by taking the medicines myself 
(Plowman, 1977). 

Many of the species I had hoped to see in the 
field could not be found in localities where they 
were formerly collected, due to both the sporadic 
occurrence of the plants and the destruction of 
their habitats by man. At other times, the plants 
were found but without much sought-after flowers 
or fruits. 

To acquire a historical perspective of the genus, 
to properly typify specific names, and to view as 
much morphological variation as possible, I bor- 
rowed herbarium specimens from many institu- 
tions in Europe, South America, and the United 
States. These specimens, numbering about 3,000 
[the number at the time of his thesis in 1974, al- 
though many more subsequently were examined 
by Plowman Eds.}, provided the basis for the 
study. This material was organized according to 
the recommendations outlined by Leenhouts's 
(1968) A Guide to the Practice of Herbarium Tax- 
onomy, which offered many helpful suggestions. 

Some 55 morphological characters were ob- 



served and measured or scored on a character 
flow sheet. For many specimens it was not pos- 
sible to record all the characters, especially those 
of the fruits, which are rarely, if ever, collected, 
and those of the corolla, which is usually poorly 
pressed and often glued fast to the herbarium 
sheet, precluding dissection of the stamen and pis- 
til. The characters and their variation were ana- 
lyzed according to methods outlined and dis- 
cussed in Benson (1962), Davis and Hey wood 
(1963), Lawrence (1965), and Anderson (1967), 
among others. Descriptions of the various taxa 
were prepared using characters observed in her- 
barium specimens and in living plants. In many 
cases the color of the flowers and other evanes- 
cent characters were not known with certainty; as 
a result, the completeness of the description varies 
somewhat from species to species. 

Pollen was acetolyzed according to the proce- 
dure of Erdtman (1960). Unacetolyzed pollen 
grains were also examined with the aid of a scan- 
ning electron microscope. Cytological material 
was collected in the field and in the greenhouse 
and fixed in Carney's solution (absolute alcohol: 
glacial acetic acid, 3:1), then stored in 95% al- 
cohol. Chromosomes were stained with acetocar- 
mine. 

Bulk material for chemical analysis was col- 
lected in the field and dried in the open sun or in 
special drying ovens. This material was sent to 
Mr. Jan-Erik Lindgren at the Department of Tox- 
icology, Karolinska Institute!, Stockholm, Swe- 
den, and to Dr. John Leary, Massachusetts Col- 
lege of Pharmacy, Boston, Massachusetts. Ex- 
tracts of these plants were tested for alkaloids and 
general constituents using precipitation reactions 
and gas and thin-layer chromatography [see Plow- 
man, 1977; Lloyd et al., 1985; Schultes & Raf- 
fauf, 1990, 1991 Eds.]. In addition, simple tests 
for alkaloids were performed by Prof. Robert F. 
Raffauf and myself on fresh plants growing in the 
greenhouse. Extracts were prepared from fresh 



FIELDIANA: BOTANY, N.S., NO. 39, AUGUST 31, 1998, PP. 1-135 



material of five species and tested for alkaloids by 
precipitation reactions. 

Sixteen species were grown in the greenhouses 
of the Biological Laboratories, Cambridge, Mas- 
sachusetts; the Arnold Arboretum, Jamaica Plain; 
and the Botanical Research Station of Northeast- 
ern University in Woburn, Massachusetts. [Other 
specimens were cultivated at the Field Museum 
of Natural History, Chicago, Illinois, in later 
years. Eds.] Material was propagated from both 
seeds and cuttings collected in the wild or from 
cultivated plants. Crosses were attempted between 
as many species as possible. The living plants 
were also a valuable source of those morpholog- 
ical characters that are lost in dried specimens and 
permitted observations to be made on the growth 
and development of various organs, particularly 
the inflorescence. 

A considerable amount of time in this study 
was devoted to searching the botanical, chemical, 
and pharmacological literature for references to 
Brunfelsia. Only those works that are cited in the 
text are included in the bibliography. 

All of the specimens cited in this work were 
examined by me unless they are denoted by "non 
vidi." The use of brackets in the citation of a col- 
lection number indicates that some uncertainty ex- 
ists or, in the case of type materials, that the num- 
ber appearing on the sheet was not given in the 
original citation of the specimen. 

Finally, the use of taxonomic characters de- 
serves some explanation. Of the 24 species of 
Brunfelsia that are recognized in South America, 
four are considered to possess one variety each in 
addition to the typical variety. The category of 
subspecies is here used for geographical or eco- 
logical races that show morphological differenti- 
ation from the typical form of the species. It 
should be noted, however, that the degree of dif- 
ferentiation and isolation varies from one species 
to another, so that the subspecific concepts are not 
entirely equivalent. Furthermore, variants exhib- 
iting considerable differentiation exist within sev- 
eral subspecies (e.g., B. grandiflora subsp. schul- 
tesii Plowman, B. brasiliensis (Spreng.) L.B. Sm. 
& Downs subsp. brasiliensis). Subsequent work- 
ers in the genus may choose formally to recognize 
these as variants at infraspecific rank. 



II. Taxonomic History of the Genus 

The genus Brunfelsia was first described by 
Charles Plumier in 1703 in his Catalogus Plan- 
tarum Americanarum, commemorating the Ger- 
man herbalist Otto Brunfels (1464-1534). Brun- 
fels was a Carthusian monk and physician who, 
in 1530, published an herbal, Herbarum Vivae Ei- 
cones, that, as the name indicates, included illus- 
trations of living plants. This early work is out- 
standing for the high quality, accuracy, and nat- 
uralistic aspect of the woodcuts, which for the first 
time departed from the often copied and stylized 
illustrations of earlier herbals (Arber, 1912). Un- 
fortunately, Plumier misspelled the genus name as 
"Brunsfelsia" an error that has tenaciously per- 
sisted in the literature [see D'Arcy, 1989 Eds.]. 

Simultaneously with his short description and 
diagnosis, Plumier published a rather good illus- 
tration of the flower and fruit of the plant, which 
is now known as B. americana L. The illustration 
was taken from a larger, more complete drawing 
that, along with more detailed descriptions in 
manuscript form, is kept at the Museum National 
d'Histoire Naturelle in Paris, France. The com- 
plete plate was published with a more complete 
description in 1756 in Burman's edition of Plu- 
mier's Catalogus Plantarum Americanarum (Fig. 

1). 

In the second edition of Genera Plantarum 
(1742), Linnaeus included Brunfelsia, based on 
Plumier's description. He classified Brunfelsia 
among the Pentandria Monogynia, indicating that 
the plant had five stamens (an aberrant condition 
that occasionally occurs in B. americana). It 
would thus seem that Linnaeus had access to 
specimens since Plumier failed to mention this 
character and it is not indicated in his drawing. 
Although there is no known specimen of B. amer- 
icana in extant Linnaean herbaria, he did indicate 
with a symbol in the fifth edition of Genera Plan- 
tarum (1754) that he knew the genus from her- 
barium material. 

In a letter to David Van Royen in 1767, Lin- 
naeus mentioned that he did, in fact, see a speci- 
men of the Brunfelsia of Plumier: " Brunsfelsiam 
Plumieri vidi exsiccatam, at ilia forte a Tua specie 
diversa erat, ideoque nihil certi de ea potui pron- 



FIG. 1. Brunfelsia americana. From Burmann's edition of Plumier's Catalogus Plantae Americanarum (1703). 



FIELDIANA: BOTANY 



TAB . LXV. 




PLOWMAN: REVISION OF BRUNFELSIA 



unciare, nee Adansonii compilationes mihi suffec- 
ere" (Bonnet, 1895). Van Royen had sent Linnae- 
us a specimen (no. 257.1, LINN) that Linnaeus 
recognized as being different from Plumier's spe- 
cies. This plant is now referable to B. undulata 
Swartz. 

The valid publication of the genus dates from 
1753, when Linnaeus published B. americana in 
Species Plantarum, in which he followed the orig- 
inal misspelling of the genus. This was apparently 
an orthographical oversight on his part with which 
he was inconsistent in later works. For example, 
in the following year (1754), he used the correct 
spelling in Genera Plantarum but misspelled it in 
subsequent editions of Species Plantarum. In the 
International Code of Botanical Nomenclature 
(Greuter et al., 1994, p. 297), Brunfelsia is listed 
in Appendix IIIA as a conserved spelling [see also 
D'Arcy, 1989 Eds.} . 

The next important date in the history of the 
genus was 1826, when Pohl published seven new 
species from Brazil as the genus Franciscea, 
which he placed in the order Scrophularinae, close 
to BrowalHa. Each of Pohl's seven new species 
was accompanied by an excellent illustration 
(frontispiece, Figs. 22, 28, 31, 39, this work). Ad- 
ditional species were soon added to Franciscea 
by Chamisso and Schlechtendal (1827) and Hook- 
er (1828). 

However, as early as 1829, David Don pub- 
lished his observations on this group, in which he 
maintained that Franciscea did not differ signifi- 
cantly from Brunfelsia. He united the two genera 
under the earlier name Brunfelsia but published 
only one new combination, B. uniflora (Pohl) D. 
Don. Don also expressed the opinion, precocious 
for the time, that Brunfelsia belonged in the So- 
laneae (Solanaceae) rather than the Scrophulari- 
nae, an idea suggested even earlier by Antoine de 
Jussieu (1791). 

Most authors of the time ignored Don's conclu- 
sions and continued to recognize Brunfelsia and 
Franciscea as distinct genera, primarily on the ba- 
sis of the nature of the capsule and the length of 
the corolla tube (Bentham, 1835; G. Don, 1837; 
Endlicher, 1839; Meisner, 1840). 

In 1846, however, in his monograph on the 
Scrophulariaceae for de Candolle's Prodromus, 
Bentham united Franciscea with Brunfelsia, ob- 
serving that although the francisceas were distin- 
guished by some authors, especially by the dry 
capsule, he himself had looked in vain for defin- 
itive structures; B. latifolia (Pohl) Benth. and B. 
guianensis Benth. had small flowers like the rest 



of the francisceas but capsules no less fleshy than 
those of B. americana, and in several species the 
fruit was completely unknown. 

Bentham's treatment of Brunfelsia included 19 
species, eight of which were formerly considered 
Franciscea. He divided the genus into three 
groups without rank: Multiflorae and Uniflorae 
(Franciscea) and Longiflorae (Brunfelsia), placed 
among the Salpiglossideae. This work bears the 
distinction of being the only revision of Brunfelsia 
that has attempted to encompass the entire genus. 
Bentham considered the Salpiglossideae to be a 
"suborder" (subfamily) of the Scrophulariaceae 
and noted their close affinity with the capsular 
Solanaceae, with which, he added, they might be 
better associated. 

Shortly after Bentham's revision appeared, 
John Miers began publishing a series of papers on 
the Solanaceae consisting of taxonomic treat- 
ments, descriptions of new genera and species, 
and a major reorganization of the tribes into two 
families, the Solanaceae and the Atropaceae. 
These replaced the older concept of Solanaceae 
and included the Salpiglossideae of Bentham. The 
text of Miers 's work was originally published in 
The Annals and Magazine of Natural History be- 
tween 1849 and 1857 and later appeared bound 
together in two volumes accompanied by 87 
plates under the name Illustrations of South Amer- 
ican Plants (Miers, 1855, 1857). This extensive 
work remains a major source of information for 
solanaceous plants. 

Miers (1849) erected the new family Atropa- 
ceae as a distinct group with imbricate or plicate 
corolla estivation that was intermediate between 
the Solanaceae (with valvate estivation) and the 
Scrophulariaceae (with imbricate estivation). The 
group included ten tribes, the last of which he 
named the Brunfelsiae. Miers firmly believed that 
Brunfelsia and Franciscea should be maintained 
as distinct genera, citing some ten differences that 
he considered diagnostic. Among these were the 
habit of the plants, the relative length and shape 
of the corolla, the color of the corolla, the shape 
of the anthers, the size and structure of the fruit, 
and the number of seeds per fruit. 

Several points of difference indicated by Miers 
between Brunfelsia and Franciscea no longer 
stand when one considers the wide range of vari- 
ation and many additional species that are now 
recognized. For example, he stated (1849, 1850) 
that brunfelsias are trees attaining 20 ft in height 
whereas francisceas are small shrubs. In fact, both 
low shrubs and small trees occur in about equal 



FIELDIANA: BOTANY 



numbers in both groups. Miers considered the 
fruit of Brunfelsia a fleshy drupe containing an 
indehiscent putamen and that of Franciscea a de- 
hiscent capsule that splits along four vertical su- 
tures. It is now clear that within both Brunfelsia 
and Franciscea, dehiscent and indehiscent, thin- 
and thick-walled, and fleshy and dry fruits occur. 
On the basis of limited material, Miers failed to 
recognize the overall similarities between these 
groups. However, several differences that he 
pointed out remain valid and serve to distinguish 
Brunfelsia and Franciscea as distinct sections of 
the genus. 

Perhaps the most disturbing fact in Miers 's 
study of Brunfelsia is his omission of B. guianen- 
sis, a species that Bentham (1846) described and 
commented on in the Prodromus. In Miers's oth- 
erwise complete list of new combinations, in 
which he transferred most of Bentham's new 
brunfelsias to Franciscea, B. guianensis is con- 
spicuously missing. Since B. guianensis possesses 
several features that are intermediate between 
Brunfelsia and Franciscea, it seems that Miers 
perhaps simply ignored it because it blurred the 
sharp distinction he wanted to demonstrate. 

The view of Bentham, to unite Franciscea with 
Brunfelsia, eventually prevailed over Miers's 
ideas, perhaps because of the wide distribution 
and general acceptance of de Candolle's Prodro- 
mus. Although some of Miers's conclusions have 
not gained acceptance in light of more recent 
knowledge, his extensive observations are none- 
theless valuable to any taxonomist interested in 
the Solanaceae. For example, Bentham and Hook- 
er (1873) drew heavily from Miers's data in de- 
scribing the Solanaceae for Genera Plantarum. 

There has been only one other major study of 
Brunfelsia since the time of Miers and Bentham. 
This is found in J.A. Schmidt's treatment of the 
Scrophulariaceae in Martius's Flora Brasiliensis in 
1864. Although only Brazilian species were in- 
cluded, then numbering about ten, Schmidt made 
many sound taxonomic judgments and correctly re- 
duced many names to synonyms. Until the present 
time, this work remained the standard reference for 
Brunfelsia in Brazil, which possesses the largest 
number of species of any country in South Amer- 
ica. However, Schmidt's treatment is now out of 
date, as may be expected, and not without error. 

In an attempt to clarify certain nomenclatural 
problems regarding his new species Brunfelsia 
mire Monach., Monachino (1953) offered a brief 
discussion of the names of the South American 
species. This was a work of limited value and 



provided little help in solving the taxonomic dif- 
ficulties in the genus, being based primarily on 
the erroneous conclusions of others. 

Besides the original descriptions of the plants, 
which for certain species are the only information 
available, only floristic works have provided ac- 
counts of the species in particular areas. Foremost 
among these, which are by no means of uniform 
quality, are the following works: Flora of the Brit- 
ish West Indian Islands (Grisebach, 1861); Botany 
of Porto Rico and the Virgin Islands (Britton & 
Wilson, 1925); Flora de Cuba (Liogier, 1957); 
Flora of Peru (Macbride, 1962); Flora Ilustrada 
Catarinense, Solandceas (Smith & Downs, 1966); 
Flowering Plants of Jamaica (Adams, 1972); and 
Flora of Panama (D'Arcy, 1974). 

Table 1 summarizes the history of classification 
of Brunfelsia, including generic, tribal, and family 
placement. It should be noted under the heading 
"Family" that the groupings of the earlier bota- 
nists were not equivalent to our modern families 
of plants. These are included for historical pur- 
poses only. 



III. Generic Relationships 

Brunfelsia is a distinct genus that is not readily 
confused with other genera. It belongs to the tribe 
Salpiglossideae of the Solanaceae, as circum- 
scribed by Bentham (1835, 1846, under Scrophu- 
lariaceae), Bentham and Hooker (1873), Wettstein 
(1895), and Baehni (1946). This group is consid- 
ered the most advanced tribe of the 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. 
Eds. 



PLOWMAN: REVISION OF BRUNFELSIA 



TABLE 1 . Summary of historical classification of Brunfelsia. 



Author 


"Family" 


Tribe 


Genus 


Linnaeus (1753) 


Pentandria 




Brunfelsia 




Monogynia 






Swartz (1788) 


Didynamia 




Brunfelsia 




Angiospermae 






Jussieu (1791) 


Personatae 




Brunfelsia 


Pohl (1826) 


Scrophularineae 




Franciscea (excluding Brunfelsia) 


D. Don (1829) 


Solaneae 




Brunfelsia (including Franciscea) 


Bentham (1835) 


Scrophularineae 


Salpiglossideae 


Brunfelsia 








Franciscea 


G. Don (1837) 


Solanaceae 


Francisceae 


Brunfelsia 








Franciscea 


Endlicher (1839) 


Scrophularineae 


Salpiglossideae 


Brunfelsia 








Franciscea 


Bentham (1846) 


Scrophulariaceae 


Salpiglossideae 


Brunfelsia (including Franciscea) 


Miers (1849) 


Atropaceae 


Brunfelsiae 


Brunfelsia 








Franciscea 


Bentham and Hooker (1873) 


Solanaceae 


Salpiglossideae 


Brunfelsia (including Franciscea) 


Baillon (1888) 


Scrophulariaceae 


Salpiglossideae 


Brunfelsia (including Franciscea) 


Wettstein (1895) 


Solanaceae 


Salpiglossideae 


Brunfelsia (including Franciscea) 


Baehni (1946) 


Solanaceae 


Salpiglossideae 


Brunfelsia (including Franciscea) 


Hutchinson (1969) 


Salpiglossidaceae 




Brunfelsia (including Franciscea) 


Plowman (1973) 


Solanaceae 


Salpiglossideae 


Brunfelsia (including Franciscea) 



sibly three heteromorphic species, and Streptoso- 
len Miers, a monotypic Andean shrub genus. 
[Plowmania Hunz., a monotypic genus from mon- 
tane Guatemala, first described as a species of 
Brunfelsia, is also considered to be closely related 
to Brunfelsia (see Hunziker & Subils, 1986). 
Eds.] Browallia is distinguished from Brunfelsia 
by its herbaceous habit; by the filaments, which 
are greatly dilated and ciliate at the apex; by the 
broad, lamellate stigma, which has recurved mar- 
gins, and by the dry capsule, which is dehiscent 
by four sutures. Streptosolen differs from Brun- 
felsia in having the corolla tube twisted at the 
base, the limb bright red-orange, and the filaments 
thickened at the apex. This shrub grows at high 
altitudes in the Andes from Colombia to Peru and 
is pollinated by hummingbirds. 

The other genera of the Salpiglossideae are pri- 
marily annuals found in southern South America: 
Salpiglossis Ruiz and Pavon, Schizanthus Ruiz 
and Pavon, Leptoglossis Benth., Reyesia Clos, Pe- 
tunia Juss., and Schwenkia L. (which also occurs 
in Africa). In addition, three genera of Australian 
shrubs are also placed in the tribe: Duboisia R. 
Br, with indehiscent, baccate fruits, and Antho- 
cercis Labill. and Anthotroche Endl., with dehis- 
cent, capsular fruits. 2 



2 Since the time of the dissertation, these genera have 
been segregated as the endemic tribe Anthocercideae 
(Haegi, 1981). and additional genera have been de- 
scribed. Eds. 



Hutchinson (1969) elevated the Salpiglossideae 
to the rank of family (Salpiglossidaceae) with the 
explanation that the genera of this group consis- 
tently arrived next to Scrophulariaceae in his key 
to the families of plants of the world. This deci- 
sion does not seem justified in view of the exten- 
sive evidence from morphology, anatomy, and 
chemistry, which links the Salpiglossideae with 
the Solanaceae. 

The origin and relationships of the Salpiglos- 
sideae remain obscure. Baehni (1946) derived the 
group from two sources, the Solaneae via the sub- 
tribe Goetzeineae and the Atropeae via Para- 
bouchetia Baillon. In the past, relationships at the 
generic and tribal levels in the Solanaceae were 
studied using relatively few characters, such as 
the curvature of the embryo, corolla estivation, 
number of anthers, or type of fruit. A definitive 
and in-depth evaluation of these higher relation- 
ships is needed in which many characters are con- 
sidered simultaneously, including those from mor- 
phology, chemistry, anatomy, cytology, and other 
areas [see footnote 1 Eds.} 



IV. General Morphology 
Habit 

All known species of Brunfelsia are woody 
shrubs, a few of which will become small trees. 



FIELDIANA: BOTANY 



No herbaceous members are known. In section 
Brunfelsia, certain species may reach 10-12 m in 
height (B. jamaicensis (Benth.) Griseb., B. amer- 
icana, B. lactea Krug & Urb.). These are some- 
what atypical, however, and most are only 2-3 m. 
Species of sections Franciscea and Guianenses 
are mostly medium-sized shrubs 1-3 m tall that 
form a component of the understory of the forest. 
Brunfelsia grandiflora D. Don of the western Am- 
azon may become a small tree 4-5 m tall or, in 
the Brazilian state of Acre, may become lianoid 
in habit, attaining up to 10 m. Brunfelsia latifolia 
is the most diminutive species, often flowering 
when only 30 cm tall and rarely reaching 1 m in 
height. 



Root 

The roots of Brunfelsia are diffuse in the seed- 
lings and young plants. However, mature plants 
often develop a deep, woody taproot. This is es- 
pecially true in species of section Brunfelsia, 
which often grow in rocky areas. In section Fran- 
ciscea, the central root may be more or less 
branched (Fig. 2). 



Branching 

Species of Brunfelsia may have one central 
trunk (B. densifolia Krug & Urb., B. mire, B. chir- 
icaspi Plowman) or they may be branched to a 
greater or lesser extent from near the base, at 
times producing two or more main trunks (B. aus- 
tralis Benth., B. grandiflora, B. brasiliensis). In 
some cultivated species, root suckers often appear 
around the main trunk and form extensive clonal 
plants (B. nitida Benth., B. undulata Sw., B. aus- 
tralis). 

In section Brunfelsia, the branches are often 
few and subvirgate, with few leaves at the end of 
a long, spindly branch (B. undulata, B. splendida 
Urb.). In sections Franciscea and Guianenses, the 
branches are more often diffuse or spreading and 
tend to arch over with age. 

The branching pattern in Brunfelsia is sympo- 
dial. When the terminal bud ceases to grow, the 
shoot is perpetuated from one or more subtermi- 
nal axillary buds or, rarely from the bracts of the 
inflorescence. In the new shoot, several prophylls 
are produced, followed by a gradual transition to 
mature leaves. 



Leaf 

The leaves of Brunfelsia are alternately ar- 
ranged with 5/13 phyllotaxy. They may be scat- 
tered along the branchlets or crowded toward the 
apex, rarely subverticillate through periodic short- 
ening of the internodes (B. mire, B. hydrangeifor- 
mis (Pohl) Benth., B. rupestris Plowman). The 
petiole is very short in relation to the leaf blade, 
varying from 1 to 10 mm in length. It is semiter- 
ete in section and canaliculate, and frequently be- 
comes transversely cracked or corrugate with age 
or drying. 

The leaves are simple with an entire margin. 
There is much variation in their size and shape, 
which provide important characters for distin- 
guishing the species. In form, the blade may be 
elliptic to lanceolate; obovate to oblanceolate; or, 
rarely, spathulate, ovate, linear, or linear-lanceo- 
late. Certain species are noted for great variability 
of leaf shape, sometimes in the same individual 
(B. uniflora (Pohl) D. Don, B. americana, B. lac- 
ted). In other species (B. portoricensis Krug & 
Urb., B. densifolia), the leaves of seedlings and 
young plants differ radically from the foliage of 
adult plants. At the apex, the leaves may be blunt, 
acute, or acuminate. It is of ecological interest that 
the leaves of species of wet, tropical forests are 
often acuminate in the form of a modified "drip 
tip" (B. grandiflora, B. macrocarpa Plowman, B. 
chocoensis Plowman, B. amazonica Morton). 

In size, leaves may be 1-30 cm long and 0.1- 
15 cm wide. Leaf size may vary considerably 
within a species (e.g., B. grandiflora, 6-23 cm 
long) and is strongly influenced by local environ- 
mental conditions such as available light and 
moisture. 

The leaf texture is mostly firm-membranaceous 
to subcoriaceous, often soft and rubbery to the 
touch. A few species (B. splendida Urb., B. 
dwyeri D'Arcy, B. obovata var. coriacea J. A. 
Schmidt, B. rupestris) are noted for their shiny, 
thick-coriaceous leaves. Coriaceous leaves also 
occur in certain populations of B. uniflora and B. 
brasiliensis that grow in unusually dry or exposed 
situations. 

In color, the leaves are light to dark green and 
paler green beneath. Dark purple pigmentation 
due to anthocyanins is present in the young flush 
growth of several species (B. pilosa Plowman, B. 
plicata Urb., B. undulata). The leaves of B. splen- 
dida are particularly striking, being very dark 
green, almost black, and very shiny. 

The nervation in Brunfelsia consists of the mid- 



PLOWMAN: REVISION OF BRUNFELSIA 




FIELDIANA: BOTANY 



rib, which is completely submerged on the adaxial 
surface and prominent beneath, and three to 12 
pairs of pinnate, lateral nerves. Each nerve 
spreads out from the costa and arcuately anasto- 
moses with the next near the margin. The veinlets 
are finely reticulate and may be obscure or some- 
what prominulous. The number of lateral nerves 
may be used to separate certain species (e.g., B. 
martiana Plowman from B. guianensis), but the 
angle of divergence and degree of arching are of- 
ten variable. Brunfelsia hydrangeiformis is unique 
in having the midrib, nerves, and major veinlets 
sunken into a shallow depression or sulcus on the 
adaxial surface, creating a very uneven surface 
texture. 



Indument 

Most Brunfelsia species are essentially glabrous 
throughout, although the young leaves and flower 
buds may be sparsely puberulent in their earlier 
stages, becoming glabrous with age. In the pu- 
bescent species, trichomes are usually encoun- 
tered in the twigs, leaves, pedicel, calyx, and co- 
rolla tube. In section Franciscea, the indument is 
best developed in B. boliviano Plowman, B. bras- 
iliensis, B. cuneifolia J.A. Schmidt, B. hydrangei- 
formis, and B. pilosa, and to a much lesser extent 
in certain populations of B. uniflora, B. pauciflora 
(Cham. & Schltdl.) Benth., and B. obovata Benth. 
In section Brunfelsia, the following species may 
be pubescent: B. americana, B. sinuata A. Rich., 
B. membranacea Urb., and B. jamaicensis. Spe- 
cies of section Guianenses are entirely glabrous 
but occasionally have sparse glandular hairs on 
the petioles and pedicels (B. burchellii Plowman, 
B. clandestina Plowman) and ciliate or villous 
bract margins (B. chocoensis, B. martiana). 

Uniseriate trichomes are found in both section 
Brunfelsia and section Franciscea. They may be 
either glandular or nonglandular, containing from 
two to ten cells. Glandular trichomes often have 
a single terminal, globose cell or may have one 
to several biseriate, terminal cells (B. uniflora). 
Several types of trichomes may occur together in 
the same organ, or only one type may be found. 
In Venezuelan populations of B. uniflora, the ca- 
lyx and pedicel may bear short glandular tri- 
chomes, while the twigs and leaves may be pilose 



to villous with nonglandular hairs and a sharp de- 
marcation at the attachment of the pedicel. 

In section Brunfelsia (B. americana, B. sinu- 
ata), simple, branched trichomes also occur on the 
leaves. These have not been observed in section 
Franciscea. 

The presence, location, and degree of pubes- 
cence appear to be highly variable in some species 
(B. uniflora, B. pauciflora, B. americana). At- 
tempts have been made in the past to distinguish 
varieties solely (or essentially so) on the basis of 
the presence or absence of trichomes (B. uniflora 
var. pubescens (Benth.) R. E. D. Baker & N. W. 
Simmonds, B. americana var. pubescens Griseb., 
B. pauciflora var. calycina (Benth.) J. A. Schmidt, 
B. brasiliensis var. acuminata (Pohl) L. B. Sm. & 
Downs). Numerous intergradations in degree of 
pubescence are found in these species, and lack- 
ing correlation with other, more stable characters, 
it would be unwise to recognize these as "varie- 
ties." Indeed, variation can be observed within 
duplicates of a single collection. [This is also 
commonly the case in other Solanaceae (see 
Knapp, 1989). Eds.] The greater development of 
pubescence is sometimes associated with plants 
growing in drier or more exposed habitats, sug- 
gesting that they may be environmentally induced 
forms or ecotypes. In any case, the unstable nature 
of these variants precludes their receiving formal 
taxonomic status. In other species, however, the 
type and location of trichomes, correlated with ad- 
ditional characters, can be useful in delimiting 
subspecies (B. brasiliensis, B. hydrangeiformis). 

In relatively few species is the indument of di- 
agnostic value at the specific level. Brunfelsia pi- 
losa is characterized by long, pilose trichomes on 
the twigs, leaves, and calyces. Brunfelsia bras- 
iliensis has a distinctive yellow-brown pubescence 
that is generally distributed in the plant. Brunfel- 
sia cuneifolia usually has a villous calyx, a lower 
leaf surface, and twigs, although the type collec- 
tion is nearly glabrous. Brunfelsia boliviana has 
villous-ciliate leaf margins and villous twigs. 



Inflorescence 

The inflorescence may be strictly terminal, ax- 
illary, or a combination of terminal and axillary 
in the same plant. 



FIG. 2. Root of Brunfelsia grandiflora subsp. schultesii. 



PLOWMAN. REVISION OF BRUNFELSIA 



The inflorescence in Brunfelsla is basically a 
cincinnus or some reduction or variation thereof 
(Danert, 1958). In Diccionario de Botdnica, Font 
Quer (1965) provided a succinct definition of the 
cincinnus as a uniparous cymose inflorescence in 
which the medial plane of each branchlet is placed 
transversely with respect to the medial plane of 
the subtending bract of the main axis (pseudaxis) 
and with the lateral branchlets alternately dis- 
posed to the right and left. It is a scorpioid cyme 
in which the various pseudaxes (branchlets) do 
not all fall in the same plane. 

The cincinnal cyme is a primitive type of inflo- 
rescence in the Solanaceae in that the lateral 
branchlets are essentially undifferentiated from the 
primary inflorescence axis. In view of the other- 
wise advanced position of Brunfelsia in the family, 
this is a striking feature [but see Olmstead & Palm- 
er, 1992 Eds.]. 

In Brunfelsia species with well-developed in- 
florescences, the cincinnus is elaborated as fol- 
lows: a single flower terminates the main or lat- 
eral axis and is subtended by one to three (rarely 
more) small, caducous bracts, each of which 
bears a bud in its axil. From one or more of these 
bracts arise lateral branchlets, which are usually 
highly reduced, each of which bears a terminal 
flower and usually two tiny bracts, one on each 
side. Each successive branchlet is formed at right 
angles to its subtending bract. In secondary and 
higher orders of branching, only one of the two 
bracts forms a branchlet, alternating to the right 
and left. This gives the older branches of the in- 
florescence a zigzag pattern, produced by the re- 
maining pedicel scars (e.g., B. latifolia, B. gran- 
diflord). 

The most common form of inflorescence in the 
genus is few-flowered, compact, and sessile, with 
the branchlets of the cincinnus very reduced. In 
some species, the individual branchlets are often 
difficult to distinguish, so condensed are the 
flower clusters. Two species (B. grandiflora, B. 
brasiliensis) are known to develop open, lax in- 
florescences in which the branches may be elon- 
gated to greater or lesser extent. In other species 
(B. mire, B. hydrangeiformis), many flowers are 
produced but without elongation of the inflores- 
cence branchlets, resulting in a very dense, cap- 
ituliform cyme. In still others, the inflorescence 
consists of a single flower (B. americana, B. 
dwyeri, B. macrocarpa, B. uniflora) or one to 
three flowers (B. pilosa, B. australis, B. cunei- 
folia, B. rupestris). 

Great plasticity is found in the expression of 



the inflorescence in a single species and is due 
to both environmental and genetic factors. Spe- 
cies that frequently produce many flowers (B. 
grandiflora, B. hydrangeiformis) in a well-de- 
veloped inflorescence may have only a few flow- 
ers per cluster. Other species that usually are sin- 
gle-flowered (B. pilosa, B. amazonicd) will oc- 
casionally develop two or three flowers. Under 
normal forest conditions, B. chiricaspi bears up 
to seven flowers in the inflorescence; however, 
one collection (Kennedy 1386), growing in full 
sun in a cut-over place, had more than 20 flow- 
ers. Similar light-related responses have been ob- 
served in B. grandiflora (Plowman 2535). This 
variability, evident to some extent in most of the 
species, limits the usefulness of the number of 
flowers per inflorescence as a taxonomic char- 
acter, particularly in the construction of keys. 



Flower 

The flowers of Brunfelsia species are com- 
posed of the following parts: a pedicel; a five- 
lobed calyx; a long-tubular corolla with a five- 
lobed limb; two pairs of stamens inserted in the 
upper part of the corolla tube; and a pistil, the 
style of which nearly equals the tube. There are 
major differences among the flowers of the three 
sections of the genus; however, within a section, 
the flowers are remarkably similar with only a 
few minor exceptions. Differences that do occur 
and that are important in distinguishing the spe- 
cies are found in the relative sizes of the flowers 
and their parts. 

The flowers are slightly zygomorphic with bi- 
lateral symmetry. This is observable in the calyx, 
which occasionally has unequal lobes; in the cur- 
vature of the corolla tube; in the unequal limb; 
and in the didynamous stamens. The stamens and 
style are curved toward the "back" or posterior 
side of the flower, in line with the uppermost co- 
rolla lobe, and the anthers and stigma are brought 
very close to that side of the tube. This construc- 
tion is probably advantageous in pollination be- 
cause the proboscis of the pollinating insect (Lep- 
idoptera) is most likely to descend along this part 
of the tube (see Fig. 3). 

The flowers of section Brunfelsia are very large 
with a long tube and a usually showy white limb 
that fades to yellow (Figs. 4, 5). They are ex- 
tremely fragrant in the evening and apparently are 
pollinated by night-flying moths (Sphingidae). In 
section Franciscea the flowers are smaller and 



10 



FIELDIANA: BOTANY 











GUIANENSES BRUNFELSIA FRANCISCEA 

FIG. 3. Floral structure of the sections of Brunfelsia. 



have a shorter tube with a purple limb and a white 
eye at the center. They are usually not fragrant 
and are pollinated by day-flying butterflies from 
several different families. Species of section Gui- 
anenses have small whitish or greenish flowers 
with a short tube and a relatively narrow limb. 
They are said to be fragrant, but the pollinators 
are unknown. 



Calyx 

The calyx is actinomorphic or slightly zygomor- 
phic, in form tubular to campanulate and often 
somewhat inflated. It varies in length from 3 to 35 
mm. In section Brunfelsia the calyx is very short 
compared to the corolla tube; in sections Francis- 
cea and Guianenses it is usually about half as long, 



PLOWMAN: REVISION OF BRUNFELSIA 



11 




12 



FIELDIANA: BOTANY 



although in some species (B. pauciflora, B. hy- 
drangeiformis) it may equal the corolla tube. 

The calyx teeth or lobes are triangular, ovate or 
lanceolate, and valvate in bud. In section Brun- 
felsia they are often blunt with a few ciliate tri- 
chomes. In sections Franciscea and Guianenses 
the teeth are acute or acuminate, and the ciliate 
margins are much reduced or obsolete. 

The calyx is glabrous or may be furnished with 
glandular (B. boliviana, B. pauciflora, B. uniflord) 
or nonglandular (B. pilosa, B. brasiliensis) tri- 
chomes. In color it is usually some light shade of 
green to yellowish (B. undulata, B. grandiflord) 
or purplish (B. pilosa, B. jamaicensis). 

The size and shape of the calyx, as well as its 
pubescence, are taxonomically important in spe- 
cific delimitation. In most species the calyx is te- 
rete, but in a few (B. obovata, B. boliviana, B. 
cuneifolia, B. nitidd) it is five-angled at the sinus- 
es and concave at the lobes. 

The nature of the calyx in fruit also provides 
useful characters. The calyx is usually persistent 
in fruit but scarcely accrescent. It may become 
thicker and firmer in texture and in section Fran- 
ciscea frequently bears lenticular outgrowths near 
the base. It may split along one or more sides to 
accommodate the developing fruit, persisting only 
at the base, or it may increase in girth to partially 
or completely enclose the fruit at maturity (B. hy- 
drangeiformis, B. pauciflora, B. brasiliensis). 
Brunfelsia macrocarpa is unique in the genus in 
having a large, truly accrescent calyx that be- 
comes thick and leathery, completely enclosing 
the fruit. 



Corolla 

The corolla differs significantly in the three sec- 
tions of Brunfelsia, as mentioned above (Fig. 3). 
In section Brunfelsia the corolla tube is very long, 
from 6 to 24 times as long as the calyx. It be- 
comes gradually dilated near the apex but never 
curved or constricted at the mouth (Fig. 5). The 
limb usually is held at an angle to the tube. The 
five lobes may be narrow or broad, plane or un- 
dulate to crenate at the margin. In color the co- 
rolla tube is greenish or white; the limb is usually 
pure white when it first opens, turning yellow with 
age. Brunfelsia americana and B. lactea may be 



streaked with purple. Anomalous species are B. 
cestroides A. Rich, and B. purpurea Griseb. of 
Cuba, the flowers of which are said to be purplish 
red. 

Section Franciscea has a much shorter tube 
that is only one to five times as long as the calyx. 
It is briefly inflated near the apex to include the 
anthers and stigma, then slightly constricted to 
form a raised, fleshy ring. The tube may be white 
or violet, and the limb is deep to pale violet, fad- 
ing to pale lavender or pure white with age. The 
ring or eye at the center may be white or yellow. 
The limb in section Franciscea may be large and 
varies from 1 .5 to 8 cm in diameter. The lobes are 
broadly rounded, overlapping at the lateral mar- 
gins. 

The corolla tube in section Guianenses is short, 
only two to three times as long as the calyx, and 
gradually dilated or inflated toward the apex. 
There is no marked constriction as in section 
Franciscea. The flowers in section Guianenses are 
white or greenish white and apparently do not 
change color with age. The corolla limb in section 
Guianenses is relatively small, rarely exceeding 
25 mm in diameter. The lobes are quite narrow 
and convex; the lateral margins appear to be re- 
flexed and are completely folded under in pressed 
specimens. This is also observed in B. densifolia 
of section Brunfelsia (Fig. 4). 

The estivation of the corolla lobes in bud is a 
combination of quincuncial and imbricate. In 
quincuncial estivation, two of the lobes lie wholly 
to the outside, two wholly on the inside, and one 
lobe has one side to the inside and one to the 
outside. In section Brunfelsia the uppermost lobe 
is wholly to the inside, one of the lower lobes is 
wholly to the outside, and the other three are im- 
bricate with one side in and one side out. 

Only the size of the corolla and the shape of 
the lobes are useful as taxonomic characters. 
Within each section the corolla is quite similar. 
Characters such as color and the shape of the cen- 
tral eye in section Franciscea are diagnostic in the 
living state, but these are unfortunately lost in 
preparation of specimens and rendered useless to 
the herbarium taxonomist. 

Androecium 

The four stamens occur in two pairs in the up- 
per part of the corolla tube, alternating with the 



FIG. 4. Brunfelsia densifolia (Knapp s.n.) from Puerto Rico. 



PLOWMAN: REVISION OF BRUNFELSIA 



13 




14 



FIELDIANA: BOTANY 



upper three corolla lobes. A fifth stamen is oc- 
casionally produced in Brunfelsia americana, but 
this is an irregular condition. The glabrous fila- 
ments are fused to the corolla tube for most of 
their length and are free for only 2-5 mm. The 
anterior pair is longer, exceeding the stigma, and 
may be briefly exserted from the tube (B. obovata, 
B. chocoensis); the posterior pair, attached to the 
"back" of the tube, is shorter than the style. Both 
pairs are curved at the apex to project the anthers 
toward the back of the tube. In section Brunfelsia 
the filaments may be curved somewhat in the re- 
gion of the stigma (B. americana, B. lactea), 

The anthers are medifixed to the filaments and 
dehisce by a single longitudinal slit. In section 
Brunfelsia the anthers are oblong-reniform and 
may appear to be slightly bilobate, although the 
cells are confluent. At anthesis, each pair may 
tend to adhere to each other, apparently from the 
sticky nature of the pollen. Brunfelsia densifolia 
is somewhat anomalous in having both pairs of 
anthers reaching the mouth of the tube; the ante- 
rior pair is much reduced in size, although it pro- 
duces a small amount of pollen. They are not 
completely sterile, as implied in Urban's original 
description (Krug & Urban, 1897). Slight inequal- 
ity in anther size is not unknown in other species 
(cf., B. martiand) and hardly merits the formation 
of a new genus as proposed by Urban (1906). In 
sections Franciscea and Guianenses the anthers 
are unilocular and more or less semicircular, ap- 
pearing round or reniform in outline. 



Pollen 

The pollen grains of all three sections of Brun- 
felsia are spheroidal. 3 Those of section Brunfelsia 
(B. americana, B. undulata) and section Guianen- 
ses (B. chocoensis, B. martiand) appear triporate 
or possibly tricolporate. Those of section Fran- 
ciscea are tricolpate or tricolporoidate (B. chiri- 
caspi, B. grandiflora, B. latifolia, B. pilosd). They 
are all medium-sized grains 30-38 to 34-42 jim 
in diameter. 

The surface of the pollen was examined by 
means of a scanning electron microscope using 



3 Pollen grains of Brunfelsia species were acetolyzed 
according to the procedure of Erdtman (1960). 



unacetolyzed grains. These revealed low-relief 
scrobiculate or reticulate exine patterns. Although 
the sculpturing appears to differ slightly from spe- 
cies to species, the preparations were not suffi- 
ciently clear to permit a detailed analysis, nor 
were enough species observed to permit general- 
ization. 



Gynoecium 

The ovary is ovoid or conical and is inserted 
on a short, nectar-producing disc that may be ob- 
scure in dried specimens or appear merely as a 
slight thickening at the base of the ovary. The 
ovary is bilocular with axile placentation, bearing 
numerous anatropous ovules. The thin dissepi- 
ments occasionally do not extend to the apex of 
the ovary, breaking down early in fruit formation 
and leaving the placenta attached only at the base. 
The fruit then may have only one locule. 

The style is filamentous and nearly equals the 
corolla tube in length. It may be somewhat thick- 
ened and curved at the apex, especially in sections 
Franciscea and Guianenses, or suberect, as in 
several species of section Brunfelsia. 

The stigma is usually included between the four 
anthers in the uppermost part of the tube. Only 
rarely does it project to the mouth of the tube (B. 
densifolia). The stigma is short and briefly bifid 
in sections Franciscea and Guianenses and faces 
the posterior wall of the corolla tube. It has the 
appearance of an open forceps at anthesis with the 
stigmatic tissue included between the lobes. After 
pollination, the lobes close together. In section 
Brunfelsia the stigma is more shallowly lobed and 
may appear subcapitate and convex. 



Fruit 

There has been considerable misinterpretation 
of the fruit of Brunfelsia from both a morpholog- 
ical and a phylogenetic point of view. This has 
occurred, I believe, because early botanists had 
very little material to work with and drew their 
conclusions from only a few specimens. The fruit 
varies considerably from group to group, and dif- 
ferent fruit types may be found in otherwise 
closely related species. In some cases, the nature 



FIG. 5. Brunfelsia lactea (Knapp s.n.) from Puerto Rico. 



PLOWMAN: REVISION OF BRUNFELSIA 



15 



of the pericarp and surface of the fruit may be 
important in differentiating species; fruit charac- 
ters should be used whenever possible. Unfortu- 
nately, the fruits of many species are poorly or 
not at all known. 

The fruit of Brunfelsia is essentially a capsule, 
not a berry as suggested by Miers ( 1 850) and oth- 
ers in the past. There is, however, great variation 
in the texture of the pericarp, which may be thin- 
walled and brittle, leathery, fleshy or hard and in- 
durated (bony). In shape it ranges from globose 
to ovoid, and in size, from 1 to 5 cm long. There 
may be two locules or only one by degeneration 
of the dissepiments. The surface may be smooth 
or rough; reticulate or areolate; and green, brown, 
or bright yellow or orange. The pericarp may be 
differentiated into three distinct layers or may be 
more or less fused into one. In all cases the en- 
docarp is a thin, somewhat cartilaginous layer that 
on drying becomes brittle. The outer layers of the 
fruit wall often separate from the endocarp when 
the fruit rots. The thin endocarp may then split in 
half to release the seeds. 

The uncertainty and controversy concerning the 
nature of the fruit in Brunfelsia (see "Ecology and 
Reproductive Biology") reflect the scarcity of 
fruiting material for many species. In reality, there 
is a wide spectrum of variation in the texture of 
the pericarp, which has led different authors to 
classify the fruit variously as a berry, a capsule, 
or a drupe. The fruits of certain species are indeed 
difficult to classify, demonstrating the limitations 
in our terminology. When the full range of forms 
is seen, however, it is clear that all the fruits of 
Brunfelsia are modifications of the capsule. Dif- 
ferent types of capsule are found in each of the 
three sections of the genus, and no type is unique 
to any one section. 

In section Brunfelsia the fruit is usually leath- 
ery or fleshy and brightly colored. The calyx per- 
sists only at the base. In B. americana and B. 
densifolia, the valves are fleshy and yellow and 
orange, respectively. The valves split apart at ma- 
turity in a true dehiscence (Fig. 6). Brunfelsia ja- 
maicensis is green at maturity, with a thick, bony 
pericarp. Brunfelsia maliformis Urb. has a large, 
fleshy to leathery fruit with a distinct reticulate- 
areolate surface. It apparently does not open ex- 
cept by rotting. (B. lactea has orange valves that 
rot, allowing the seeds to be shaken out. Eds.). 
Some of the Cuban species have fruits that are 
thin-walled, indehiscent, and usually brightly col- 
ored (B. nitida, B. clarensis Britton & P. Wilson). 



These may function as a berry in dispersal by an- 
imals. 

In section Franciscea the fruit is usually a rel- 
atively small capsule that is green or brown and 
has a coriaceous pericarp that becomes thin and 
brittle on drying. The persistent calyx encloses the 
fruit, at least in part. These capsules may split 
open at maturity, but only sparingly. It may be 
that many species are completely indehiscent and 
their seeds are released only upon rotting on the 
forest floor. Two anomalous species are B. dwyeri 
(from Panama) and B. macrocarpa (from South 
America), in which the fruits have a leathery and 
a fleshy pericarp, respectively. The calyx of B. 
macrocarpa increases greatly in size and itself be- 
comes fleshy, completely enclosing the bright yel- 
low fruits. 

In section Guianenses we find fleshy, yellow 
fruits in two of the six species, with the short ca- 
lyx present only at the base. The fruits of B. ama- 
zonica and B. burchellii are thin-walled and dry 
at maturity; the fruit of B. martiana is as yet un- 
known. 



Seed 

The seeds are oblong or ovoid and often an- 
gular or compressed on one or more sides. They 
range in size from 2 to 13 mm long and in di- 
ameter from 1 to 7 mm. The surface texture is 
reticular-pitted and reddish brown in color. The 
surface pitting is formed by many tightly com- 
pressed surface outgrowths, resembling globulose 
hairs, that slough off as the seeds reach maturity. 
[See Soueges, 1907; Edmonds, 1983; Lester & 
Durands, 1984. These hairs are lignified thicken- 
ings of the lateral testal cell walls. Eds.] Each 
of these hairs leaves a tiny pit in the surface. The 
embryo is relatively large and straight and is em- 
bedded in endosperm. The cotyledons are flat and 
ovate to elliptic. The size of the seeds may be 
useful as a taxonomic character, but their shape 
and number per capsule appear to be quite vari- 
able. 



V. Cytology 

The chromosomes of very few species of Brun- 
felsia have been investigated. Four counts for the 
genus are known from the literature, but these are 
of limited value at the specific level because no 
voucher herbarium specimens were prepared. 



16 



FIELDIANA: BOTANY 




FIG. 6. Fruit of Brunfelsia americana. 



PLOWMAN: REVISION OF BRUNFELSIA 



17 



TABLE 2. Chromosome counts in Brunfelsia. 



Species 



Chromosome no. 



Author 



B. americana 

B. americana 

B. australis 

B. calycina (= B. pauciflora) 

B. dwyeri 

B. americana 

B. australis 

B. calycina (- B. pauciflora) 

B. dw\eri 



2n = 22 

n = 11 

n = 12 

In = 22 

2n = 22 

2n = 22 

n = 11 

n = 12 

2n = 22 

2n = 22 



Bhaduri (1933) 

Madhavadian (1968) 

Ratera (1944) 

Darlington and Janaki Ammal (1945) 

Davidse (1981) 

Bhaduri (1933) 

Madhavadian (1968) 

Ratera (1944) 

Darlington and Janaki Ammal (1945) 

Davidse (1981) 



These findings are summarized in Table 2. On the 
basis of this evidence, Darlington and Wylie 
(1955) hypothesized that the base number for 
Brunfelsia is x = 11. 

The mitotic chromosomes of Brunfelsia are dif- 
ficult to prepare because the young root tips pro- 
duced in flower pots tend to be hard and brittle, 
making squashes very difficult. Treatment in 1% 
HC1 softens the material somewhat, but despite 
this treatment, no good mitotic figures could be 
found in three species examined (B. australis, B. 
latifolia, B. pauciflora). Sufficient seeds of Brun- 
felsia were not available to test the possibilities of 
preparing squashes from the roots of young seed- 
lings. 

A second method of obtaining chromosome 
preparations is to examine pollen mother cells un- 
dergoing meiosis. Young flower buds were col- 
lected both in the field and from cultivated spec- 
imens in the greenhouse and immediately placed 
in Carney's solution for at least 24 hr and refrig- 
erated when possible. Material was collected 
about midday on sunny days. It was difficult to 
obtain a sufficient number of buds because brun- 
felsias generally have few flowers per inflores- 
cence, and these are usually in various stages of 
maturation; not infrequently only one flower per 
inflorescence per day will be available for cytol- 
ogy. Compounded by the fact that most species 
flower only sporadically and some not at all under 
greenhouse conditions, securing adequate material 
was a task. 

Meiosis occurs in Brunfelsia when the buds are 
very small (about 1.0-2.5 mm long) and well be- 
fore the corolla emerges from the calyx. The 
young anthers were teased out in 2% acetocar- 
mine and gently heated, then squashed with even 
pressure. Relatively few countable meiotic figures 
were obtained out of numerous attempts. The re- 



sults obtained in five species and one subspecies 
are summarized in Table 3. 

All counts were obtained from meiotic figures 
with the exception of B. pauciflora, which appar- 
ently represents a somatic division in a tapetal 
cell. The meiotic figures obtained were fixed in 
late anaphase I or telophase II. The chromosomes 
are small, from 2.0 to 5.5 (jtm (cf., Swanson, 
1957), and exhibit some size variation between 
the different species examined. In the case of B. 
nitida and B. americana, the smaller size is due 
to the fact that only two chromatids are present 
(telophase II). The somatic chromosomes of B. 
pauciflora appear greatly enlarged. This is prob- 
ably the result of endomitosis, which produces a 
high degree of endopolyploidy within the cell. 
Unfortunately, no normal mitotic cells could be 
obtained for this species. 

The chromosomes obtained from these species 
were very compact in the stages observed, sug- 
gesting a high degree of contraction due to chro- 
mosomal coiling. No details of chromosomal 
structure could be ascertained from the meiotic 
figures. It is possible, however, to see the centro- 
mere in the somatic chromosomes of B. pauciflo- 
ra. 

In an extensive investigation, Gottschalk (1954) 
examined the structure of the pachytene chromo- 
somes of many species of Solanaceae, with par- 
ticular reference to the presence and position of 
heterochromatic and euchromatic areas. Because 
of the high degree of contraction, he was unable 
to obtain pachytene chromosomes in what he re- 
ferred to as B. macrophylla (probably B. pauciflo- 
ra). He did, however, observe the presence of 
chromocenters in the interphase nucleus, indicat- 
ing that the chromosomes of this genus are dif- 
ferentiated into euchromatic and heterochromatic 
regions. Similar results were found in species in 



18 



FIELDIANA: BOTANY 







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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 



19 



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- 
sachusetts. 

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- 
cessful. 

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 
seeds. 

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 
among them. 



VII. Ecology and Reproductive 
Biology 

Habitat 

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- 



20 



FIELDIANA: BOTANY 



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 
collections. 

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- 
tions. 

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 
Andes. 

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 
habitats. 

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 



21 



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. 



Life Cycle 

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- 
nisms." 



Pollination Biology 

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 
all self-incompatible. 

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 
while feeding. 

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. 



22 



FIELDIANA: BOTANY 



TABLE 4. Pollinators of Brunfelsia grandiflora. 



Family 


Subfamily 


Genus 


Voucher 


Locality 


Subspecies schultesii 










1. Nymphalidae 


Satyrinae 


Euptychia sp. 1 


Plowman 2091 
Plowman 2533 


Putumayo, Colombia 
Loreto, Peru 


2. Nymphalidae 
3. Nymphalidae 
4. Pieridae 
5. Lycaenidae 


Satyrinae 

9 

Riodininae 


Euptvchia sp. 2 
7 

Eurema sp. 
Euselasia sp. 


Plowman 2533 
Plowman 2091 
Plowman 2091 
Plowman 2533 


Loreto, Peru 
Putumayo, Colombia 
Putumayo, Colombia 
Loreto, Peru 


Subspecies grandiflora 










6. Hesperidae 
7. Hesperidae 


Hesperiinae 
Hesperiinae 


Calpodes group 


Plowman 2912 
Plowman 2912 


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. 



Dispersal Mechanisms 

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 
seeds. 

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, 
1952). 

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. 



Plant-Insect Relationships 

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 



23 




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 
host. 

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. 



24 



FIELDIANA: BOTANY 




PLOWMAN: REVISION OF BRUNFELSIA 



25 



TABLE 5. Brunfelsia species recorded as larval food 
plants for species of Methona (Lepidoptera: Nymphali- 
dae: Ithomiinae). 

M. M. M. M. 

Species confusa megisto singularis themisto 

B. amazonica 

B. australis 

B. dwyeri 

B. grandiflora 

B. pauciflora 

B. mire 

B. martiana 

B. uniflora 



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 
the plant. 



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- 
ciscea. 

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. 



26 



FIELDIANA: BOTANY 



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- 
sociated characters. 

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 



27 



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 
branches. 

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- 
ovata. 

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 



28 



FIELDIANA: BOTANY 



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 
their sections. 

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 
Relationships 8 

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 



29 



TABLE 6. Trends in specialization of Brunfelsia. 



Less specialized 



More specialized 



Plants of humid, tropical forests 

Leaves scattered on stem 

Leaves large, soft-leathery 

Inflorescence terminal and subterminal 

Inflorescence lax, branched 

Flowers few-many, small 

Flowers scentless 

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 
changing color 

Corolla lobes narrow, not overlapping, 
plane 

Fruits dehiscent 

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 

Flowers fragrant 

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 

Fruits indehiscent 

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 
been undertaken. 

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 



30 



FTELDIANA: BOTANY 



JAMAICA HISPANI 

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 



BJQLOGY LIBRARY 
101 BWWLL HALL 



31 



CCD 



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. 
Eds. 



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. 



32 



FIELDIANA: BOTANY 



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 
present-day species. 

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- 
eastern Brazil. 

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 



33 



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 
speciation developed. 

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 Linnaeus 



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 



34 



FIELDIANA: BOTANY 



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 
America. 



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 

Section Brunfelsia 

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 



35 



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 

subsp. 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. 

pauciflora 

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- 
minate 
30. Pedicels 6-12 mm long; calyx in fruit nearly as long as 

capsule. SE Brazil 9. B. bonodora 

30. Pedicels 2-6 mm long; calyx in fruit half as long as 
capsule or less. W South America . . 14b. B. grandi- 
flora subsp. schultesii 

PLOWMAN: REVISION OF BRUNFELSIA 37 



Brunfelsia sect. Guianenses Plowman in Hawkes, 
J. Linn. Soc. (Bot.) 76: 294. 1978. 

TYPE SPECIES Brunfelsia guianensis Benth. in 
DC., Prodr., 10: 200. 1846. 

Shrubs to 2 m tall. Inflorescence terminal and 
axillary. Flowers small, 1-7 per inflorescence. 
Corolla white or greenish white, tube 2-3 times 
as long as calyx, open at mouth, not constricted, 
the lobes narrow, usually with deflexed margins. 
Stigma briefly bifid, in the form of a forceps. Cap- 
sule dry or fleshy at maturity. South America. 

1. Brunfelsia amazonica C.V. Morton, Proc. 
Biol. Soc. Wash. 62: 151. 1949. Type: BRA- 
ZIL Amazonas: Manaus, Estrada do Raiz, sil- 
va secundaria, non inundabilis, manacd, 24 
Mar. 1937, Ducke 430 (holotype, us 
1693434; isotypes, A, F-902265, K, MO- 
1156896, NY, R-75434, s). Figure 11. 

Shrub or small tree 1.5-2.5 m tall. Branches 
spreading, knobby. Bark buff to reddish brown, 
cracked in longitudinal fissures. Brant hk-ts shiny, 
glabrous, light, yellowish brown. Leaves scattered 
on branchlets, blade 6.5-12 cm long, 2.5-5 cm 
wide, elliptic-oblong to lanceolate, abruptly acu- 
minate, acumen often oblique, broadly cuneate to 
narrowed at base, glabrous, dark green above, 
pale green beneath, firmly membranaceous, lateral 
nerves 5-8(10), straight; petiole 2-5(8) mm long, 
transversely corrugate with cracked epidermis, 
glabrous. Inflorescence terminal or axillary, ses- 
sile with 1-2(5-6) flowers. Flowers white or 
greenish white, fragrant. Bracts 1-2 per flower, 
2-6 mm long, linear to lanceolate, glabrous, ca- 
ducous. Pedicel curved in bud, becoming erect, 
1 3-20 mm long, 1 mm in diameter, slender, thick- 
ened toward apex, glabrous. Calyx 8-12 mm 
long, 5-8 mm in diameter, tubular-campanulate, 
slightly inflated, obovate in bud, glabrous, green, 
membranaceous, teeth 2-6 mm long, erect, some- 
what unequal, ovate-lanceolate, apically blunt to 
acuminate; calyx in fruit, enclosing capsule at 
base. Corolla tube 20-25 mm long, 1.5-3.0 mm 
in diameter, twice as long as calyx, cylindric, 
slightly dilated toward mouth, glabrous, minutely 
glandular at mouth; limb 15-25 mm in diameter, 
spreading, lobes 6-11 mm long, subequal or the 
posterior lobe somewhat larger, oblong to obo- 
vate, somewhat convex, blunt or rounded at apex, 
the sides of the lobes reflexed after anthesis, nar- 
rowed slightly toward base. Stamens inserted at 



middle of tube, included; filaments subligulate, 
upper pair 5 mm long, lower pair 2-3 mm long; 
anthers 1-1.5 mm in diameter, orbicular-reniform. 
Ovary 1.5 mm long, 1 mm in diameter, narrowly 
conical; style 16-18 mm long; stigma 1 mm long, 
bifid, upper lobe somewhat larger. Fruit 10 mm 
in diameter, globose, apiculate at apex, green, 
smooth, pericarp thin, about 0.2 mm thick, dry at 
maturity, sparingly dehiscent. Seeds 8-10, 5-6 
mm long, 3 mm in diameter, oblong-reniform, 
flattened on 1 or 2 sides, angular, dark brown, 
reticulate-pitted with a medial hilum. Embryo 3 
mm long, slightly curved; cotyledons about 0.5 
mm long, elliptic. 

DISTRIBUTION Brazil (Amazonas). See Figure 
15. 

SPECIMENS EXAMINED BRAZIL. Amazonas: 
Manaus, Estrada da Raiz, 18 Mar. 1943, Ducke 
430 (paratype, MG, us); perto da Estrada da Raiz, 
22 Mar. 1932, Ducke sp. nov. (GH, RB); Mata do 
Aleixo, 4 Mar. 1945, Froes 20535 (NY, us); Uyp- 
iranga, Rio Negro, perto de Manaus, 21 Feb. 
1923, J.G. Kuhlmann 943 (GH, RB); road Manaus 
(Cacau-Pireira) to Manacapuru, km 25, 3 Jan. 
1967, Prance, Pena & Ramos 3880 (F, INPA, K, 
MG, P, R, s, us); ad oram meridionalem Rio Negro, 
usque ad concursum flum. Solimoes, May 1851, 
Spruce 1495 (EM, BR, CGE, F, G, GH, K, LE, MG, NY, 
p, RB, w); *Rio Negro, Tapuruquara, Santa Izabel, 
Black 48-2905 (IAN). 

This species was first collected by Spruce in the 
middle of the last century and distributed under 
the name B. spruceana to the world's herbaria. 
This name was recorded but not validly published 
by Schmidt, who placed it in synonymy with B. 
maritima Benth. 

Brunfelsia amazonica is a very local species, 
known only from the environs of Manaus at the 
mouth of the Rio Negro in Brazil. Notwithstand- 
ing the intensity of collecting in this region, B. 
amazonica has not been collected often. It is a 
very distinct species of section Guianenses and is 
easily separated from B. guianensis and B. cho- 
coensis by the longer pedicels, the oblong-lanceo- 
late leaves, and dry, thin-walled fruit, and from 
B. martiana by the longer pedicels and smaller 
leaves. 

Brunfelsia amazonica grows in open secondary 
forests on the southern bank of the Rio Negro in 
white sandy soil. It is known to flower from Jan- 
uary to May; fruits have been collected only in 



38 



FIELDIANA: BOTANY 




FIG. 1 1 . Brunfelsia amazonica. 



PLOWMAN: REVISION OF BRUNFELSIA 



39 



May. Like many brunfelsias in Brazil, this species 
is locally called manacd. 

2. Brunfelsia burchellii Plowman, Fieldiana Bot. 
n. s. 8: 9. 1981. Type: BRAZIL. Goias: Porto 
Real (now Porto Nacional), ford of Igarape, 
1828-1830, Burchell 8527 (holotype, P; iso- 
types, K, L). H Figure 12. 

Shrub. Mature branchlets 2-3 mm in diame- 
ter, spreading, with shiny, yellowish to dark 
brownish, longitudinally cracked bark. Leaves 
scattered along stem, short petiolate, blade 70- 
180 mm long, 30-75 mm wide, broadly to nar- 
rowly ovate, apically acuminate with a long, 
pointed, often falcate acumen, sometimes abruptly 
acuminate, minutely ciliolate at margin, basally 
rounded to obtuse, glabrous on both sides, dull or 
somewhat shiny, medium green above, dull, paler 
green beneath, firmly membranaceous to charta- 
ceous, the lateral nerves 5-9, strongly arcuate, 
anastomosing near margin, nerves prominent on 
lower surface; petiole 1-6 mm long, with a few 
scattered glandular hairs, glabrescent. Inflores- 
cence terminal on current year's branchlets, very 
briefly pedunculate, with 1-3 flowers. Peduncle 
14 mm long, persistent, sparsely glandular-pubes- 
cent, glabrescent. Bracts 1-3 per flower, 4-20 
mm long, linear to lanceolate, apically acuminate, 
glabrous or villous at nerves and margin, cadu- 
cous. Flower color unknown. Pedicel 4-8 mm 
long, 1 mm in diameter, sparsely glandular-pubes- 
cent, glabrescent. Calyx 1 1-20 mm long, 5-9 mm 
in diameter, tubular or tubular-campanulate, te- 
rete, with scattered glandular hairs, glabrescent, 
striately nerved, teeth 2-6 mm long, unequal, tri- 
angular-ovate, apically acute to acuminate, mi- 
nutely gland-tipped. Corolla tube 24-28 mm 
long, 1.5-2 mm in diameter, \Vi-2 times as long 
as calyx, straight, glabrous or bearing few scat- 
tered glandular hairs, orifice 5 mm in diameter; 
limb 1 8-30 mm in diameter, spreading, lobes sub- 
equal, rounded, overlapping at the lateral margins. 
Stamens inserted in upper part of corolla tube; 
filaments 1 mm wide, straplike, the anterior pair 
3.5-6 mm long, apically slightly incurved or sub- 
erect, the posterior pair 3-4 mm long; anthers 1 
mm in diameter, orbicular-reniform. Ovary 1.5 
mm long, conical-ovoid; style 20-21 mm long, 
broadened and incurved at apex; stigma included 
between pairs of anthers, 1 mm long, briefly bifid, 



11 All Burchell localities are taken from his itinerary, 
published by Smith and Smith (1967). 



upper lobe slightly larger. Fruit and seed un- 
known. 

DISTRIBUTION Basin of the upper Rio Tocan- 
tins (Goias State) and adjacent Maranhao State, 
Brazil. See Figure 15. 

SPECIMENS EXAMINED BRAZIL. Goias: Porto 
Real, Porto Real to Igarape, 1828-1830, Burchell 
8415 (L); about village, Burchell 8494 (K); at en- 
trance to village, Burchell 8653 (K); "about the 
Manga" (cf., Rio dos Mangues, where it meets 
the Tocantins), Burchell 8752 (CH). Maranhao: 
Island of Sao Luiz, Estrada do Barreto, Feb.-Mar. 
1939, Froes 11620 (A, F, NY, s, us). 

Brunfelsia burchellii is known from only a few 
collections by William J. Burchell in the vicinity 
of Porto Real (now called Porto Nacional) on the 
upper Rio Tocantins 150 years ago. Unfortunately, 
these specimens bear no field data, and the plant 
has not been recollected in this region. A modern 
collection by Froes from Sao Luiz Island, more 
than 1,000 km north of Porto Nacional, is tenta- 
tively assigned to B. burchellii but lacks mature 
flowers or fruits and differs somewhat in leaf ve- 
nation. 

Brunfelsia burchellii appears to be most closely 
related to B. guianensis, from which it differs by 
the ovate, acuminate leaves, the longer tubular ca- 
lyx, and the broadly rounded corolla lobes. With 
B. guianensis, B. burchellii is provisionally placed 
in section Guianenses on the basis of the gradu- 
ally dilated and not apically constricted corolla 
tube. 

Two additional collections with ovate acumi- 
nate leaves appear to be related to B. burchellii 
but cannot be placed with certainty. One of these, 
Ducke s.n. (RE 18141), was collected at Bragan?a 
in Para and bears small, white, terminal, solitary 
flowers and immature leaves. The other, Sucre & 
da Silva 9204 (F, RB), was collected at Buriti dos 
Lopes, Piaui, and has rather shiny, thick leaves 
and immature fruits completely enclosed in per- 
sistent, accrescent calyces. 

3. Brunfelsia chocoensis Plowman, Bot. Mus. 
Leafl. 23(6): 245, t. 14. 1973. Type: CO- 
LOMBIA. Antioquia: Chigorodo, forest just 
southeast of Chigorodo, 45 km S of Turbo, 
alt. 50 m. Shrub 2 m high. Flowers rather 
conspicuous, of a peculiar flat white color, 
like unglazed paper, 15 Apr. 1945, Haught 
4563 (holotype, uc M 048365; isotypes, NY, 
COL 109727). Figure 13. 



40 



FIELDIANA: BOTANY 





BRUNFELSIA 




burchellii Plowman 



FIG. 1 2. Brunfelsia burchellii. Reproduced courtesy of the Field Museum of Natural History. 



PLOWMAN: REVISION OF BRUNFELSIA 



41 



BRUNFELSIA chocoensis 

Plowman 




FIG. 13. Brunfelsia chocoensis. Reproduced courtesy of the Botanical Museum of Harvard University. 



Shrub 2 m tall. Branches sparse, terete, some- long, 9-11 cm wide, elliptic to oblong, abruptly 

what knobby at nodes, naked, gray or grayish to long acuminate at apex, acumen often subfal- 

brown. Branchlets 3 mm in diameter, glabrous, cate, obtuse at base, glabrous, dull, dark green 

shiny, light grayish brown. Leaves few, usually above, pale green beneath, firmly membrana- 

crowded at ends of branchlets, blade 18-28 cm ceous, lateral nerves 6-7, spreading; petiole 10- 



42 



FIELDIANA: BOTANY 



15 mm long, stout, brown. Inflorescence subses- 
sile, terminal, 1 -flowered, or, on previous branch- 
es, axillary with 1-3 flowers. Flowers white, fra- 
grant. Bracts 2-4 mm long, ovate to ovate-lan- 
ceolate, concave, ciliolate at margin, caducous. 
Pedicel 5-6 mm long, erect, slender, glabrous. 
Calyx 8-12 mm long, 3-6 mm in diameter, tu- 
bular-campanulate, globose in bud, glabrous, firm- 
ly membranaceous, veinlets prominulous, reticu- 
late, teeth 2-4 mm long, erect, ovate, blunt to 
acute; calyx in fruit to 13 mm long, persistent, 
dotted with lenticels. Corolla tube 22-27 mm 
long, 2-2.5 mm in diameter, twice as long as ca- 
lyx, gradually dilated toward mouth, glabrous; 
limb 22-25 mm in diameter, spreading from 
rounded mouth 5 mm in diameter, the lobes 8-10 
mm long, subequal, rounded at apex, rarely emar- 
ginate, slightly overlapping at margin above base. 
Stamens included in upper third of tube; filaments 
5 mm long, somewhat flattened, lower pair curved 
at apex, upper pair suberect, glabrous; anthers 2 
mm long, orbicular-reniform, light brown. Ovary 
2.5 mm long, ovoid; style 10-20 mm long, slen- 
der; stigma 1.5 mm long, bifid, gaping, the upper 
lobe somewhat larger. Fruit 2-3 cm long, 2-3 cm 
in diameter, globose to ovoid, smooth, yellow 
when ripe, mesocarp soft, fleshy, up to 1.5 mm 
thick, endocarp thin, cartilaginous. Seeds few, 2- 
5, 8-10 mm long, 6-7 mm in diameter, ovoid- 
ellipsoid, angular, dark brown, reticulate-pitted. 
Embryo 8 mm long, straight; cotyledons 3.5 mm 
long, widely elliptic. 

DISTRIBUTION Panama, Colombia. See Figure 
15. 

SPECIMENS EXAMINED PANAMA. Darien: 
Cerro Pirre, 11 Apr. 1967, Bristan 566 (ECON, 
osu), Bristan 569 (ECON, osu), * Bristan 1220 
(MO), *Duke 5300 (MO), ^Gentry & Clewell 7138 
(MO); *along Rio Armila, WSW of Puerto Obal- 
dia, Mori 6828 (MO). COLOMBIA. Choco: Hy- 
dro Camp No. 14, Rio Salaqui, 6 days upstream 
from Rio Sucio, alt. ca. 200 m, 23 May 1967, 
Duke 11351 (ECON); *hills below first rapids on 
Rio Truando, Duke 13331 (MO). 

Brunfelsia chocoensis is restricted to the north- 
ernmost part of the Choco region of Colombia in 
the low-lying basin of the Rio Atrato, an area 
bound by the Serrania de Baudo in the west and 
the Serrania de Abibe in the east. Its range ex- 
tends northward to Cerro Pirre across the Pana- 
manian border. Although additional collecting 
may reveal a larger area for the species, B. cho- 



coensis appears to be endemic at this low eleva- 
tion (50-200 m) in swampy forest. In 1945 it was 
reported as a common species at Chigorodo, the 
type locality. I failed to find the plant on a col- 
lecting trip there, primarily because of the large- 
scale destruction of the forests for agriculture and 
pasturage. 

Most closely related to B. guianensis of the 
Guianas and northern Brazil, this species differs 
in having much larger leaves with longer petioles, 
longer pedicels, and broader corolla lobes. The 
fruit of B. chocoensis is very similar to the fleshy 
capsules of both B. guianensis and B. macrocar- 
pa. Although B. chocoensis was originally 
thought to be related to B. macrocarpa, it now 
seems that the resemblance is only superficial. 
The flowers of B. macrocarpa are very large and 
purple, fading to white with age in the manner of 
species of section Franciscea. In addition, the ca- 
lyx of B. macrocarpa is accrescent and encloses 
the mature fruit, whereas the calyx in B. chocoen- 
sis is small and present only at the base of the 
fruit. 

Data from herbarium labels (Bristan 566 and 
569) indicate that the fleshy, yellow fruits of this 
species are edible and that the flowers are fra- 
grant. 

4. Brunfelsia clandestina Plowman, Fieldiana 
Bot. n.s. 8: 11. 1981. Type: BRAZIL. Bahia: 
Municipio Itapebi, Fazenda Lombardia, BR 
101 ao lado leste. Arbusto de 3 m de altura. 
Flor branca. Capoeira, 12 Aug. 1971, T. S. 
dos Santos 1777 (holotype, CEPEC 7086; iso- 
type, F 1849031). Figure 14. 

Shrub or treelet to 6 m tall. Trunk to 8 cm in 
diameter. Bark on trunk and branches yellowish 
brown, cracking longitudinally and transversely, 
shedding in thin, irregular flakes. It ranch lets 1 .5- 
2 mm in diameter, glabrous, grayish to dark, red- 
dish brown, more or less shiny, cracked longitu- 
dinally. Leaves short petiolate, the blade 35-120 
mm long, 15-50 mm wide, elliptic to oblong-lan- 
ceolate, rarely lanceolate or obovate, apically 
acute to acuminate, the apex itself obtuse, basally 
acute or obtuse, glabrous or rarely sparsely pu- 
berulent on the costa beneath, medium green 
above, somewhat lighter green beneath, shiny on 
both surfaces, chartaceous to subcoriaceous, rare- 
ly coriaceous, the lateral nerves 6-8, mostly 
straight, forming an angle of 45-70 with midrib, 
anastomosing with the arcuate marginal nerve 3- 
6 mm from margin; petiole 26 mm long, glabrous 



PLOWMAN: REVISION OF BRUNFELSIA 



43 



BRUNFELSIA clandestiiia Plowman 




FIELDIANA: BOTANY 



or with scattered glandular hairs. Inflorescence 
terminal or subterminal on mature or newly 
formed twigs of current season, with or without a 
short peduncle, 1- or 2-flowered. Bracts 0-3, 2- 
6 mm long, linear or cymbiform, truncate at apex, 
sparsely pubescent or glandular-pubescent, cadu- 
cous. Flowers white. Pedicel 3-9 mm long, 1 mm 
in diameter, glabrous, becoming thicker in fruit, 
to 3 mm in diameter, warty-lenticellate. Calyx 8- 
16 mm long, 3-7 mm in diameter, tubular or tu- 
bular-campanulate, terete, glabrous, light green, 
membranaceous, teeth 2-5 mm long, subequal, 
triangular-ovate, apically acute or acuminate, the 
apex itself blunt and minutely glandular-papillose; 
calyx in fruit persistent, 11-16 mm long, shiny, 
coriaceous, striately nerved, tightly enclosing ba- 
sal half of capsule. Corolla 18-25 mm long, 1-3 
mm in diameter, tube 1.5-2 times as long as ca- 
lyx, straight, glabrous, rarely with a few glandular 
hairs, orifice 3-5 mm across; limb 15-26 mm in 
diameter, spreading, inclined, lobes 6-10 mm 
long, subequal, expanding somewhat with age, 
broadly obovate to rounded. Stamens inserted in 
upper part of corolla tube; filaments 0.6-1 mm 
wide, strap-shaped, the anterior pair 3-5 mm long, 
incurved at apex, included, the posterior pair 1 .5- 
3 mm long; anthers 1 mm in diameter, orbicular- 
reniform. Ovary 1-1.5 mm long, 0.8-1.2 mm in 
diameter, conical-ovoid, with about 10-12 ovules 
per locule; style 16-20 mm long, incurved at 
apex; stigma 1-1.5 mm long, briefly bifid, upper 
lobe slightly larger. Capsule dry at maturity, 13- 
20 mm long, 13-15 mm in diameter, ovoid to 
subglobose, apiculate, smooth, shiny, dark green 
at maturity; pericarp thin, 0.5-1 mm thick, crus- 
taceous. Seeds ca 9-15, 5-7 mm long, 2.5-3 mm 
in diameter, oblong-reniform, terete, or somewhat 
flattened on one side, dark brown, reticulate-pit- 
ted. Embryo 3-6 mm long, straight; cotyledons 
1-2 mm long, ovate to elliptic, radicle 2-4 mm 
long. 

DISTRIBUTION States of Bahia and Espirito 
Santo, Brazil. See Figure 15. 

SPECIMENS EXAMINED BRAZIL. Bahia: 
"Prov. Jacobina," 1841, Blanchet 3354 (G, LE, w), 
1843, Blanchet s.n. (c, G, w); "Igreja Velha," 
1841, Blanchet 3352 (BM, BR, G, P); Serra de Sin- 
cora, Brejao a Iracema, 17 Feb. 1943, Froes 
20210 (IAN, NY, us); Municipio Jaguaquara, Ja- 



guaquara a Apuerama, 4 Oct. 1972, Pinheiro 1980 
(CEPEC, F, NY); Municipio Itacare, Itacare-Ubai- 
taba, 14 Apr. 1980, T.S. dos Santos 689 (CEPEC, 
F); Rodovia BA 654, km 6 ao oeste de Itacare, 
approx. 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, 
1981 Eds.}. 

Brunfelsia clandestina superficially resembles 
several other brunfelsias and may be confused 
with them. Complete specimens, including field 
data and flower color, are essential for making 
positive identifications. Brunfelsia clandestina ap- 
pears to be most closely related to B. martiana, a 



FIG. 14. Brunfelsia clandestina. Reproduced courtesy of the Field Museum of Natural History. 



PLOWMAN: REVISION OF BRUNFELSIA 



45 




FIG. 15. Distribution of species in section Guianenses: Brunfelsia amazonica (open square), B. burchellii (open 
triangle), B. chocoensis (open circle), B. clandestina (solid triangle), B. guianensis (solid square), and B. martiana 
(solid circle). 



species that also grows in the moist forests of Ba- 
hia. Brunfelsia clandestina differs in having much 
smaller leaves with fewer lateral nerves and one- 
or two-flowered inflorescences that are borne ter- 
minally. In dried specimens, B. clandestina may 
also be confused with B. uniflora, a species 
known from Bah fa but belonging to a different 
section of the genus (section Francisced). Brun- 
felsia clandestina differs mainly in having dark 
brown or reddish brown branchlets, glabrous or 
nearly glabrous leaves and twigs, and a tubular- 
campanulate rather than narrowly tubular calyx. 
The flowers of B. clandestina are white; those of 
B. uniflora are violet, fading to white with age. 

Brunfelsia clandestina grows in the moist 
coastal forests of southern Bah fa and Espirito 
Santo. Earlier collectors in Bahia also found this 



species further inland in areas formerly covered 
with mesophytic forest. Most of these areas have 
now been converted to pastures (Mori & Mattos 
Silva, 1979). The last collection in the drier, in- 
terior part of Bahia was made in 1943. 

5. Brunfelsia guianensis Benth. in DC., Prodr., 
10:200. 1846. Lectotype (designated here): 
SURINAM. Without locality or date, Host- 
mann 1278 (lectotype, K; isolectotypes, G, s, 
w). Lectoparatype: FRENCH GUIANA. Cay- 
enne, 1820, Perrottet s.n. (G, G-DC, non vidi; 
photograph, NY). Figure 16. 

Shrub to treelet 1-3 m tall. Branches lax, 
spreading. Branchlets terete, becoming knobby at 
nodes, glabrous, rarely pubescent, epidermis split- 



46 



FIELDIANA: BOTANY 



BRUNFELSIA guianensis Benth 




FIG. 16. Brunfelsia guianensis. Reproduced courtesy of the Linnean Society, London. (First published in Plowman, 
1978: Botanical Journal of the Linnean Society, 76: 294-295.) 

ting in longitudinal cracks, eventually shedding in glabrous, dull, dark green above, paler green be- 

flakes, light to dark brown. Leaves with blade 6- neath, firmly membranaceous, lateral nerves 4-5, 

15 cm long, 2-6.5 cm wide, obovate to elliptic, arcuate; petiole 3-8 mm long, slender, glabrous, 

rarely lanceolate, abruptly acuminate at apex, acu- rarely pubescent. Inflorescence axillary or ter- 

men often subfalcate, cuneate to narrowed at base, minal, with 1-2 flowers. Flowers white, odorless. 



PLOWMAN: REVISION OF BRUNFELSIA 



47 



Bracts 1-2 per flower, 2-10 mm long, linear-lan- 
ceolate or squamiform, glabrous, caducous. Ped- 
icel 2-5 mm long, slender, short, glabrous. Calyx 
7-10 mm long, 4-7 mm in diameter, ovoid-cam- 
panulate, inflated, rarely tubular-campanulate, 
ovoid in bud, glabrous, green, membranaceous, 
sparsely veined, teeth usually unequal, two pos- 
terior teeth somewhat shorter, 2-5 mm long, 
ovate, acute to acuminate at apex; calyx in fruit 
persistent, becoming somewhat coriaceous, split- 
ting on 1 or more sides. Corolla tube 20-26 mm 
long, 1-1.5 mm in diameter, 2.5-3 times as long 
as calyx, slender, straight or curved, gradually di- 
lated above middle, glabrous, greenish white to 
white, limb 15-22 mm in diameter, spreading, 
lobes 6-8 mm long, subequal, oblong-obovate to 
spathulate, narrowed at base, rounded or blunt at 
apex, lateral margins strongly reflexed, white, oc- 
casionally with traces of purple. Stamens includ- 
ed within tube; filaments slender, suberect, ligu- 
late, anterior pair 4-5 mm long, posterior pair 2- 
4 mm long; anthers 1 mm in diameter, orbicular- 
reniform. Ovary 1.5-2 mm long, 1 mm in di- 
ameter; ovoid-conical, style slender, broadened to- 
ward apex; stigma included between anthers, brief- 
ly bifid, upper lobe slightly larger. Fruit 2.5-3.2 
cm in diameter, 2.5-4 cm long, globose to ovoid, 
smooth, yellow when ripe, exocarp thin, rubbery, 
mesocarp soft, fleshy, 5-8 mm thick, endocarp 
thin, cartilaginous. Seeds 7-10, 10-13 mm long, 
5-7 mm in diameter, oblong, flattened on 1 or 2 
sides, angular, reddish brown to nearly black, re- 
ticulate-pitted. Embryo 8 mm long, straight; cot- 
yledons 2 mm long, elliptic. 

DISTRIBUTION Surinam, French Guiana, Bra- 
zil. See Figure 15. 

SPECIMENS EXAMINED SURINAM. In monti- 
bus Nassau, in forest near km 0.1, 17 Feb. 1949, 
Lanjouw & Lindeman 2137 (K, NY); near km 6, 7, 
10 Mar. 1949, Lanjouw & Lindeman 2564 (K, NY); 
along creek with large rapids, 24 Mar. 1949, Lan- 
jouw & Lindeman 2888 (K, NY); Saramacca River, 
Jacob Kondre, 19 Jun. 1944, Maguire 23894 (NY); 
Jodensavanne-Mapane creek area; Dist. Broko- 
pondo, N of Brokopondo, 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 
4631 (F). 

Brunfelsia guianensis was for a long time con- 
sidered an anomalous species among the South 
American brunfelsias. With the recent discovery 
of several related species, it is now known to be- 
long to a distinct section of the genus, of which 
it is the type species (Plowman, 1978). Brunfelsia 
guianensis differs from other species in this group 



48 



FIELDIANA: BOTANY 



in having obovate leaves and a somewhat inflated, 
ovoid-campanulate calyx. The fruit is large and 
fleshy and yellow at maturity. 

A lowland tropical forest species, B. guianen- 
sisis is distributed throughout the lower Amazon 
basin, chiefly north of the river, reaching well into 
the Guianas. It is known to flower from October 
to April, and fruits have been collected throughout 
the year. In the Brazilian state of Para, this species 
is used in local medicine under the name manaca 
with properties similar to those of B. uniflora (see 
LeCointe, 1947; Plowman, 1977). 

6. Brunfelsia martiana Plowman, Bot. Mus. 
Leafl. 24(20): 37. 1974. Type: BRAZIL. 
Provinciae Rio Negro (Amazonas): habitat in 
sylvis ad flum. Japura prope Sao Joao do 
Principe, Dec. 1819, Martins 3247 (holotype, 
M; isotypes, M). Figure 17. 

Shrub to 1 m tall. Branches few, naked below, 
somewhat knobby at nodes, glabrous. Bark 
cracked longitudinally, dark reddish brown, shiny, 
furnished with lenticels. Leaves scattered along 
branchlets, subsessile, blade 10-25 cm long, 4-8 
cm wide, mostly oblong, sometimes elliptic-ob- 
long or oblong-obovate, acuminate at apex, broad- 
ly cuneate to blunt at base, glabrous on both sides, 
upper surface dark green, nitid or dull, lower sur- 
face paler green, sometimes nitid, firmly membra- 
naceous to subcoriaceous, midrib dark reddish 
brown, lateral nerves 8-12, spreading, straight; 
petiole very short, 1-4(12) mm long, glabrous, 
dark brown, becoming cracked, rugose. Inflores- 
cence terminal or axillary in the upper leaf axils, 
glabrous, axis 2-10(15) mm long. Flowers 1-7 
per inflorescence, sometimes with short peduncle 
that articulates with pedicel, greenish white to 
white. Bracts 1-3 per flower, 1-10 mm long, lin- 
ear-lanceolate, concave, ciliolate at margin, ca- 
ducous. Pedicel 3-6(10) mm long, short, slender, 
glabrous. Calyx 8-12 mm long, 3-8 mm in di- 
ameter, tubular to tubular-campanulate, glabrous, 
rarely striately veined, teeth 2-6 mm long, sub- 
equal, triangular-ovate to ovate-lanceolate, acute 
to acuminate at apex. Corolla tube 20-24 mm 
long, 1-3 mm in diameter, twice as long as calyx, 
straight, cylindric, inflated at apex, glabrous; limb 
15-22 mm across, spreading, somewhat undulate, 
lobes 5-10 mm long, subequal, oblong-obovate, 
abruptly reflexed at lateral margins, rounded-sub- 
truncate at apex. Stamens included within upper 
part of corolla tube; filaments subligulate, anterior 
pair 4 mm long, posterior pair 3 mm long; anthers 



about 1 mm in diameter, orbiculate-reniform, 
slightly unequal, the upper pair somewhat smaller. 
Ovary 2 mm long, oblong-ovoid; style about 15 
mm long, filamentous, a little broader and curved 
at apex; stigma about 1 mm long, briefly bifid, 
upper lobe slightly larger. Fruit and seed incom- 
pletely known. 

DISTRIBUTION Guyana, Brazil (Amazonas, Ba- 
hia, Maranhao, Para). See Figure 15. 

SPECIMENS EXAMINED GUYANA. Essequibo- 
Demerara River, Madray-Bubu Trail, Topy Trysil 
Forest, 8 Feb. 1944, Forest Department 4422 (K). 
BRAZIL. Amazonas: Manaus and vicinity, road 
Manaus-Caracarai, km 22, 21 Mar. 1967, Prance 
et al. 4704 (wis); Rio Negro, Cachoeira Baixa de 
Tarumo, 11-14 Apr. 1973, Schultes & Rodriguez 
26132A (ECON), Schultes & Rodrigues 26133A 
(ECON); *Mun. de Barcelos; Rio Demeni, Froes 
28393 (IAN); *vicinity of Manaus, Prance et al. 
11611 (MO). Bahia: Dist. Ilheus, Ferradas, Dec. 
1818, Martius s.n. (M); Feira de Santana, Apr. 
1850, collector unknown (G). Maranhao: Ilha 
Sao Luis; estrada que vai do Rio Anil para Maio- 
ba, Froes 25650 (IAN); Mun. Sta. Luzia; Fazenda 
Cacique, W of Sta. Inez and E of Entroncameen- 
to, Taylor et al. 1084 (F). Para: Faro, 22 Jan. 
1910, Ducke s.n. (MG); Bella Vista, Rio Tapajoz, 
matta das immediacoes da Campina do Perdido, 
12 Feb. 1917, Ducke s.n. (MG); Regiao do Jutah 
y de Almerim. Palhal, 16 Apr. 1923, Ducke s.n. 
(RB); Gurupa, 25 Feb. 1923, Ducke s.n. (RB); 
Para, May 1819, Martius 3300 (M). Without 
state: Schott 5361? (w), Schott s.n. (w). 

Brunfelsia martiana belongs to section Gui- 
anenses. The type specimen collected by Martius 
(Schedulae 3247) on the Rio Japura includes a 
handwritten description of the plant that Martius 
apparently intended to publish as B. acuminata, a 
name that was later used by Pohl and Bentham 
for a form of B. brasiliensis. 

Brunfelsia martiana has been very poorly col- 
lected, as is true of many of the species of section 
Guianenses. The few collections available were 
collected over an extensive geographical area 
spanning the entire breadth of Brazil, and nearly 
all are preserved in European and Brazilian her- 
baria. The species has been collected only twice 
in the last 50 years; no collections have been 
made in the state of Bahia since 1850. One ap- 
parently quite early collection with six duplicates 
is preserved in Vienna. The label bears only the 
data "Schott 5361, Brasilia." This appears to be 



PLOWMAN: REVISION OF BRUNFELSIA 



49 




BRUNFELSIA martiana Plowman 

FIG. 17. Brunfelsia martiana. Reproduced courtesy of the Botanical Museum of Harvard University. 



50 



FIELDIANA: BOTANY 



the wrong label for the specimen because Schott 
collected only in the state of Rio de Janeiro and 
no other collection of the species has ever been 
made there. Mislabeling of the collections of 
Schott and Pohl has been recognized since the 
time of Martius, who made a special note of it in 
Flora Brasiliensis [1(1): 82 (1906)]: "Dolendum 
quod, quae specimina a cl. POHL, quae a cl. 
SCHOTT collecta sint, ex plagulis herbarii pala- 
tini Vindobonensis non semper statui potest." 

The origin of this collection remains question- 
able, yet it must be taken into consideration be- 
cause it represents one of the more complete gath- 
erings of the species and includes an immature 
fruit. 

Brunfelsia martiana is a small shrub with 
greenish white flowers. It is related to both B. 
amazonica and to B. guianensis and occurs sym- 
patrically with both of these species. Brunfelsia 
martiana differs from B. amazonica in having 
larger leaves (10-25 vs. 6-12 cm) with more lat- 
eral nerves (8-12 vs. 5-8) and much shorter ped- 
icels (3-6 vs. 13-20 mm). 

Brunfelsia martiana is closely allied with B. 
guianensis as well, and specimens of the former 
were originally considered to be exceptional 
forms of B. guianensis. Additional material 
proved the distinctiveness of B. martiana. It is 
now distinguished from B. guianensis by having 
larger (10-25 vs. 6-15 cm), oblong leaves with 
more lateral nerves (8-12 vs. 4-5), a calyx that 
is usually narrow-tubular instead of ovoid-cam- 
panulate, and a corolla tube that is rarely more 
than twice as long as the calyx. In B. guianensis 
it is frequently 2%-3 times as long. 

The flowers of all three of these related species 
are remarkably similar in the shape and color of 
the corolla, especially in the curious form of the 
corolla lobes, the lateral margins of which are de- 
flexed, giving them a rectangular appearance. The 
fruit of B. guianensis is a thick-walled, fleshy cap- 
sule; that of B. amazonica is thin-walled and dry 
at maturity. The collection labeled Schott 5361 
mentioned above includes a small, immature fruit 
enclosed by a coriaceous and striately veined ca- 
lyx. The pericarp appears cartilaginous but not 
fleshy in this early stage. 

A second collection, which for B. martiana has 
exceptionally small leaves reminiscent of B. gui- 
anensis, was made by Martius near Belem do 
Para. This specimen included the remnants of a 
fruit that is small, definitely thin-walled, and brit- 
tle, perhaps the remains of the endocarp only. 
Martius tentatively named this plant B. flexuosa 



and wrote a description (Schedulae 3300) that ap- 
pears on the same page attached to the type spec- 
imen of B. martiana. The nature of the fruit in 
this species remains problematical. When, indeed, 
it is fully known, it should be very useful in de- 
termining the relationship of B. martiana to other 
species. 

Brunfelsia martiana is one of the most widely 
distributed species of section Guianenses, grow- 
ing in tropical lowland forests from Guyana south 
to Bahia in Brazil and westward to the Colombian 
frontier. The curious distribution between the Am- 
azon basin and the coastal forests of Bahia has 
been observed in other groups of plants (Plow- 
man, 1979). 

Brunfelsia sect. Franciscea (Pohl) Griseb., Fl. 
Brit. W. I.: 432. 1861. Franciscea Pohl, PI. 
Bras. Icon. Descr. 1: 1, t. 1-7. 1826. Lecto- 
type species (designated here): Franciscea 
uniflora Pohl. 

TYPE SPECIES Brunfelsia uniflora (Pohl) D. 
Don, Edinburgh New Philos. J. 86. Jul. 1829. 

Shrubs or small trees, 0.3-10 m tall. Inflores- 
cence terminal, rarely axillary. Flowers large or 
small, 1-many per inflorescence. Corolla purple 
fading to pale violet or white with age, with white 
eye at mouth, tube 1-4 times as long as calyx, 
slightly inflated near apex, then slightly constrict- 
ed to form thickened ring, the lobes broadly 
rounded, overlapping at lateral margins. Stigma 
briefly bifid, in the form of a forceps. Anthers 
orbicular-reniform in outline. South and Central 
America. 

Note: Pohl, in establishing his new genus Fran- 
ciscea, included seven species. In recognizing this 
name as a section of Brunfelsia, it is necessary to 
designate a type species. I have chosen B. uniflora 
(Pohl) D. Don for the following reasons. It is the 
oldest known species of the genus, having ap- 
peared in 1 648 in the work of Piso under the name 
Manacd, accompanied by a good illustration 
showing the flowers and fruits (Fig. 18). The ho- 
lotype collected by Pohl and preserved in Vienna 
is a reasonably good specimen showing the char- 
acters of the species. The species is one of the 
most widespread of the genus and is well repre- 
sented in herbaria. Lastly, B. uniflora (as B. ho- 
peand) was the only species mentioned by Gri- 
sebach (1861) in setting up Franciscea as a sec- 
tion of Brunfelsia. 



PLOWMAN: REVISION OF BRUNFELSIA 



51 




FIG. 18. Manaca (Brunfelsia uniflora). From Piso, W., De Medicina Brasiliensis (1648). 



52 



FIELDIANA: BOTANY 



Although the date 1827 appears on the title 
page of Pohl's work Plantarum Brasiliae Icones 
et Descriptiones, it has been pointed out (Stafleu, 
1967; Stafleu & Cowan, 1983) that the first part 
of the work, including the text and plates of Fran- 
ciscea, actually appeared in August 1826. 

7. Brunfelsia australis Benth. in DC., Prodr. 10: 
200. 1846. Lectotype (designated by Hunt, 
1978): BRAZIL. Brasilia meridionalis, Rio 
Uruguay, Baird s.n. (K). Lectoparatypes: AR- 
GENTINA. Buenos Aires, in hortis Bonarien- 
sibus sub nomine Jasmini Paraguayensis cul- 
ta, Tweedie s.n. (K); Estrada de Minas, Mar- 
tius s.n. (?). No collection of Martius with 
this data has been located. Query was made 
by Bentham. Sellow 1573, also cited by Ben- 
tham, is excluded as a type; it belongs to B. 
pilosa Plowman. Figure 19. 

Franciscea australis (Benth.) Miers, Ann. Mag. Nat. 
Hist. ser. 2, 5: 250. 1850. 

Brunfelsia hopeana var. australis (Benth.) J.A. 
Schmidt in Mart., Fl. Bras. 8(1): 262. 1864. 

Brunfelsia paraguayensis Chodat, Bull. Herb. Bois- 
sier, ser. 2, 1: 406. 1901. Type: PARAGUAY. In 
silvis pr. Rio Apa, May, without year, Hassler 
725a (holotype, G). 

Brunfelsia uniflora f. typica Hassl., Feddes Repert. 
Spec. Nov. Regni Veg. 15: 243. 1919. Non sensu 
D. Don. Type: PARAGUAY. In silvis pr. Rio 
Apa, May, without year, Hassler 725a (holotype, 

G). 

Brunfelsia uniflora f. obovatifolia Hassl., Feddes Re- 
pert. Spec. Nov. Regni Veg. 15:243. 1919. Type: 
PARAGUAY. Cordillera de Altos, in silvis, fru- 
tex vel arbor parva 3-5 m, corolla albo-coeru- 
lescens vel violacea in eadem planta, Sep. 1898- 
1 899, Hassler 3246 (holotype, o; isotypes, A, G, 
NY, P, w). 

Brunfelsia uniflora f. intermedia Hassl., Feddes Re- 
pert. Spec. Nov. Regni Veg. 15: 243. 1919. Lec- 
totype: PARAGUAY. In regione lacus Ypacaray, 
Cordillera do Altos, arbusto 2-2.5 m, fl. lila- 
morado, monies y orillas, Cordilleras y bajos, 
azucena, Sep. 1913, Hassler 12257 (lectotype, 
G; isolectotypes, c, GH, K, L, MO, NY). Lectopar- 
atypes: PARAGUAY. In regione lacus Ypacaray. 
Picada a Bernalcue, Mar. 1913, Hassler 12147 
(A, G); San Bernardino, orillas monies, fl. morado 
despues blanca, arbusto 3-4 m, Aug. 1915, 
Hassler (cited as Rojas) 1333 (G). 

Shrub to small tree 2-4 m tall. Trunk often 
branched from near base, to 8 cm in diameter. 
Bark rugose, yellowish brown. Branches ascend- 
ing and spreading, leafy, glabrous, green with yel- 
lowish brown cracks, furnished with lenticels. 
Branchlets glabrous, green. Leaves scattered 
along branchlets, blade 4.0-12.5 cm long, 2.5-6.0 



cm wide, broadly elliptic to obovate, rarely orbic- 
ular or rhombic, blunt to rounded at apex, occa- 
sionally acute, cuneate at base, often briefly pu- 
bescent-ciliate when young, becoming glabrous, 
or sparsely glandular beneath at costa, dull dark 
green above, pale green beneath, membranaceous 
to subcoriaceous, lateral nerves 5-7, prominulous 
beneath, straight or somewhat arching, veinlets 
often obscure; petiole 3-8 mm long, glabrous or 
sparsely pubescent. Inflorescence terminal at the 
tips of the branchlets, sessile, with 1-4 flowers. 
Bracts present or absent, to 2 mm long, lanceo- 
late, concave, ciliolate, early caducous. Flowers 
showy, violet, fading to pure white with age, fra- 
grant. Pedicel stout, 4-7 mm long, glabrous, be- 
coming thicker and corky-verrucose in fruit. Ca- 
lyx 7-12 mm long, 4-10 mm diameter, campan- 
ulate, more or less inflated, glabrous, rarely pa- 
pillose-punctate, pale green, firmly membrana- 
ceous, reticulately veined, teeth 3-6 mm long, 
ovate-lanceolate, acute; calyx in fruit coriaceous, 
partially enclosing fruit, smooth. Corolla tube 
25-32 mm long, 1.5-2.0 mm in diameter, 2-3 
times as long as calyx, glabrous, often sparsely 
glandular in bud, glabrescent; limb 30-40 mm in 
diameter, plane, spreading, thickening at mouth 
prominent, round, white or yellowish, lobes equal, 
broadly rounded, overlapping laterally, abruptly 
narrowed at base. Stamens included; filaments 
slender, upper pair 2-3 mm long, lower pair 3-5 
mm long; anthers about 1 mm in diameter, reni- 
form-orbicular. Ovary 2.5 mm long, 1 mm in di- 
ameter, ovoid-conical; style 20-28 mm long; stig- 
ma about 1 mm long. Capsule 10-22 mm long, 
10-20 mm in diameter, subglobose to ovoid, apic- 
ulate at apex, smooth, glabrous, dark green, 
pericarp to 0.5 mm thick, dry at maturity, inde- 
hiscent or tardily dehiscent. Seeds 20-40, 4-6 
mm long, 3-4 mm in diameter, oblong-ellipsoid, 
angular, dark brown. Embryo 3-4 mm long, 
slightly curved; cotyledons ovate. 

DISTRIBUTION Brazil (Parana, Rio Grande do 
Sul, Sao Paulo), Paraguay, Argentina (Chaco, 
Corrientes, Formosa, Misiones, Santa Fe), and 
Uruguay; cultivated in some areas, especially in 
Central America. See Figure 20. 

SPECIMENS EXAMINED BRAZIL. Parana: 
*Municipio Foz do Igua9u; Vila Postes, Buttura 
157 (F); *Ceu Azul, Hatschbach 43192 (F). Rio 
Grande do Sul: Neu Wurttemberg, 450 m, 15 
Oct. 1904, Bornmuller 219 (A, G, GH, M, w); Sao 
Leopoldo, 16 Oct. 1934, Rambo 321 (C); Porto 
Alegre, 12 Sep. 1945, Rambo 37819 (A, c, w); 



PLOWMAN: REVISION OF BRUNFELSIA 



53 



Morra da Gloria, Rambo 29147 (BR, MO). *Mun- 
icipio Itaqui, Dobereiner & Tokarnia 841 (RB). 
Sao Paulo: *Parque Estadual do Turvo, Tenente 
Portela, Paiva, Stehmann s.n. (F). PARAGUAY. 
Cordillera de Altos: Cerros de Tobati, 18 Aug. 
1902, Fiebrig 49 (A, F, G, L, M, PR); Hassler 725 
(G); Sep. 1913, Hassler 12559 (A); Villarrica, Sep. 
1931, Jorgensen 3661 (A, c, F, MA, MO, NY, PH, s, 
us); Dep. Cantera, Zona Grande Bosques, Herrera 
Vegas, 220 m, 11 Aug. 1939, Monies 4703 (BAB); 
Dep. San Pedro, Colonia Primavera, 11 Sep. 
1955, Woolston 571 (c, GH, NY, s, uc); without 
locality, 26 Aug. 1874, Balansa 2240 (G, GOET, K, 
LD, LE, PE, s); 4 Sep. 1854, Palmer 155 (us); *Par- 
aguari, Cerro San Jose, a civitate Ybicui, Bernardi 
18138 (F); *Neembucu, Curupaity ad Laureles, 
Bernardi 20495 (F). ARGENTINA. Chaco: Dep. 
Resistencia, Margarita Belen, 8 Sep. 1946, Agui- 
lar 841 (s), 18 Sep. 1948, Aguilar 1302 (w); Dep. 
Primer de Mayo, Colonia Benitez, Oct.-Nov. 
1928, Schulz 85 (BAB), 26 Nov. 1906, Stuckert 
16307 (G); Las Palmas, Jorgensen 2098 (GH, MO, 
uc); Rio de Oro, km 140 de Resistencia, 29 Aug. 
1944, Rojas 12133 (uc). Corrientes: Capital, Feb. 
1821, Bonpland 528 (P); 1908, Llamas s.n. (BAB); 
Quinta La Eloisa, 77 m, 6 May 1969, Plowman 
2713 (ECON, GH). Dep. Conception, Tabay, 1 Nov. 
1965, Krapovickas & Cristobal 11706 (uc); 7 
June 1969, Krapovickas & Cristobal 15385 (c, 
MO); 7 May 1969, Plowman 2723 (ECON, GH); 
Dep. Mburucuya: Estancia "Santa Teresa," 6 Sep. 
1949, Pedersen 418 (BR, c, GH, p, s, us); Estancia 
"Santa Maria," Campo "Dios Gracia," 18 Dec. 
1949, Pedersen 418 (BR, c, NY, p, s); Estancia 
Santa Ana, 21 Oct. 1944, Schwarz 93 (A); Canada 
Paso Clarios, 11 Nov. 1949, Schwarz 8661 (BH, 
G, w). Dep. Bella Vista, Bella Vista, Oct. 1904, 
N. Rojas Acosta 13004 (BAB); Dep. Saladas: La- 
guna Salada, 28 Sep. 1944, Schwarz 36 (A); La- 
guna Soto, 28 Sep. 1944, Schwarz 48 (A); Paso 
del Deseo, 1 Nov. 1944, Schwarz 139 (A, uc); 2 
Dec. 1949, Schwarz 8972 (BH, G, w). Dep. Itu- 
zaingo, Salto Apipe, 6 Oct. 1949, Schwarz 8114 
(BR); Dep. General Paz, Laguna Rincon, 2 Nov. 
1949, Schwarz 8547 (LD); Dep. San Roque, San 
Roque cercanias, 20 Apr. 1945, Ybarrola 2963 
(uc, w); Dep. San Luiz del Palmar, Colonia J. M. 
Llano, 19 Sep. 1945, Ybarrola 3270 (B, w); Cer- 
rudo Cue, 5 Sep. 1946, Ybarrola 3430 (G, p); vi- 
cinity of Goya, 15-30 m, 2-10 Nov. 1913, Curran 



s.n. (us); *Dep. Santo Tome, Garruchos, Krapov- 
ickas et al. 26092 (MO); *Dep. San Martin, Carlos 
Pellegrini, Krapovickas 29261 (MO); *Dep. San 
Martin, Sussini, Schinini et al. 18580 (F). For- 
mosa: Dep. Pilcomayo, Estancia Riacho Negro, 3 
Oct. 1947, Morel 3783 (BR, MO); Salvation, 17 
Nov. 1947, Morel 4144 (c); S.O. a 12 km de Fi- 
lipina, Est. Salaberry, 21 Nov. 1949, Morel 8921 
(BH, c, G, w); Formosa, 29 Oct. 1900, Kermes 
1363 (BAB); Fontana, Aug. 1933, Meyer 166 (GH). 
Misiones: Dep. Iguazii, Salto Iguazu, 2 Oct. 1947, 
Pierroti 6563 (G); 6 Oct. 1910, Rodriguez 484 (A, 
us); Dep. Candelaria, Martires, 24 Oct. 1917, Ber- 
toni 3434 (w); Loreto, 26 Sep. 1932, Muniez 99 
(BAB); Dep. San Javier, Acaragua, 16 Sep. 1946, 
Bertoni 2939 (B); Dep. G. M. Belgraw, "El Ca- 
burei," 450 m, 22 Sep. 1957, Monies 27471 (M, 
NY); without locality, Fox s.n. (CGE). Santa Fe: 
Dep. General Obligado, Florencia, 30 Oct. 1923, 
Chiar s.n. (BAB); Villa Guillermina, 14 Feb. 1939, 
Ragonese 3646, (F); Indio Muerto, 6 Mar. 1939, 
Meyer 3076 (A). URUGUAY. Banda Oriental, 
1816-1821, St. Hilaire C-2197 (p). 

CULTIVATED SPECIMENS OF INTEREST BRA- 
ZIL. Rio Grande do Sul: Porto Alegre, 20 Mar. 
1893, C. Lindman A597a (s); Santa Maria, Sil- 
vacultura, 14 Oct. 1955, Michel 8 (B). Rio de Ja- 
neiro: Rio de Janeiro, Quinta de Boa Vista, 13 
Oct. 1879, Glaziou 12110 (BR, c, G, LE, p, R); Rio 
de Janeiro, Oct. 1915, Vincent s.n. (L). Santa Ca- 
tarina: Brusque, 4 Oct. 1961, Reitz & Klein 
11250 (us). ARGENTINA. Tucuman: Capital, 
Institute Lillo, 9 Sep. 1947, Meyer 12621 (B); 
Dep. Trancaas, San Pedro de Colalao, 1,200 m, 
Apr. 1926, Venturi 4408 (A). 

Bentham described B. australis from three 
specimens, two of which are included in the pres- 
ent concept. Baird s.n. was chosen (by Hunt, 
1978) as the lectotype. One of the other speci- 
mens, collected by Tweedie, was made from hor- 
ticultural material in Buenos Aires. The third col- 
lection cited by Bentham, Sellow 1573, is here 
referred to B. pilosa, a closely allied species. A 
fourth collection by Martius (with a query by 
Bentham) cannot be located. The locality "Estra- 
da de Minas" indicates another species was in- 
volved because B. australis does not occur north 
of Rio Grande do Sul in Brazil. 



FIG. 19. Brunfelsia australis. Reproduced courtesy of Curtis 's Botanical Magazine and Christabel King. 



54 



FIELDIANA: BOTANY 







Brunfelsia australis 



CHRISTABEL KING del. 



PLOWMAN: REVISION OF BRUNFELSIA 



55 




FIG. 20. 
circle). 



Distribution of Brunfelsia australis (solid triangle), B. boliviano (solid square), and B. bonodora (solid 



Schmidt in Martius (1864) considered B. aus- 
tralis to be a variety of B. uniflora (B. hopeana}. 
He cited the Sellow collection mentioned above 
and two additional specimens. One of these, Rie- 
del s.n., collected near Macae, Rio de Janeiro, 
agrees with B. bonodora. The other collection, 
Martius s.n., from Para, is referred to B. guianen- 
sis, although most of this material is sterile. 

Brunfelsia australis is closely related to B. un- 
iflora and B. pilosa and by some authors (Schmidt 
1864; Hassler, 1919) has been considered synon- 
ymous with B. uniflora. However, these workers 
suffered from the disadvantage of working exclu- 
sively with dried herbarium specimens (Schmidt) 
or with local populations without understanding 
the entire morphological range of the species in- 
volved (Hassler). Studies of both B. australis and 



B. pilosa in the field and of all three entities in 
cultivation and from herbarium specimens reveal 
that these plants are morphologically, ecological- 
ly, and geographically distinct and merit the rank 
of species. Experimental crosses suggest that re- 
productive isolation exists between B. pilosa and 
B. australis. 

Brunfelsia australis is distinguished from B. 
uniflora and B. pilosa by the broadly elliptic to 
obovate leaves, the variable number (one to four) 
of flowers per inflorescence and the uniformly 
glabrous, campanulate calyx. 

Even though their ranges overlap, almost no in- 
termediates have been found between B. australis 
and B. pilosa. One collection from Rio Grande do 
Sul, Bornmuller 219, has oblong leaves and a 
more deeply incised calyx and may represent 



56 



FIELDIANA: BOTANY 



some introgression from B. pilosa, which also oc- 
curs in this region. 

Commonly growing as a tall shrub or treelet, 
B. australis is found primarily in the relatively 
recent drainage systems of the Rio Parana, Rio 
Uruguay, and Rio Jacui (Rio Grande do Sul), 
growing in the understory of low-elevation forests 
near the rivers and in adjacent uplands. It occurs 
from sea level to 450 m altitude. In the low-lying, 
often swampy river basin of the Rio Parana, B. 
australis is found in the scattered woodlands that 
appear on higher and drier ground. 

Two reports of widely disjunct populations are 
now known to represent cultivated plants. Venturi 
4408 from Tucuman, Argentina, was distributed 
to some herbaria without the notation "culta." 
Specimens of Glaziou 12210 were distributed 
with the fictitious provenance "Rio de Janeiro, 
Serra da Palmital." However, a duplicate speci- 
men at the Museu Nacional de Rio de Janeiro 
bears the correct locality, "Quinta da Boa Vista 
(Rio), Arbusto cult." This erroneous data may be 
the result of direct falsification by Glaziou, which 
was pointed out in the Melastomataceae by Wur- 
dack (1970). 

Brunfelsia australis flowers from August to 
November; mature fruits appear from April to 
June. Frequently the entire plant is covered with 
fragrant, showy purple and white flowers at the 
same time, making this species a valuable orna- 
mental. It is widely cultivated in gardens between 
latitudes 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). 
Figure 21. 

Shrub 1-2 m tall (fide Cardenas). Bark on 
branches thin, yellowish to grayish brown, lon- 
gitudinally rugose, not exfoliating. Branchlets 2- 
3 mm in diameter, more or less villous, becoming 
glabrous. Leaves scattered along branchlets or 
crowded at tips of lateral short shoots, 40-135 
mm long, 23-58 mm wide, blade obovate, rarely 
elliptic-obovate, apically rounded with a short 



acumen 5-10 mm long, the acumen itself blunt to 
acute, somewhat revolute at margin, basally atten- 
uate, glabrescent on both surfaces except at the 
midrib, which bears villous and glandular hairs, 
ciliate-villous on the adaxial surface of the revo- 
lute margin, dull green above, paler, yellowish 
green beneath, firmly membranaceous or some- 
what coriaceous, the lateral nerves 4-9, straight, 
arcuately anastomosing toward the margin, prom- 
inulous beneath; petiole short, 2-6 mm long, more 
or less villous, more densely so on upper side. 
Inflorescence corymbiform, terminal on last sea- 
son's branchlets, short pedunculate, branched, 
with 6-15 flowers. Peduncle 5-12 mm long, more 
or less villous. Bracts small, 14 mm long, lan- 
ceolate or cymbiform, sparsely to densely villous 
especially at margin, caducous. Flowers light vi- 
olet fading to white (fide Cardenas). Pedicel short, 
2-6 mm long, 1 mm in diameter, scarcely thick- 
ened at apex, glabrous or with sparse glandular 
hairs. Calyx 9-15 mm long, 3-4 mm in diameter, 
tubular, truncate at base, 5-angled in cross section, 
appearing plicate in pressed specimens, glabrous 
or with scattered glandular hairs, teeth 2-4 mm 
long, subequal, ovate, apically short acuminate, 
the acumen itself blunt and glandular-papillose; 
fruiting calyx persistent, withering. Corolla tube 
21-25 mm long, 2-3 mm in diameter, about twice 
as long as calyx, straight; limb 18-30 mm in di- 
ameter, spreading, lobes 8-14 mm long, subequal, 
rounded. Stamens inserted in upper part of co- 
rolla tube; filaments strap-shaped, the anterior pair 

3 mm long, suberect, briefly exserted from mouth 
of corolla tube, the posterior pair 2 mm long, in- 
cluded; anthers 1-1.2 mm in diameter, orbicular- 
reniform, subequal or the upper pair slightly larg- 
er. Ovary 2-3 mm long, 1.8-2.2 mm in diameter, 
narrowly ovoid, with about 30 ovules; style 19 
mm long, incurved at apex; stigma 2 mm long, 
briefly bifid. Capsules 3-6 per infructescence, 
20-25 mm long, 20-25 mm in diameter, dry at 
maturity, subglobose, smooth, green; pericarp 
thin, 1-1.5 mm thick, endocarp thin, crustaceous, 
with 22-28 seeds per capsule. Seeds 5-8 mm 
long, 2-3 mm in diameter, oblong to subreniform, 
subterete, dark brown, reticulate-pitted. Embryo 

4 mm long, straight; cotyledons 1.5 mm long, 
ovate; radicle 2.5 mm long. 

DISTRIBUTION The eastern Andes of southern 
Bolivia. See Figure 20. 

SPECIMENS EXAMINED BOLIVIA. Chuquisa- 



PLOWMAN: REVISION OF BRUNFELSIA 



57 



BRUNFELSIA boliviana Plowman 




FIG. 21. Brunfelsia boliviana. Reproduced courtesy of the Field Museum of Natural History. 



58 



FIELDIANA: BOTANY 



ca: mountain above Bartolo, on road from Mon- 
teagudo to Sucre, 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, 
1977). 

9. Brunfelsia bonodora (Veil.) J. F. Macbr., Field 
Mus. Nat. Hist., Bot. Ser. 13, 5-B(l): 152. 
1962. Figure 22. 



Besleria bonodora Veil., Fl. Flumin. 261. 1829 
("1825"). Ic. 6, t. 80. 1831 ("1827"). Lectotype 
(designated here): Vellozo, J.M. da C., Fl. Flu- 
min. Ic. 6, t. 80. 1831 ("1827"). Representative 
collection: BRAZIL. Rio de Janeiro, Serra de Es- 
trella, Mar. 1833, Riedel 84 (LE). 

Shrub 1-2 m tall. Branches ascending, spread- 
ing, naked, glabrous. Bark thin, light brown. 
Branchlets glabrous or sparsely puberulent, gray- 
ish brown. Leaves scattered along branchlets, 
blade 5-14.5 cm long; 2-5 cm wide, narrowly el- 
liptic to lanceolate, acuminate at apex, blunt to 
cuneate at base, glabrous, at times puberulent at 
midrib on lower surface, shiny, dark green on up- 
per surface, pale green beneath, firmly membra- 
naceous, lateral nerves 5-8, more or less arcuate; 
petiole 2-6 mm long, slender, glabrous. Inflores- 
cence terminal, compact, unbranched, axis 3-8 
mm long. Flowers 3-15, rarely reduced to 1 or 2; 
blue fading to white with age, fragrant. Bracts 2- 
12 mm long, lanceolate, glabrous or puberulent, 
caducous. Pedicel 6-12 mm long, slender, thick- 
ened slightly at apex, glabrous. Calyx 11-14 mm 
long, about 5 mm in diameter, tubular-campanu- 
late to campanulate, ellipsoid to obovoid in bud, 
rarely somewhat inflated, glabrous, firmly mem- 
branaceous, striately veined, teeth 3-4 mm long, 
triangular to ovate, acute to blunt at apex, rarely 
apiculate, calyx to 15 mm long in fruit, subcor- 
iaceous, conspicuously veined, enclosing fruit. 
Corolla tube 17-25 mm long, 2 mm in diameter, 
twice as long as calyx, glabrous or sparsely glan- 
dular; limb 20-35 mm in diameter, plane spread- 
ing, lobes rounded to oblong, subtruncate or 
rounded at apex, rarely emarginate, overlapping 
at sides, narrowed abruptly at base. Stamens in- 
cluded in upper part of corolla tube; filaments 
about 4 mm long, subligulate; anthers 1.5 mm 
long, reniform. Ovary 1.5 mm long, ovoid-coni- 
cal; style slender, equaling the filaments; stigma 1 
mm long, briefly bifid, upper lobe slightly larger, 
obtuse. Capsule 8 mm long, 6-8 mm in diameter, 
partially enclosed by calyx, ovoid-turbinate, apic- 
ulate at apex, smooth, pericarp thin-walled, drying 
crustaceous, sparingly dehiscent. Seeds about 5, 
incompletely known. 

DISTRIBUTION Brazil (Rio de Janeiro, Sao Pau- 
lo). See Figure 20. 

SPECIMENS EXAMINED BRAZIL. Rio de Ja- 
neiro: Serra de Estrella, 700 m, 29 Oct. 1946, 
Brade 18643 (RB), 12 Nov. 1823, Riedel 18 (A, 
LE, mixed collection; NY, mixed collection; us), 14 
Dec. 1823, Riedel 59 (A, LE, mixed collection); 



PLOWMAN: REVISION OF BRUNFELSIA 



59 




( Tal) 



60 



FIELDIANA: BOTANY 



Macahe, May 1832, Riedel sp. nov. (LE); Barra 
Mansa, Fazenda Paraiso, 3 Dec. 1960, Duarte 
5477 (RB, us). Sao Paulo: Taubate, Dec. 1817, 
Martius 541 (M). Without state: Riedel 112 (us). 

Brunfelsia bonodora was first described in the ge- 
nus Besleria by Vellozo in Flora Fluminensis, a 
work that appeared in 1829 although 1825 ap- 
pears on the title page (Carauta, 1973). Vellozo's 
scanty description was accompanied by a some- 
what stylized illustration. This drawing must now 
be considered the lectotype of the species since 
no specimens of Vellozo exist (Fig. 22). 

In his monograph of Brunfelsia in 1846, Ben- 
tham erroneously placed Vellozo's species in syn- 
onymy with his B. latifolia (based on Franciscea 
latifolia Pohl). This concept also included mate- 
rial now recognized as B. grandiflora subsp. 
schultesii of the western Amazon and Andes. This 
problem was discussed by Monachino (1953) and 
Macbride (1962), but neither of these workers 
were able to resolve the issue, primarily because 
they saw neither the type materials nor represen- 
tative collections from southeastern Brazil. 

It is now clear that the name B. latifolia and its 
synonym, B. maritima Benth., apply to a distinct 
species known only from the sand restingas 
around the city of Rio de Janeiro. Brunfelsia bon- 
odora is represented by several collections from 
the mountains of the state of Rio de Janeiro that 
agree with Vellozo's description and plate. This 
concept is, however, not identical with Macbride's 
interpretation of the species since he included ma- 
terial here referred to as B. grandiflora subsp. 
schultesii. 

Nevertheless, B. bonodora is related to both B. 
latifolia and B. grandiflora subsp. schultesii, 
which is in part responsible for the confusion of 
these concepts. Brunfelsia bonodora is distin- 
guished from B. latifolia in having a more robust, 
upright habit; longer, lanceolate, and apically acu- 
minate leaves; and longer pedicels. In addition, B. 
bonodora differs ecologically, growing in the 
montane rain forests of the Serra do Mar and not 
in open, coastal formations (restingas). There 
seems to be some overlap in these two species, 
but the status of possible hybrids cannot be de- 
termined with so few specimens at hand, most of 
which lack adequate collection data. In fact, only 



two modern collections of B. bonodora are 
known. 

Macbride (1962) considered B. bondora to be 
conspecific with the plant here treated as B. gran- 
diflora subsp. schultesii. On close examination, 
however, there are several points of difference be- 
tween them. Brunfelsia bonodora has longer ped- 
icels (6-12 vs. 2-5 mm), a more compact inflo- 
rescence, larger bracts, and the capsule almost 
completely enclosed by the calyx in fruit. Ac- 
cording to the specific epithet, B. bonodora is fra- 
grant; B. grandiflora almost never is. 

Geographically, B. bonodora and B. grandiflo- 
ra are separated by several thousand miles. They 
exhibit, however, a distribution similar to those of 
B. hydrangeiformis and B. mire and may be con- 
sidered to be vicarious species that arose from a 
common ancestor. Brunfelsia bonodora is known 
only from the coastal montane rain forests in the 
state of Rio de Janeiro, at elevations up to 700 m. 
It flowers from October to December and fruits 
during May and June. Additional collections of 
this plant with complete field data will greatly aid 
in understanding its specific characters and rela- 
tionships. 

10. Brunfelsia brasiliensis (Spreng.) L. B. Sm. & 
Downs in Reitz, Fl. Ilustr. Catar., pt. 1, Sola: 
303. 1966. 

Gerardia brasiliensis Sprang., Syst. Veg. 2: 806. 
1825. Type: BRAZIL. Brasilia meridionalis, 
without locality, Sellow s.n. (type destroyed at 
Berlin; representative collection and possible 
isotype, M). 12 

Shrub 0.3-2 m tall. Bark roughish, reddish to 
light brown. Branches numerous, arising from 
near base, leafy, yellowish brown, pilose, es- 
pecially so when younger, or glabrous. Branch- 
lets of current year leafy, slender, erect, densely 
pubescent with yellowish brown, nonglandular 
hairs. Leaves scattered along branchlets, some- 
times more crowded toward apex, blade 3-13 cm 
long, 1-4.5 cm wide, lanceolate to oblong-lanceo- 



12 In the manuscript Plowman gave no obvious indi- 
cation that he wished to lectotypify this species as cited. 
We have thus left the lectotypification of this species to 
later Brunfelsia specialists. Eds. 



FIG. 22. Lectotype of Brunfelsia bonodora (originally as Besleria bonodora). From Vellozo, J. M. C, Flora 
Fluminensis (1881). 



PLOWMAN: REVISION OF BRUNFELSIA 



61 



late, more rarely obovate-oblong, acute to acu- 
minate at apex, occasionally blunt, cuneate to nar- 
rowed at base, the upper surface sparingly pubes- 
cent, dark to light green, rarely nitid, subscabrous 
or glabrate, the lower surface variously pubescent, 
often villous at midrib, rarely glabrous, paler 
green, firmly membranaceous to coriaceous, lat- 
eral nerves 6-10, mostly straight; petiole 1-5 mm 
long, puberulent to villous. Inflorescence terminal 
and subterminal, corymbiform, compact and near- 
ly sessile or lax-elongate, with 5-many flowers, 
rarely reduced to 1-2, the axis 1-2 cm long, pu- 
bescent- villous. Bracts small, 2-15 mm long, lan- 
ceolate, ciliolate at margin, pubescent, caducous. 
Flowers often showy, violet fading to pale lav- 
ender or white with age, odorless, short pedun- 
culate. Peduncles 2-8 mm long, thickened at apex 
and articulating with pedicel, persistent, villous or 
glandular-pubescent, bearing 1-3 bracts. Bracts 
minute, 1-2 mm long, linear-lanceolate. Pedicel 
2-15 mm long, erect, pubescence yellowish to 
yellowish brown, sparsely pilose to villose, rarely 
glandular, thickening in fruit. Calyx 8-22 mm 
long, 3-8 mm in diameter, tubular to tubular-cam- 
panulate, with variable yellow to yellowish brown 
pubescence, sparsely to densely villous, rarely pi- 
lose or glandular, light green to purple, firmly 
membranaceous, teeth 1-5 mm long, erect, some- 
times recurved in fruit, triangular to ovate, acute 
at apex; calyx in fruit campanulate to urceolate, 
becoming coriaceous, striate-nerved. Corolla tube 
17-25 mm long, 1-3 mm in diameter, 1-2 times 
as long as calyx, sparsely pilose or glabrous, pale 
violet below, white toward apex, limb 18-32 mm 
in diameter, plane spreading, lobes subequal, 
broadly rounded to truncate at apex, lateral mar- 
gins. Stamens included in upper part of tube; fil- 
aments slender, slightly broader at base, glabrous, 
white, lower pair 2-4 mm long, upper pair 4-5 
mm long; anthers 1 mm long, oblong-reniform. 
Ovary 1-5 mm long, ovoid-conical, green; style 
17-18 mm long, filiform, bright green; stigma 1- 
2 mm long, briefly bifid, the upper lobe somewhat 
larger, bright green. Capsule 8-13 mm long, 8- 
10 mm in diameter, enclosed by calyx, ovoid to 
subglobose, apically apiculate, smooth, shiny, 
light to dark green, pericarp thin, 0.5 mm thick, 
cartilaginous to crustaceous, dry at maturity, spar- 
ingly dehiscent. Seeds (2)7-10, 4-6 mm long, 3 
mm in diameter, oblong-ovoid, more or less an- 
gular, dark reddish brown, reticulate-pitted. Em- 
bryo 4 mm long, very slightly curved; cotyledons 
1.5 mm long, ovate-elliptic. 



DISTRIBUTION Brazil. See Figure 23. 



Key to the Subspecies of Brunfelsia brasiliensis 
(Spreng.) L. B. Sm. & Downs 

1 . Inflorescence terminal, nearly sessile, compact, 
with 1-8 flowers; peduncles shorter than ped- 
icels; pedicels 5-15 mm long; calyx 15-22 mm 
long; leaves often glabrescent 

lOa. subsp. macrocalyx 

1. Inflorescence terminal and subterminal, lax or 
dense, often branched, with 8-many flowers; 
peduncles equaling or longer than pedicels; 
pedicels 2-5 mm long; calyx 7-17 mm long; 
leaves usually pubescent-villous beneath 

lOb. subsp. brasiliensis 

lOa. Brunfelsia brasiliensis subsp. brasiliensis. 
Figure 24. 

Franciscea acuminata Pohl, PI. Bras. Icon. Descr. 1: 
4, t. 3. 1826. Type: BRAZIL. Rio de Janeiro: in 
umbrosis inter frutices ad Mandioca, Schott 6163 
(holotype, BR). 

Brunfelsia acuminata (Pohl) Benth. in DC., Prodr. 
10: 199. 1846. 

Brunfelsia brasiliensis var. acuminata (Pohl) L. B. 
Sm. & Downs in Reitz, Fl. Ilustr. Catar, pt. 1, 
Sola: 305. 1966. 

Franciscea ramosissima Pohl, PI. Bras. Icon. Descr. 
1: 5, t. 4. 1826. Type: BRAZIL. Rio de Janeiro: 
ad Serra Tingua, Schott 3720 (holotype, w; iso- 
types, GH, w). 

Brunfelsia ramosissima (Pohl) Benth. in DC., 
Prodr., 10: 199. 1846. 

Franciscea confertiflora Pohl, PI. Bras. Icon. Descr. 
1: 6, t. 5. 1826. Type: BRAZIL. Minas Gerais: 
inter frutices locis umbrosis in via de Villa Fan- 
ado vel Bom Successo dicta ad Villa do Principe, 
Oct.-Nov. 1820, Pohl 3478 (holotype, w; iso- 
type, w). 

Brunfelsia confertiflora (Pohl) Benth. in DC., 
Prodr., 10: 199. 1846. 

Brunfelsia ramosissima var. confertiflora (Pohl) J. 
A. Schmidt in Mart., Fl. Bras. 8(1): 260. 1864. 

Franciscea divaricata Pohl, PI. Bras. Icon. Descr. 
1: 6, t. 6. 1826. Type: BRAZIL. Minas Gerais: 
inter frutices, locis umbrosis in via de Sao Joao 
d'El Rey, ad Villa Paracatudo Principe, floret ab 
Octubri usque ad Martium, 1818, Pohl 593, 6165 
(holotype, w; isotypes, F, GH LE, w). 

Brunfelsia ramosissima var. laxiflora J. A. Schmidt 
in Mart., Fl. Bras. 8(1):260. 1864. Type: BRA- 
ZIL. Brasilia meridionalis, without locality, Sel- 
low s.n. (holotype, M). 

Brunfelsia ramosissima var. parcifolia J. A. 
Schmidt in Mart., Fl. Bras. 8(1):260. 1864. Type: 
BRAZIL. Minas Gerais: without locality, Claus- 
sen 171 (holotype, BR). 



62 



FIELDIANA: BOTANY 




FIG. 23. Distribution of Brunfelsia brasiliensis (solid square) and B. chiricaspi (solid circle). 



Leaves 3-13 cm long, 1-4.5 cm wide, puber- 
ulent or scabrous above, often glabrescent, vari- 
ably villous beneath, often densely so at midrib, 
firmly membranaceous to subcoriaceous. Inflo- 
rescence variable, dense to lax, occasionally elon- 
gate and well-developed, with 10-many flowers, 
rarely reduced to about 5. Peduncles equaling or 
longer than pedicels, 2-10 mm long, rufo- villous, 
not glandular. Pedicels 2-5 mm long, villous. Ca- 
lyx 7-15 mm long, variably villous with dense, 
yellowish brown hairs, rarely pilose or glabres- 
cent, eglandular. Corolla tube 17-22 mm long, 
1.5-2 times as long as calyx; limb 16-30 mm in 
diameter. 

DISTRIBUTION Brazil (Distrito Federal, Goias, 

Minas Gerais, Parana, Rio de Janeiro, Sao Paulo). 

SPECIMENS EXAMINED BRAZIL. Distrito 



Federal: gallery forest ca. 10 km NW of Planal- 
tina, 950 m, 27 Feb. 1966, Irwin et al. 13194 (us); 
gallery forest ca. 10 km E of Lagoa Paranoa, near 
DF-6, 1000 m, 25 Feb. 1970, Irwin et al. 26556 
(wis); *Brasilia; Bacia do Rio Sao Bartolomeu, 
Corrego Forquilha, Heringer et al. 6148 (F). 
Goias: *25 km NW of Brasilia, Irwin et al. 15799 
(F). Minas Gerais: Mun. Santo Luzia, Fazenda 
da Chicaca, 1 100 m, 20 Nov. 1945, Assis 102 (GH, 
R, us), Assis 137 (GH), Assis 142 (GH), 13 Dec. 
1945, Assis 181 (GH, uc, us); Lagoa Santa, Oct.- 
Nov. 1863, Engler 1006 (c), 14 Jan. 1864, Engler 
1009 (c), 30 Oct. 1863, Engler sp. nov. (c), 25 
Mar. 1933, Mello Barreto 7786 (F), Warming 103 
(NY, s); Serra do Cipo, km 120, 2 Sep. 1933, Mel- 
lo Barreto 7788; Mun. Belo Horizonte, Morro do 
Candido, Jan. 1934, Atamp. 6560 (R); Matta da 
Caixa d'Areia, 15 Dec. 1918, Gehrt 3276 (SP); 



PLOWMAN: REVISION OF BRUNFELSIA 



63 




FIG. 24. Brunfehia brasiliensis subsp. brasiliensis (originally as Franciscea confertiflora). From Pohl, J. E. 
Plantarum Brasiliae Icones et Descriptiones (1826). 



64 



FIELDIANA: BOTANY 



estrada para Lagoa Santa, 15 Jan. 1951, Joly 1133 
(SP); Villa Paraiso, 20 Nov. 1922, Mello Barreto 
7789 (F); Barreiro, 21 Jan. 1933, Mello Barreto 
7791 (F); Serra do Taquaril, 18 Jan. 1933, Mello 
Barreto 7791 (F), 29 Mar. 1933, Mello Barreto 
7793 (F); Jardim Botanico, 17 Nov. 1933, Mello 
Barreto 7794 (R); Serra da Mutuca, 27 Jan. 1945, 
Moreira 5775 (GH); Fazenda da Baleia, 4 Mar. 
1940, Oliveira 21 (us); Morro das Pedras, 3 km 
S of Belo Horizonte, 1000 m, 12 Feb. 1945, Wil- 
liams & Assis (GH); near Lagoa Pampulha, 1000 
m, 8 Mar. 1945, Williams & Assis 6088 (F, GH, 
MO, NY); Mun. Carandai, Carandai, km 416, 25 
Nov. 1946, Duarte 631 (GH, RB); Hermilo Alves, 
10 Jan. 1965, Duarte 8741 (GH, RB); Mun. Ouro 
Preto, Itacolomi, 1400 m, 31 Dec. 1950, Macedo 
2829 (MO, NY, s); Mun. Araxa, Barreiro, 10 Feb. 
1951, Macedo 3131 (s, SP, us); Mun. Santa Bar- 
bara, Serra do Cara9a, 14 Apr. 1933, Mello Bar- 
reto 7784 (F); Mun. Tombos, Fazenda das Antil- 
has, 21 Jan. 1936, Mello Barreto 7767 (F, R); 
Mun. Diamantina, Serra dos Crystais, 6 Nov. 
1937, Mello Barreto 9603 (F, R); Po de Araca, 17 
Nov. 1937, Mello Barreto 9833 (F, R); Mun. Ja- 
boticatubas, Lagoa de Dona Ignacia, 6 Jan. 1940, 
Mello Barreto 10530 (R); Mun. 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 



65 



42 (F, c, P). Novo Friburgo, 9 Dec. 1918, Curran 
619 (GH, s, SP), Curran 623 (F, GH); Morro Pos- 
sole, Sitio Dr. Dungs, 1300 m, 28 Jan. 1968, Pabst 
9115 (HB), 23 Dec. 1887, Glaziou 17170 (BR, c, 
LE, P, us); Petropolis, 1883, Glaziou 14171 (BR, c, 
c); pres de la roche de Juraracacu, 25 Mar. 1879, 
Glaziou 11394 (c, P); Carangola, Sep. 1943, Goes 
& Constantino 543 (RB); Retire, Nov. 1943, Goes 
& Constantino 710 (RB); Caetetu, Dec. 1943, 
Goes & Constantino 883 (RB); Estrada de Con- 
torno, 650 m, 2 Feb. 1968, Sucre 2360, Braga 203 
(GH, RB); Mata do Judeo, 700 m, 7 Dec. 1968, 
Sucre 4213, Braga 1169 (RB), 1858-1860, Wawra 
& Maly 426 (LE, w); Terexopolis, Parque Nacional 
da Serra dos Orgaos, 16 Dec. 1942, Duarte de 
Barros 1195 (RB); Posso, Morro das Antenas de 
Televisao, 12 Feb. 1968, Sucre 2339, Braga 182 
(GH, RB); Serra dos Orgaos, 3000 ft, 7 Jan. 1837, 
Gardner 564 (BM, CGE, G, GH, K, NY, us, w), 1885, 
Glaziou 15132 (c, G, K, LE), May 1839, Guillemin 
954 (P); Imbuhy Lane, Jan. 1838, Miers 4495 (us), 
900 m, Dec. 1952, Vidal 11-5784 (R); Santa Ana, 
1822-1823, Beyrich s.n. (P, s); Montagnes pres 
Sumidoro, Laland s.n. (P); Cantagallo, Peckolt 
181 (w); ad confines Rio de Janeiro-Minas Ger- 
ais, in graminosis partis superioris montis Itatiaia, 
2000-2500 m, Sep. 1901, Wettstein & Schiffner 
s.n. (w); without locality, 1834, Gaudichaud 442 
(G, P), Raben 703 (BR), 1840, Regnell 173 (s), 
1832, Riedel s.n. (GH, GOET, L, LE, NY, P, s, us, w), 
St. Hilaire A-746 (P), 1838-1842, U.S. South Pa- 
cific Exploring Expedition s.n. (NY), Oct. 1916, 
Vincent s.n. (L), Widgren 486 (s); *Morro Quei- 
mado, Almeida 1286 (F); *Floresta da Tijuca; Ped- 
ra do Archer, Angeli 250 (F); *Morro Queimado, 
Brade 18785 (F); *Rio de Janeiro, Pedra da Gav- 
ea, base da Chamine, Carauta 2420 (F); *Gua- 
nabara; Pedra da Gavea, Mesa, Carauta and de 
Oliveira 1523 (F); *Guanabara; Tijuca, Castellan- 
os 23973 (F); *Municipio de Macae, Pico de Fra- 
de, Farney et al. 584 (F); *Municipio de Macae, 
Pico de Frade, Farney et al. 597 (F, RB); *Rio de 
Janeiro; Alto de Boa Vista, Estrada da Vista Chi- 
nesa, mata atras do Decam, Landrum et al. 4150 
(F); *Tijuca, Bom Retire, Pabst 7411 (F); *Estrada 
da Vista Chinesa, Strang 172 (F); *Pedra da Gav- 
ea, Sucre 7446 (F); *Morro Queimado, Sucre 
8002 (F); *Serra da Carioca, no topo do Morro 
Queimado, Vianna et al. 1574 (F). Sao Paulo: 
Jundiahy, region of Morro da Mursa, 1000 m, 4 
Apr. 1915, Brade 7020 (SP); Sao Paulo, Vila Ema, 
Dec. 1932, Brade 12119 (R), Jan. 1952, Brade 
21218 (RB); Campos do Jordao, 10 Sep. 1937, 
Campos Porto 3381 (R), Campos Porto s.n. (GH, 



RB), 22 Oct. 1938, Hashimoto 1 (RB), 27 Jul. 1898, 
Lofgren 3986 (SP); Atibaia, Nov. 1931, Constan- 
tino s.n. (RB); Guaratingueta, 27 Jan. 1920, Gehrt 
3667 (SP); Mogi das Cruzes, 18 Apr. 1921, Hoeh- 
ne 5481 (SP, us), Oct. 1833, Riedel 1466 (G, LE, 
NY, us); Sao Jose dos Campos, 17 Dec. 1908, Lof- 
gren 86 (RB, s); ca. 7.7 km SW em linha reta da 
pra9a principal de Sao Jose dos Campos, 300 m, 
NW da Via Dutra, 600 m, 25 Oct. 1961, Mimura 
53 (us); Ipiranga, Dec. 1912, Luderwaldt 1086 
(RB, SP), Jun. 1907, Usteri 23/306 (NY); Taubate, 
Lund 754, Lund 756 (c); inter Lorena et Taubate, 
Apr. 1833, Riedel 1465 (K, LE, NY); Serra da Bo- 
caina, Jan. 1925, Lutz s.n. (R); in nemorosis editis 
retro Bananal in via publica versus Sao Paulo, 
Dec. 1817, Martius s.n. (M); Morro Formozo, 
Dec. 1817, Martius s.n. (M); Serra da Mantiqueira, 
Fazenda de Cruzeiro do Dr. Major Novaes, 3-12 
Jan. 1884, Saldanha 8587 (R); *Municipio Sao 
Jose de Barreiro; Parque Nacional da Serra da Bo- 
caine, Fazenda das Garrafas, Lima, H.C. de, & 
Ramamoorthy 1224 (F); *Municipio Registro; 
Rod. Registro-Sete B arras, Hatschbach & Gui- 
mardes 45004 (F); *Serra da Bocaina, W. Hoehne 
6154 (F), *W. Hoehne 6155 (F); *Salesopolis; Es- 
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 



66 



FIELDIANA: BOTANY 



calyx or up to H/2 times as long; limb 25-32 mm 
in diameter. 

DISTRIBUTION Brazil (Minas Gerais, Parana, 
Rio de Janeiro, Santa Catarina, Sao Paulo). 

SPECIMENS EXAMINED BRAZIL. Minas Ger- 
ais: Mun. Diamantina, Guinda, 5 Nov. 1937, Mel- 
lo Barreto 9479 (F, R). Parana: Ipiranga, Monte 
Alegre, 1100 m, Dusen 3388 (R); Bauhado, 13 
Dec. 1911, Dusen 13562 (GH, NY, s); Guaratuba, 
21 Feb. 1952, Reitz 4375 (us); *Palmeira, Hatsch- 
bach 11878 (MBM); *Barra Rio Papagaios (Balsa 
Nova), Hatschbach 18747 (MBM); *Municipio 
Guaratuba; Rio Sai, Hatschbach 19447 (MBM); 
*Municipio Jaquariaiva; Fazenda Chapada S. An- 
tonio, Hatschbach 20997 (GH); *Municipio Pir- 
quara; Serra do Emboque, Hatschbach 26532 
(MBM); *Municipio Pirai do Sul; Joaquim Murtin- 
ho, Hatschbach 39209 (GH, MBM); *Municipio 
Palmeira, Rio Tibagi, Kummrow 970 (GH); *Bran- 
dalize, Lindeman and Haas 1174 (u). Rio de Ja- 
neiro: Alto Macahe de Nova Friburgo, 22 Jan. 
1874, Glaziou 6908 (BR, c, s), Glaziou 13478 (c, 

F, LE); 1882, Glaziou 14172 (c, G, LE); Petropolis 
a Itamaraty, 25 Mar. 1879, Glaziou 11395 (BR, c, 

G, LE, R); Serra dos Orgaos, 18 Nov. 1887, Gla- 
ziou 17169 (BR, c, p, R), 2,200 m, Jun. 1915, Liitz- 
elburg 6187 (M); Morro Assu, 2400 m, Jan. 1916, 
Lutzelburg 6559 (M, NY), Feb. 1838, Miers s.n. 
(BM); Serra do Itatiaia: 1900 m, Sep. 1913, Brade 
& Tamandare 6576 (SP), 1915, Campos Porto 173 
(R, RB), 2100 m, Jun. 1902, Dusen 569 (R), 18 
Oct. 1903, Dusen 2023 (K, R, s, us), 1750 m, 28 
Oct. 1927, Ginzberger 93 (F), Moreira s.n. (R); 
Parque Nacional de Itatiaia, km 15, 17 Aug. 1948, 
Occhioni 1131 (GH, RB); prope Registro, 1800 m, 
14 Nov. 1954, Pabst & Brade 10283 (us); Ma- 
cieras, 1900 m, Sep. 1934, Brade 14074 (B, R, RB), 
1800-2000 m, 23 Oct. 1927, Zerny s.n. (w); entre 
Maromba e Macieiras, 1820 m, 29 May 1969, 
Plowman 2891, Sucre 5191 (ECON, GH, MICH, PH, 
RB, uc, YEN), 1,700 m, Plowman 2906, Sucre 5206 
(GH, MICH, PH, RB), 1200-1900 m, 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. & 
Downs. 

Sprengel described G. brasiliensis from a spec- 
imen collected by Sellow in Brazil, and his de- 
scription, albeit brief, more or less agrees with the 
present concept of the species. There is, however, 
one point of difference in Sprengel's description, 
where he writes "pedunculis axillaribus solitar- 
iis." Usually this species has terminal, branched 
inflorescences, although there may be subterminal 
axes produced in the uppermost leaf axils. 

Unfortunately, Sprengel's herbarium was dis- 
mantled and sold after his death by his son. Some 
of the specimens survived and were eventually 
deposited at the Berlin herbarium, where they 
were destroyed in the Second World War (Stafleu, 
1967; Stafleu & Cowan, 1985). Sellow apparently 
made several collections of this species with nu- 
merous duplicates, none of which bears collection 
numbers or localities. It is thus impossible to de- 
termine which specimens might be isotypes of the 
original specimen that Sprengel described. As a 
representative collection, I selected another spec- 
imen of Sellow that was sent to Munich from Ber- 
lin in exchange [this is not to be taken as neotyp- 
ification of the name (see footnote 11) Eds.}. 

In 1826 Pohl published seven new species of 
his new genus Franciscea, including F. ramosis- 
sima, F. acuminata, F. confertiflora, and F. di- 
varicata. He stated at that time that these "spe- 
cies" were all closely related but believed that 
their differences were great enough for them to 
be considered separate entities. Pohl's specific 
concepts were generally followed by Bentham, 
who in 1846 transferred all of Pohl's species of 



PLOWMAN: REVISION OF BRUNFELSIA 



67 



Franciscea to the genus Brunfelsia, omitting only 
F. divaricata, which he placed in synonymy with 
B. confertiflora. According to Bentham, B. con- 
fertiflora differed from B. ramosissima in having 
a many-flowered, crowded inflorescence, an in- 
flated calyx, and the corolla tube only W2 times 
as long as the calyx. 

Brunfelsia acuminata as maintained by Ben- 
tham was merely a glabrous form of B. ramosis- 
sima. The type specimen from which Pohl's orig- 
inal drawing was made is preserved in Brussels 
(BR). A specimen at Vienna (w), collected and ap- 
parently annotated by Schott as F. acuminata, has 
a well-developed indument and does not conform 
to the description of this species. As mentioned 
above in the discussion of B. martiana, there has 
been considerable confusion of collection data 
and numbers in the Brazilian collections of Pohl 
and Schott. For example, the same collection in 
different herbaria may bear the same number or 
numbers with either or both names. For this rea- 
son the numbers of these two collectors should be 
used with some caution. 

Schmidt (1864), in his account of Brunfelsia for 
Flora Brasiliensis, treated this group of "species" 
as a single variable species, B. ramosissima, with 
three varieties. Variety laxiflora included B. ra- 
mosissima and B. acuminata of Bentham, and va- 
riety confertiflora conformed more or less with 
his B. confertiflora. Schmidt described a third va- 
riety, parcifolia, for a form with small, coriaceous 
leaves. Annotations in Schmidt's handwriting on 
the type and other specimens read "parvifolia," 
indicating either a typographical error in publi- 
cation or that he changed his mind. The former is 
more likely since these plants have quite small, 
but not few, leaves. This form is now considered 
only a minor variant. 

Brunfelsia brasiliensis is here treated as a sin- 
gle polymorphic species, widely distributed in 
southern Brazil, with one subspecies, macrocalyx, 
occurring at higher elevations from the Serra do 
Mar and Serra da Mantiqueira southward in the 
mountains to Santa Catarina. Brunfelsia bras- 
iliensis exhibits a wide range of morphological 
variation throughout its area of distribution, which 
consists mostly of mountainous, ecologically di- 
versified terrain. In this region, primarily in the 
states of Minas Gerais, Rio de Janeiro, and Sao 
Paulo, there exist ample opportunities for geo- 
graphical and ecological isolation of populations 
with subsequent morphological differentiation. 

Environmental factors also play an important 
part in the phenotypic variability seen in this spe- 



cies, and different forms may be interpreted as 
ecotypes. For example, in exposed, well-drained 
places such as the crest of a peak or rocky out- 
crop, the plants are frequently low in stature with 
horizontal branches, small, often coriaceous 
leaves, and fewer and smaller flowers. Lower 
down the same slope, in shaded, more protected, 
and probably more fertile places, the plants may 
attain a height of up to 3 m with well-developed 
inflorescences and larger flowers and leaves. 

The great morphological variation in B. bras- 
iliensis occurs chiefly in the presence, type, and 
location of indumentum, in the size and form of 
the inflorescence, and to a lesser extent in the size 
and form of the flowers and leaves. 

An important diagnostic feature of this species 
is the presence of a persistent peduncle, a short 
stalk that articulates with the pedicel of each flow- 
er. This structure is scarcely, if ever, developed in 
any other species of Brunfelsia. Some populations 
of B. uniflora, however, may have a similar stalk 
in its solitary flowers. This "peduncle" is perhaps 
more accurately described as an ultimate branch 
of the inflorescence that immediately precedes a 
flower. It commonly bears 1-3 small bracts near 
the base, the axillary buds of which may produce 
additional flowers, thereby proliferating the inflo- 
rescence. The peduncle may be shorter or longer 
than the pedicel or, more rarely, obsolete. The dis- 
tal end is dilated into a short, flat torus that be- 
comes conspicuous when the flower falls. 

The inflorescence arises at the tips of the bran- 
chlets and is usually much-branched and many- 
flowered. It may be strictly terminal and compact 
to lax and open with several subterminal branches 
also developing into flowering axes. Alternatively, 
the flowers may be reduced to a few, often sub- 
sessile ones and rarely to a single flower in sub- 
species macrocalyx. The calyx in B. brasiliensis 
may be tubular to nearly campanulate in form and 
varies greatly in size. The corolla tube may be 
scarcely exserted from, or up to twice as long as, 
the calyx. 

The presence of a well-developed indument, 
particularly in the inflorescence, is also diagnostic 
in B. brasiliensis. The trichomes are nonglandular, 
uniseriate, long or short, and straight or curved. 
In dried material, they are frequently yellowish or 
yellowish brown. Often the indument may be 
densely villous to tomentose. More rarely it ap- 
pears quite sparse and pilose, or the plants may 
be glabrescent. The upper surface of the leaves is 
frequently glabrescent, with short, subscabrous 
hairs that break off near the base. The lower sur- 



68 



FIELDIANA: BOTANY 



face of the leaves is usually pubescent, especially 
at the nerves and midrib. 

Brunfelsia brasiliensis is easily distinguished 
from most other species of the genus by the reg- 
ularly oblong-lanceolate leaves, the distinctive 
yellowish brown pubescence, and the presence of 
the short peduncle articulating with the pedicel. 
Rare glabrous forms of B. brasiliensis (= B. ac- 
uminata (Pohl) Benth.) resemble B. bonodora but 
are distinguished by their stalked pedicels. Brun- 
felsia brasiliensis appears to be most closely re- 
lated to the B. bonodora-B. grandiflora complex 
by the nature of the more or less branched inflo- 
rescence and floral and fruit morphology. 

Brunfelsia brasiliensis subsp. brasiliensis is 
distributed in southeastern Brazil from Minas 
Gerais south to Sao Paulo, between 700 and 1400 
m elevation. The species shows a fairly wide eco- 
logical tolerance but prefers well-drained sites. It 
is frequently encountered in the mares de morros, 
"the seas of hills," which occur in southeastern 
Brazil, particularly in Minas Gerais. Here the 
plant grows on granitic peaks and rocky outcrops, 
often in dry, exposed habitats. It seems to occur 
less commonly in the humid forests on the slopes 
of the Serra do Mar. It also grows in association 
with the campos, or open grassland communities, 
and in capaes, small patches of woodland found 
scattered in the campos. Brunfelsia brasiliensis is 
also encountered in capoeiras, a general term for 
cut-over, secondary woods, and is one of the few 
brunfelsias that survive cutting of the primary for- 
ests. Around Brasilia, it has been reported from 
gallery forests. Brunfelsia brasiliensis is a freely 
flowering plant with the flowers appearing pri- 
marily from September to March. The fruits have 
been collected from October to March. This spe- 
cies has been cultivated in European conservato- 
ries since its discovery, although it is much rarer 
in cultivation than other species. To my knowl- 
edge, it is not planted in the American tropics 
outside of Brazil. Peckolt (1909) says that it is a 
common garden plant in that country. Brunfelsia 
brasiliensis is known as manacd, manacd da ser- 
ra, erva de macaco, and manacd assu (Peckolt, 
1909). From herbarium labels (Occhioni s.n., V. 
Assis 102, 142) we know that it is considered poi- 
sonous. 

At higher elevations in the mountains of Rio de 
Janeiro, Sao Paulo, and south to Santa Catarina, 
there occurs a distinct subspecies of B. bras- 
iliensis. This was first described by Dusen in 1905 
as B. hopeana var. macrocalyx from a specimen 
collected on Mount Itatiaia. This type superficial- 



ly resembles B. uniflora (=B. hopeana) in having 
the inflorescence reduced to a single flower, but 
this is somewhat exceptional for the subspecies as 
now circumscribed. 

Brunfelsia brasiliensis subsp. macrocalyx dif- 
fers from B. brasiliensis subsp. brasiliensis in 
having fewer flowers per inflorescence (one to 
eight), a very short peduncle in which the flowers 
may appear sessile, a longer pedicel (5-15 mm) 
that always exceeds the peduncle, and a usually 
longer calyx (15-22 mm). The corolla tube of 
subspecies macrocalyx is short relative to the ca- 
lyx and is just barely exserted. The indumentum 
is frequently less well-developed, especially in the 
leaves, which may be glabrescent and bright 
green. In addition, the trichomes may be glandu- 
lar, especially in the calyx and pedicels, a char- 
acter not normally encountered in subspecies 
brasiliensis. 

Morphologically intermediate populations are 
known to exist between subspecies brasiliensis 
and subspecies macrocalyx. In collections from 
the Serra dos Orgaos and Alto Macahe in the state 
of Rio de Janeiro, subspecies brasiliensis shows 
a tendency to develop a large calyx (to 17 mm) 
similar to that of subspecies macrocalyx. How- 
ever, because these plants are, in general, larger 
in all parts, they may be considered exceptionally 
vigorous individuals perhaps growing in a partic- 
ularly favorable place (cf. Glaziou 6908, 13478, 
17169). Other collections (cf. Occhioni 1131) 
from Mount Itatiaia are referable to subspecies 
macrocalyx, although they exhibit a multiflorous 
inflorescence. These plants may indeed suggest 
putative hybridization between the two subspe- 
cies; however, this cannot be proved at this stage 
of knowledge of the group. Several collections 
from the states of Parana and Santa Catarina have 
been referred to B. brasiliensis subsp. macrocalyx 
(e.g., Dusen 13562, Reitz 3985). These popula- 
tions are somewhat heterogeneous and in some 
characters resemble B. cuneifolia (see discussion 
under that species), such as the indument and the 
reduction in number of flowers, often to one. Fur- 
thermore, the calyx tends to be smaller than in 
more northern populations of this subspecies, 
somewhat resembling that of subspecies bras- 
iliensis. 

Because of the complex pattern of variation 
that is encountered in this region, I believe this 
problem cannot be resolved completely without 
additional materials and a knowledge of the plants 
in the field. 

Brunfelsia brasiliensis subsp. macrocalyx in the 



PLOWMAN: REVISION OF BRUNFELSIA 



69 



northern part of its range occurs at altitudes be- 
tween 1650 and 2400 m on relatively few moun- 
tain peaks and high campos: Mount Itatiaia, Alto 
Macahe, and Serra dos Orgaos in Rio de Janeiro; 
and Alto da Serra, Serra de Bocaina, and Campos 
do Jordao in Saao Paulo. In Sao Paulo (Campo 
Grande) and southward, subspecies macrocalyx 
occurs at somewhat lower elevations (800-900 
m). 

Ecologically, it grows in wet, montane cloud 
forests in association with Podocarpus and also 
occurs in moist, high-elevation campos. In the 
Parque Nacional de Itatiaia, plants have been fre- 
quently collected between Maromba and Maci- 
eiras at 1600-1900 m elevation and are repre- 
sented by two distinct forms. One of these (cf. 
Plowman 2891, Sucre 5191) is sparsely branched 
with dark leaves and pubescent calyx with erect 
lobes. The other (cf. Plowman 2906, Sucre 5206), 
which grows sympatrically a few hundred yards 
away, is profusely branched with bright green, 
glabrous leaves and a glabrous calyx with strong- 
ly recurved lobes. Though strikingly different to 
the casual observer, these plants must be regarded 
as minor phenotypic variants of a single subspe- 
cies. Analysis in the field of situations such as this 
gives a measure of the kind of variability that is 
present in a population and a better basis for un- 
derstanding comparable problems where field 
studies are not possible. 

Efforts were made to rear these plants in the 
greenhouse from seed to study the nature of the 
characters and their inheritance. However, the 
seedlings proved to be exceedingly sensitive to 
fluctuations in temperature and humidity and soon 
died. 

11. Brunfelsia chiricaspi Plowman, Bot. Mus. 
Leafl. 23(6): 255, t. 17. 1973. Type: CO- 
LOMBIA. Putumayo: Umbria, 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 
23. 

SPECIMENS EXAMINED COLOMBIA. Putu- 
mayo: Orito, near Texaco drilling site, 11 Feb. 
1972, Kennedy 1386 (GH); alrededores de Puerto 
Limon, 300-400 m, 15 Feb. 1953, Mora 1040 
(COL); El Whiskey at Finca "Santa Marta," 13 km 
S. of Umbria near road, 400-500 m, 27 Nov. 
1968, Plowman 2080 (ECON, GH); forest about 2 
km SW of San Pedro, N of Puerto Asis, 400-500 
m, 28 Nov. 1968, Plowman 2081 (ECON, GH); Rio 
Guamues, San Antonio, 5 Sep. 1966, Pinkley 420 
(ECON); Santa Rosa, 28 Nov. 1966, Pinkley 563 
(ECON); trail between Santa Rosa and road to Hor- 
miga, 2 Dec. 1968, Plowman 2092 (ECON, GH); 
*Rio Putumayo, Puerto Ospina, Schultes 3422 
(GH). ECUADOR. Napo: * Along Rio Aguarico 
upstream from Santa Cecilia, Duke 16047 (MO); 
*Coca (Puerto Francisco de Orellana), S side of 
Rio Napo, Marling & Andersson 11771 (GB); 
*Coca (Puerto Francisco de Orellana), Marling & 
Andersson 11804 (GB, GH); *Hacienda Cotapino 



70 



FIELDIANA: BOTANY 



BRUNFELSIA chiricaspi Plowman 




FIG. 25. Brunfelsia chiricaspi. Reproduced courtesy of the Botanical Museum of Harvard University. 



(Conception), Marling et al. 7021 (GB, GH); *Path 
Cotapino (Conception), Rio Bueno, Marling et al. 
7151 (GB, GH); *Coca de Orellana, lower Rio Pay- 
amino, Marling et al. 7772 (GB, GH); *Hacienda 
San Carlos at Rio Napo, ca. 15 km E of Coca 



(Puerto Francisco de Orellana), Lugo 2724 (GB, 
GH); *Tierra Colorada, ca. 3 km N of Coca (Puerto 
Francisco de Orellana), Lugo 2942 (GB, GH); *Ca- 
non de los Monos, road Coca (Puerto Francisco 
de Orellana), Lago Agrio, ca. 12 km N of Coca, 



PLOWMAN: REVISION OF BRUNFELSIA 



71 



Lugo 2981 (GB, GH); *road from Lago Agrio-E 1 
Chaco, Lugo 3476 (GB, GH); *7 km N of Coca; 
Estacion Experimental de INIAP-Payamino, Zar- 
uma et al. 56 (F). 

Brunfelsia chiricaspi was originally known 
only from a few collections from a small area of 
southwestern Colombia on the eastern flank of the 
Cordillera Occidental. [Plowman predicted that 
the distribution would prove to be much greater, 
and this species is now also known from many 
recent collections in adjacent Ecuador (see 
above). Eds.] It grows as an understory shrub in 
humid, primary forests at elevations of 325-500 
m. The fruit is as yet imperfectly known because 
of a lack of material. 

Brunfelsia chiricaspi is related to B. grandiflora, 
which is widespread in the western Amazon, and 
to B. mire of Bolivia and adjacent Peru and Brazil. 
Brunfelsia chiricaspi differs from B. grandiflora in 
having larger leaves, longer pedicels (6-13 vs. 2- 
6 mm) and usually fewer flowers. It differs from 
B. mire in the basally obtuse, elliptic to lanceolate 
leaves; the few-flowered cyme; and smaller flowers 
(corolla tube 22-25 vs. 25-38 mm long). 

Known generally as chiricaspi in the Colom- 
bian Putumayo, this species is used as a strong 
medicine and narcotic by the Kofan Indians (see 
Plowman, 1977). 

12. Brunfelsia cuneifolia J. A. Schmidt in Mart., 
Fl. Bras. 8(1): 259. 1864. Type: BRAZIL. In 
Brasilia australiore, Sellow 4016 (lectotype, 
designated here, destroyed at Berlin; photo- 
graph of lectotype, F 621824, NY, us; isolec- 
totypes, F, M). 13 Riedel 1467, also cited by 
Schmidt, is excluded as a type. It belongs to 
B. obovata Benth. var. obovata. Figure 26. 

Shrub to about 1 m tall, much branched. 
Branches terete, naked. Bark grayish to yellow- 
ish brown. Branchlets short, leafy, sparsely to 
densely villous, rarely glabrescent, hairs often yel- 
lowish to reddish brown, sometimes glandular. 



13 It is clear that Plowman was intending to lectotypify 
this species using the destroyed B sheet as good photo- 
graphs of it exist. He appears to specifically not consider 
the unnumbered Sellow sheets at F and M as good can- 
didates for lectotypes (in the strict sense). Eds. 



Leaves scattered or crowded near the tips of the 
branchlets, blade 3.5-10 cm long, 1.8-3 cm wide, 
oblong-obovate to elliptic-lanceolate, cuneate to 
blunt at apex, rarely acuminate, narrowed to 
broadly cuneate at base, the upper surface gla- 
brescent or with long, scattered hairs, the lower 
surface velutinous, often densely so at midrib and 
nerves, rarely glandular or glabrescent, somewhat 
shiny above, dark green, pale green beneath, firm- 
ly membranaceous to subcoriaceous, lateral 
nerves 4-7; petiole 2-4 mm long, velutinous or 
glandular. Inflorescence terminal on tip of 
branchlets, sessile, with 1-3 flowers. Bracts 2-8 
mm long, small, lanceolate, glandular, ciliolate, 
caducous. Flowers violet fading to white with 
age. Pedicel 3-5 mm long, slender, thickened at 
apex, velutinous or glandular pubescent. Calyx 
12-18 mm long, 4-6 mm in diameter, tubular, in- 
flated, 5-angled, drying plicate, pubescent with 
short glandular hairs, especially near base, or vil- 
lous, pale green, membranaceous, teeth 3-7 mm 
long, ovate-lanceolate, acute to acuminate at apex. 
Corolla tube 18-28 mm long, 1.5-2 mm in di- 
ameter, \Vi times as long as calyx, pentagonal at 
base, pilose or minutely glandular in upper part; 
limb 25-37 mm in diameter, spreading, lobes 10- 
15 mm long, undulate, broadly rounded. Stamens 
inserted in upper part of tube; upper pair of fila- 
ments 3-4 mm long, suberect, exserted slightly 
from mouth; lower pair 2-3 mm long, included, 
curved at apex; anthers 1.5 mm long, orbicular- 
reniform. Ovary 2 mm long, conical-ovoid, style 
about 23 mm long, filamentous; stigma 1 mm 
long, briefly bifid. Fruit and seed unknown. 

DISTRIBUTION Brazil (Parana, Rio Grande do 
Sul, Santa Catarina, Sao Paulo). See Figure 28. 

SPECIMENS EXAMINED BRAZIL. Parana: 
Mun. Mangeirinha, Lageado San Antonio, 20 Oct. 
1966, Hatschbach 15159 (US); *Municipio de 
Guarapuara; Guarapuara, Frenzel 666 (MBM); *In- 
acio Martins, Hatschbach & Guimardes 30302 
(MBM); *Municipio Campo Largo; Serra do Pu- 
runa, Kummrow 2432 (F). Rio Grande do Sul: 
Farroupilha, 15 Oct. 1957, Camargo 2123 (B), 3 
Oct. 1957, Camargo s.n. (s); *Municipio Cacequi, 
Dobereiner & Tokarnia 732 (RB); *Esmeralda; 
Estacion Ecologica de Aracuri, Jarenkow 132 (F). 
Santa Catarina: Mun. Lajes, Passo do Socorro, 



FlG. 26. Brunfelsia cuneifolia. 



72 



FIELDIANA: BOTANY 




I 

\ 



N? 1995963 

FIELD MUSEUM 

OF 
NATURAL HISTORY 



~t 



PREFEITURA MUNICIPAL DE CURITIBA 
MUSEU BOTANICO MUNICIPAL 



i io li -a ZT- A- 



icuaa Sta. Clara d 




50573 



Arbuato 2 B| flor lllas. Interior aata te galria. 



PMC Off 004 



PLOWMAN: REVISION OF BRUNFELSIA 



73 



900 m, 31 Oct. 1963, Klein 4343 (us); Mun. Novo 
Horizonte, Lauro Miiller, 400 m, 24 Oct. 1958, 
Reitz & Klein 4076 (NY, us); Campos Novos, 12 
Sep. 1963, Reitz & Klein 16158 (GH, us); Mun. 
Campos Alegre, lower fazenda of Ernesto Schei- 
de, Campo Alegre, 900 m, 11 Dec. 1956, Smith 
& Klein 8567 (us); Mun. Chapeco, Guatambu, ca. 
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 
isolectotypes. 

Known from only a few collections, the avail- 
able material of B. cuneifolia shows a consider- 
able amount of variation in the form of the leaves 
and type and amount of indument. Brunfelsia cu- 
neifolia is distinguished from other species of 
southern Brazil by the apically cuneate leaves and 
a five-angled tubular calyx that appears plicate 
when pressed and dried. The leaves and calyces 
are always pubescent with either of two types of 
trichomes present. The type collection and mate- 
rial from Rio Grande do Sul have short glandular 
hairs. Most specimens from Santa Catarina have 
a dense villous to velutinous indument consisting 
of longer, nonglandular hairs. The form of the 
leaves and calyx associated with both types is 
fairly uniform. 

Brunfelsia cuneifolia is not easily distinguished 
from other species of southernmost Brazil. It su- 
perficially resembles B. pilosa, which is sympatric 
with B. cuneifolia. Brunfelsia pilosa, however, 
usually has a more pronounced acumen in the leaf 
and a terete, tubular calyx that is purple and bears 
only a few, long pilose hairs. In addition, the 



flowers of B. pilosa are somewhat larger than 
those of B. cuneifolia. 

The villous pubescence of B. cuneifolia is very 
reminiscent of the yellowish indument character- 
istic of B. brasiliensis. Populations interpreted 
here as B. brasiliensis subsp. macrocalyx extend 
as far south as Santa Catarina and overlap with 
the range of B. cuneifolia. Collections of these 
two species in this region are morphologically 
very similar and may be easily confused. Brun- 
felsia brasiliensis subsp. macrocalyx may be sep- 
arated by the presence of the persistent peduncle, 
which articulates with the pedicel of the flowers, 
and by the terete, campanulate or tubular-campan- 
ulate calyces. This species usually has longer ped- 
icels and shorter calyx teeth than B. cuneifolia. 
Nevertheless, the intermediate nature of these 
populations indicates that some hybridization may 
have occurred between these two species and pos- 
sibly involved B. pilosa as well. 

The true affinities of B. cuneifolia appear to lie 
with B. obovata var. obovata, which occurs far- 
ther north in Sao Paulo and Minas Gerais [but see 
also B. boliviana. Eds.]. This plant resembles B. 
cuneifolia in the tubular pentagonal calyx, which 
also nearly equals the corolla tube in length, and 
in the obovate leaves. In both species the upper- 
most pair of anthers may be briefly exserted from 
the corolla tube, a character that is unusual in sec- 
tion Franciscea. Brunfelsia cuneifolia differs in 
having the leaves cuneate at the apex, fewer flow- 
ers per inflorescence, and a pubescent calyx that 
is much more slender than in B. obovata (4-6 vs. 
6-10 mm). These two species also differ striking- 
ly in their ecological preference. Brunfelsia obov- 
ata is a plant of marshes and flooded areas, 
whereas B. cuneifolia occurs in uplands. 

Brunfelsia cuneifolia is a relatively low shrub 
that is found in habitats apparently similar to 
those of B. pilosa. It is specifically reported from 
Araucaria forests (arau-carieto) and also from 
gallery forest and capoeiras, or secondary forest, 
and between 400 and 900 m altitude. It flowers in 
October and November and is known by the com- 
mon name flor de Trovoada, or "thunderstorm 
flower." 

13. Brunfelsia dwyeri D'Arcy, Ann. Missouri 
Bot. Gard. 57:259. 1971 ("1970"). Type: 
PANAMA. Panama: Cerro Jefe in "Cleusia" 



FIG. 27. Brunfelsia dwyeri. 



74 



FIELDIANA: BOTANY 




1709790 



FIELD MUSEUM 

OF 
NATURAL HISTORY 



PLANTS OF PANAMA 

Brunfelsia dwyeri D'Arcy 

det. by R.L. Wilbur, 1971. 

Provincia de Panama': slopes of Cerro Jefe 
beyond Cerro Azul between 4-8 miles in 
mostly heavily wooded slopes. 

Common shrub 2-5 m. tall. Corolla deep 
violet blue becoming paler with age. 



R.L. Wilbur 
R.E. Weaver 



#11358 

25 January 1970 



PLOWMAN: REVISION OF BRUNFELSIA 



15 



(sic) forest, 2700-3000 ft, shrub, flowers pur- 
ple to whitish purple, 27 Jan. 1966, Tyson, 
Dwyer & Blum 3312 (holotype, MO 1820936; 
isotype, MO 1917560). Duke & Dwyer 13933 
(MO) is excluded from the cited paratypes. 
Figure 27. 

Shrub or small tree 2-10 m tall. Trunk to 6- 
8 cm in diameter, branched from near the base. 
Bark gray to grayish brown. Branches terete or 
somewhat angular, naked. Branchlets leafy. 
Leaves with blade 5-11 cm long, 2.5-5 cm wide, 
obovate, elliptic or oblong-lanceolate, acuminate 
at apex, slightly revolute at margin, cuneate or 
narrowed at base, glabrous, shiny and light to 
very dark green above, pale green beneath, cori- 
aceous, lateral nerves 5-7, veinlets reticulate, 
prominulous, especially on lower surface; petiole 
3-10 mm long, becoming transversely corrugate- 
rugose. Flowers solitary, terminal or axillary in 
the upper leaf axils, purple, fading to white with 
age, with irregular white spot at mouth. Peduncle 
1-5 mm long, sometimes obsolete. Bracts 1-3 
mm long, linear to ovate, puberulent, ciliate-mar- 
gined, caducous. Pedicel 5-15 mm long, about 2 
mm in diameter, glabrous. Calyx 12-22 mm long, 
5-10 mm in diameter, tubular to tubular-campan- 
ulate, inflated or not so, glabrous, subcoriaceous, 
reticulately veined, yellowish green or purplish, 
teeth incumbent or erect, triangular to ovate-lan- 
ceolate, at apex acute or blunt; calyx in fruit co- 
riaceous, often becoming dotted with lenticels, 
verrucose near base, splitting completely along 
one or two sinuses. Corolla tube 25-40 mm long, 
2-4 mm in diameter, 2-3 times as long as calyx, 
glabrous; limb 30-80 mm in diameter, spreading, 
lobes 10-20 mm long, broadly rounded, the upper 
lobe slightly larger. Stamens inserted in upper 
part of corolla tube; filaments slender, terete, up- 
per pair about 3 mm long, lower pair about 5 mm 
long; anthers included within corolla tube, 2 mm 
in diameter, orbicular-reniform. Ovary about 2 
mm long, conical, ovoid, weakly 4-angled; style 
filiform; stigma briefly bifid, the lobes subequal, 
blunt. Capsule 1.5-2.5 cm long, 1-1.5 cm in di- 
ameter, globose to ovoid, usually 2-celled, at ma- 
turity often appearing 4-lobed with 4 prominent 
cruciform grooves, pericarp to 4 mm thick, tough 
and leathery, drying hard, surface smooth and 
green at first, becoming rugulose-verrucose, light 
brown. Seeds 3-15, 8-10 mm long, 3-4 mm in 
diameter, oblong-ovoid, prismatic, dark reddish 
brown, reticulate-pitted. Embryo straight, coty- 
ledons ovate. 



DISTRIBUTION Panama, known only from Cer- 
ro Jefe, Provincia de Panama. See Figure 28. 

SPECIMENS EXAMINED PANAMA. Panama: 
Cerro Jefe, beyond Goofy Lake, 16 Nov. 1967, 
Correa & Dressier 471 (MO); along main road 
before turnoff to summit, 6 Jan. 1971, Croat 
13136 (GH, MO); Cerro Jefe, 8 Apr. 1970, D'Arcy, 
Dressier & Correa 3948 (MO), D'Arcy 3959 (MO), 
21 Jan. 1967, Duke 9429 (MO), 2 Apr. 1969, 
Dwyer, Durkee & Castillon 5051 (MO), 12 Mar. 
1968, Dwyer 7280 (GH, MO), ca. 2,900 ft, 29 Jul. 
1967, Dwyer & Ganger 7349 (GH, MO); along road 
from Cerro Jefe to La Eneida (Cerro Azul), Croat 
13072 (GH, MO), 12 Feb. 1971, Escobar 359 (COL, 
ECON, GH), 26 Jul. 1969, Kennedy 316 (ECON), 12 
Apr. 1972, Plowman 3160 (COL, ECON, GH, MICH, 
uc), 9 Jul. 1966, Tyson, Dwyer & Blum 4335 
(MO), 2300-2700 ft, 19 Jan. 1969, Tyson 5315 
(MO), 25 Jan. 1970, Wilbur & Weaver 11358 (GH). 

Brunfelsia dwyeri was discovered and de- 
scribed during studies on the flora of Panama. So 
far as is known, it occurs only on Cerro Jefe in 
central Panama between 700 and 1,000 m eleva- 
tion, an area now famous for its high floristic en- 
demism (Lewis, 1971). The collection Duke & 
Dwyer 13933 (det. W. G. D'Arcy, 1973) from 
Cerro Pilon in Provincia Cocle, which was cited 
in the original description as B. dwyeri, is actually 
Schultesianthus megalandrus (Dunal) Hunz. 

The thick-walled fruit and shiny, coriaceous 
leaves are distinctive features of B. dwyeri. It ap- 
pears to be most closely related to B. macrocarpa 
of coastal Colombia and Ecuador, the flowers of 
which are similarly large and solitary and which 
also possesses a thick-walled fruit. The two spe- 
cies are unique in section Franciscea in possess- 
ing a fleshy pericarp. The resemblance of B. 
dwyeri to B. lactea of Puerto Rico, pointed out by 
D'Arcy (1971), is indeed "superficial" because 
these species are placed in separate sections of the 
genus and have no close affinities with one an- 
other. 

Brunfelsia dwyeri is a beautiful shrub with at- 
tractive, shiny foliage and large, showy flowers. 
It is locally common in the cloud forests on the 
upper slopes of Cerro Jefe, occurring in secondary 
woods with seasonal rainfall. This locality and 
species represent the northernmost extension of 
Brunfelsia sect. Franciscea. As pointed out by 
Lewis (1971), the entire forest of Cerro Jefe and 
surrounding areas is rapidly being destroyed for 
raising poultry and cattle. Brunfelsia dwyeri and 



76 



FIELDIANA: BOTANY 




FIG. 28. 
square). 



Distribution of Brunfelsia cuneifolia (solid triangle), B. dwyeri (solid circle), and B. grandiflora (solid 



many other endemic species of plants are in grave 
danger of extinction. 

This species flowers and fruits intermittently 
throughout the year, although I failed to find flow- 
ers in mid-April. Local people reported that the 
bark and roots of this plant are used for a "rem- 
edy." 

14. Brunfelsia grandiflora D. Don, Edinburgh 
New Philos. J.: 86. Jul. 1829. Type: PERU. 
San Martin: Uchiza, 1798, Tafalla s.n. (ho- 
lotype, BM; isotype, MA). Figure 29. 

Franciscea grandiflora (D. Don) Miers, Ann. Mag. 
Nat. Hist., ser. 2, 5:250. 1850. 

Shrub or small tree 1-6(10) m tall. Trunk to 
7 cm in diameter near base, much branched. Bark 



thin, roughish, light to dark brown. Branches 
slender, ascending or spreading, often subvirgate 
and arching, leafy, glabrous. Branchlets glabrous, 
rarely pubescent, green. Leaves more or less two- 
ranked, scattered along branchlets, blade 6-23 cm 
long, 2-8 cm wide, lanceolate to oblong, often 
arched with ascending laminas, apically long to 
short acuminate, the acumen subfalcate, cuneate 
to narrowed at base, glabrous or sparingly pubes- 
cent beneath at midrib, dull, dark green above, 
paler green beneath, rarely glaucescent or nitid, 
firmly membranaceous to subcoriaceous; petiole 
2-6 mm long, glabrous or minutely pubescent, be- 
coming corky, transversely corrugate with age. 
Inflorescence terminal and subterminal, simple or 
branched, dense or lax, the axis 5-45 mm long. 
Flowers 5-many, showy, scentless, violet fading 
to white with age, with rounded, white ring at 



PLOWMAN: REVISION OF BRUNFELSIA 



77 



mouth. Bracts 1-3 per flower, 1-5(10) mm long, 
lanceolate to ovate, ciliolate, pubescent or glabra- 
te, caducous. Pedicel 2-5(10) mm long, glabrous 
or with few sparse glandular hairs, becoming 
thicker and corky- verrucose in fruit. Calyx 5-13 
mm long, 3-7 mm in diameter, tubular-campanu- 
late, somewhat narrowed toward base, globose to 
obovate in bud, somewhat inflated or not so, gla- 
brous, rarely punctate or sparsely glandular with- 
in, smooth or striate-nerved, light yellow-green to 
gray-green, firmly membranaceous to subcoria- 
ceous, teeth 1-5 mm long, triangular to ovate, 
blunt to short acuminate, erect or incumbent, re- 
curved slightly with age; calyx in fruit persistent, 
coriaceous, becoming corky-verrucose especially 
near base, often splitting on one or more sides. 
Corolla tube 15-40 mm long, 1.5-3 mm in di- 
ameter, 2-5 times as long as calyx, glabrous, pale 
violet to white; limb 20-50 mm in diameter, 
spreading, violet fading to white with age, lobes 
7-15 mm long, subequal or uppermost somewhat 
larger, rounded or emarginate at apex, overlapping 
at sides, abruptly narrowed at base. Stamens 
completely included in upper part of corolla 
tube; filaments thin, upper pair 4 mm long, lower 
pair 3 mm long, white; anthers 1-1.5 mm long, 
orbicular-reniform, light brown. Ovary 1.5-2 
mm long, sessile, conical to ovoid, pale yellow; 
style slender, slightly dilated at apex; stigma 
about 1 mm long, briefly bifid, unequal, the up- 
per lobe somewhat larger, obtuse, green. Fruit 
8-20 mm long, 8-20 mm in diameter, ovoid to 
subglobose, obtuse or apiculate at apex, smooth, 
nitid, dark green turning brownish, with corky- 
punctate or -verrucose outgrowths, pericarp thin, 
0.3 mm thick, crustaceous, drying brittle, tardily 
dehiscent. Seeds 10-20, 5-7 mm long, 2-3 mm 
in diameter, variable in shape, ellipsoid to ob- 
long, angular, dark reddish brown, reticulate-pit- 
ted. Embryo about 4 mm long; slightly curved, 
cotyledons 1.5 mm long, ovate-elliptic. 

DISTRIBUTION Colombia, Venezuela, Ecuador, 
Peru, Brazil, Bolivia. See Figure 28. 



Key to the Subspecies of Brunfelsia grandiflora 

1. Corolla tube 30-45 mm long, 2-3.5 mm in 
diameter, opening at mouth linear-oblong; limb 
35-42 mm in diameter, capsule 17-22 mm 
long 14a. subsp. grandiflora 

1. Corolla tube 15-30 mm long, 1-2 mm in di- 
ameter, opening at mouth elliptic-obovate; 



limb 20-40 mm in diameter; capsule 11-16 
mm long 14b. subsp. schultesii 

14a. Brunfelsia grandiflora subsp. grandiflora 

Brunfelsia calycina f. grandiflora (D. Don) Voss in 
Vilm., Blumengartner, ed. 3, 1: 743. 1896. 

Brunfelsia tastevinii Benoist, Bull. Soc. Bot. France 
75: 295. 1928. Type: BRAZIL. Acre: region de 
Rio Jordao, Tastevin s.n. (holotype, p; isotypes, 
F, P). 

Leaves 10-23 cm long, 3-8 cm wide, glabrous 
or sparingly pubescent at midrib; petiole 3-12 
mm long. Pedicel 2-10 mm long. Calyx 9-13 
mm long, teeth 2-5 mm long, ovate-lanceolate. 
Corolla tube 30-45 mm long, 2.5-3.5 mm in di- 
ameter, the mouth 6-9 mm long; limb 35-52 mm 
in diameter, spreading. Capsule 17-22 mm long. 

DISTRIBUTION Colombia, Ecuador, Peru, Bra- 
zil, Bolivia. 

SPECIMENS EXAMINED COLOMBIA. Meta: 
*25 km S from Granada toward San Juan, Luteyn 
el al. 4769 (F, MO). ECUADOR. Loja: *City of 
Loja, Zarucchi & Andrade 2316 (F). Napo: Mera, 
4 Mar. 1940, Lugo 23 (s); Canton Napo, near 
Achidona, 650 m, 19 Apr. 1935, Mexia 7275 (F, 
s, uc, us); *Costado S del Rio Napo, E de Puerto 
Napo, por carretera nueva hacia Rio Arajuno, 
Neill 6546 (F); *Rio Napo 8 km abajo de Puerto 
Misahualli; Reserva Biologica Jatun Sacha, Neill 
& Palacios 7101 (F). Zamora-Chinchipe: Reg. 
Oriental, Huamboya, 1500-1600 m, 15 Feb. 
1944, Acosta-Solis 7576 (F); Camino Gualaquiza 
al Rio Zamora (Las Jibarias), 850 m, 25 Dec. 
1948, Scolnik 1495 (us); *Zamora-Zumba (along 
Rio Jamboe), Marling & Andersson 13906 (GB, 
GH); *Guadalupe-San Jose de Yacuambi (28 de 
Mayo), along Rio Yacuambi, Harling & Anders- 
son 13934 (GB, GH). PERU. Cusco. *Prov. Pau- 
cartambo; Callanga, Woytkowksi 514 (F). Huan- 
uco: woods above Pozuzo, 6000 ft, Aug. 1863, 
Pearce 290 (K); Monzon, 950 m, 17 Jun. 1958, 
Woytkowski 5026 (F, LE, P, s); Divisoria, 1700 m, 
16 Sep. 1946, Woytkowski 34536 (F, MO, uc); 
*Prov. Tingo Maria; Rio Huallaga, Boeke 1224 
(F). Loreto: Pongo de Manseriche, Colon, 1924, 
Tessmann 5541 (o, s); Rio Ucayali, Canchahuayo, 
20 Nov. 1898, Huber 1490 (MG); Santa Catalina, 
11 Dec. 1898, Huber 1562 (MG); Prov. Coronel 
Portillo, Plantacion Margarita, cerca a la Diviso- 
ria, 1500-1600 m, 14 Aug. 1946, Ferreyra 985 
(B, M, us). San Martin: Rio Huallaga, Rio Chi- 
purana, Tarapoto, 6 Dec. 1898, Huber 1556 (MG); 



78 



FIELDIANA: BOTANY 



BRUNFELSIA grandiflora D.Don 



subsp. 
grandiflora ; 



subsp. sell ul te si i 




FIG. 29. Brunfelsia grandiflora subsp. grandiflora and subsp. schultesii. Reproduced courtesy of the Botanical 
Museum of Harvard University. 



PLOWMAN: REVISION OF BRUNFELSIA 



79 



*Prov. Mariscal Caceres; Dist. Campanilla, Rio 
Sion, Caserio de Sion, Schunke, V. 3480 (NY); 
*Prov. Mariscal Caceres; Dist. Tocache Nuevo, 
Quebrada de Huaquisha, Schunke, V. 4051 (NY); 
*Prov. Mariscal Caceres; Dist. Tocache Nuevo, 
Quebrada de Huaquisha, Schunke, V. 4034 (NY). 
Without department: E of the Andes, 4000- 
6000 ft, Jul. 1884, Pearce 295 (K); woods near 
Rio Bambusi, one of the tributaries of Maranon, 
Mar. 1868, Pearce s.n. (K); in flum. Maranon ripis 
inundatis, Apr. 1855, Spruce 3988 (K); 1909- 
1914, Weberbauer 6151 (F). BRAZIL. Acre: ba- 
sin of Rio Purus near mouth of Rio Macauhan 
(tributary of Rio Yacu), lat. 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; 
isotype, COL). 

Leaves 6-20 cm long, 2-8 cm wide, lanceolate, 
glabrous; petiole 2-6 mm long. Inflorescence 
variable, compact or lax. Pedicel 2-6 mm long. 
Calyx 5-10 mm long, teeth 1-3 mm long, trian- 
gular to triangular-ovate. Corolla tube 15-30 mm 
long, 1-2 mm in diameter, curved toward apex; 
limb 20-40 mm in diameter, spreading, mouth 3- 
5 mm long. Capsule 11-16 mm long, 10-16 mm 
in diameter. 

DISTRIBUTION Colombia, Venezuela, Ecuador, 
Peru, Bolivia, Brazil. 

SPECIMENS EXAMINED COLOMBIA. Amazo- 
nas: Leticia, about 1 km NE of town on road to 
brick factory, 30 Jan. 1969, Plowman & Kennedy 
2310 (ECON); Rio Amazonas, forest along river 
about 2 km downriver from Puerto Narino near 
mouth of Rio Loretoyacu, 5 Feb. 1969, Plowman 
et al. 2404 (GH, INPA, uc); *Rio Amazonas, down- 
river from Puerto Narino near mouth of Rio Lor- 
etoyacu, Plowman et al. 2407 (F). Caqueta: Rio 
Orteguaza, La Maria (Hacienda de Ramon 
Achiardy), 23 km SE of Venecia, 400 m, 7 Mar. 
1944, Hermann 11237 (NY); La Rastra, 22 Apr. 
1956, Vogel 23 (us); Caqueta, Dec. 1930, Uribe 
Uribe P. s.n. (us); Florencia, 420 m, Feb. 1930, 
Perez-Arbelaez 688 (COL, us). Meta: llanos of 
San Martin, 30 Sep. 1916, Da\ve 241 (K, us); San 
Martin, 400 m, 2 Aug. 1946, Uribe Uribe 1353 
(COL). Putumayo: selva higrofila del Rio Putu- 
mayo, Puerto Porvenir, arriba de Puerto Ospina, 
hacia la Loma, 230-250 m, 17 Nov. 1940, Cua- 
trecasas 10611 (COL); Puerto Ospina, 230 m, 25 
Nov. 1940, Cuatrecasas 10795 (COL, us); Mocoa, 
bosque higrofilo en la Quebrada del Rio Mulato, 
570-600 m, 26 Dec. 1940, Cuatrecasas 11275 
(COL, F); camino entre Mocoa y Puerto Asis, 500 
m, 21 Jun. 1935, Garcia Barriga 4586 (COL, us), 
500-700 m, 28 Aug. 1963, Juajibioy Chindoy 277 
(ECON); Pepino, 20 Nov. 1968, Plowman 2039 
(ECON), 1800-2400 ft, 16 Mar. 1953, Schultes & 
Cabrera 19115 (GH); Rio Caqueta, Floresta, about 
15 km downriver from Puerto Limon, 20 Dec. 
1968, Plowman 2183 (ECON); Uchupayaco, en la 
planada entre Urcusique en las orillas del Rio 
Uchupayaco, 300 m, 22 Feb. 1942, Schultes 3340 
(GH); San Antonio del Rio Guamues, 7 Sep. 1966, 



80 



FIELDIANA: BOTANY 



Pinkley 444 (ECON); along trail between San An- 
tonio and Santa Rosa, 1 Dec. 1968, Plowman 
2090 (ECON). VENEZUELA. Barinas: Barinitas, 
Mar. 1953, Aristeguieta 1669 (NY, YEN); Parque 
Barinitas, 600 m, 22 Jun. 1956, Bernardi 3310 
(G). ECUADOR. Napo: Rocafuerte, high forest 
of Hacienda Arcadia, Rio Curaray, 200 m, 7 July 
1933, Heinrichs 496 (F, G), Rio Aguarico, Dureno, 
8 Dec. 1966, Pinkley 575 (ECON); Confluence of 
Quiwado and Tiwaneo Rivers, Davis & Yost 958 
(F); *Limoncocha on Rio Napo, Drummond 7330 
(MO); *Nuevo Rocafuerte, Rio Yasuni, Marling et 
al. 7474 (GB, GH); *Rfo Aguarico, Dureno (30 km 
E of Santa Cecilia), Marling et al. 7700 (GB, GH); 
*Canon de Los Monos, ca. 12 km N of Coca 
(Puerto Francisco de Orellana), Marling & An- 
dersson 11724 (GB); *environs of Limoncocha, 
Madison et al 5390 (F); *Archidona, Pinkley 457 
(F). Pasta/a: *basin of Rio Pastaza, Pacayacu- 
Sarayacu region, Gill 45 (NY). PERU. Cuzco: 
Prov. Paucartambo, Villa Carmen, 720 m, 1 May 
1965, Vargas 16301 (us). Huanuco: *Prov. Leon- 
cio Prado; Tingo Maria, Castillo, Plowman 5825 
(F); *Prov. Pachitea; Dist. Honoria, Bosque Na- 
cional de Iparia, Schunke V. 1480 (NY). Loreto: 
Prov. Maynas, Iquitos region, ca. al Cementerio 
de Iquitos, 120 m, 18 Sep. 1964, Dodson 2822 
(MO); Mishuyacu near Iquitos, 100 m, Jan. 1930, 
Klug 741 (us); Rancho Indiana, 110 m, 24 Jan. 
1932, Mexia 6444 (BM, F, G, GH, K, MO, NY, PH, s, 
uc, us); Rio Nanay, upland forest near Samito, 19 
Feb. 1969, Plowman 2494 (ECON, GH); Rio Mazan, 
Gamitanacocha, 100-125 m, 12 Jan. 1935, Schun- 
ke 8 (A, F, NY, uc, us); Masisea, 275 m, Killip & 
Smith 26844 (us); Yurimaguas, lower Rio Hual- 
laga, 135 m, Aug.-Sep. 1929, Killip & Smith 
27667 (us), 25056 (NY), 281 78 (us), Jan. 1831, 
Poeppig 2206 (w); Balsapuerto (lower Huallaga 
Basin), 150-350 m, 28-30 Aug. 1929, Killip & 
Smith 28644 (us), 220 m, Feb. 1933, Klug 2880 
(A, F, G, GH, MO, NY, s); Pucallpa, 200 m, 4 Aug. 
1946, Soukup 3041 (F); Pongo de Manseriche, 
1924, Tessmann 4921, Rio Maranon, Tessman 
5005 (G); Prov. Ucayali, lower Pisque, Nueva Be- 
liza, 1923, Tessmann 3243 (G, NY, s); *Prov. May- 
nas; Moena Cano between Iquitos and Rio Itaya, 
Gentry et al. 15707 (MO); *IVITA, km 59 Pucall- 
pa-Tingo Maria road, Gentry et al. 18573 (MO); 
*Bosque Nacional von Humboldt, km 86 Pucall- 
pa-Tingo Maria road, Gentry et al. 18664 (MO); 
*Prov. Maynas; Negro Urcu, Rio Napo, McDaniel 
15328 (MO); *Prov. Maynas; Pebas, mouth of Rio 
Ampiyacu, Plowman et al. 6455 (F); *Prov. May- 
nas; Pebas, Plowman et al. 6737 (F); *Prov. May- 



nas; Pebas, Plowman et al. 6939 (F); *Prov. May- 
nas; Dist. Iquitos, Caserio Momoncillo, Revilla 
304 (MO); *Prov. Maynas; Dist. Iquitos, cerca a la 
boca del Rio Nanay, Revilla 1724 (MO); *lower 
Rio Nanay, Williams, LI., 584 (F). Madre de 
Dios: *ca. 5 km from Puerto Maldonado near Rio 
Tambopata, Gentry 16249 (MO). San Martin: Tar- 
apoto and vicinity, W side of Rio Huallaga, S of 
Shapaja 1-4 km, 900 ft, 28-30 Jul. 1937, Belshaw 
3130 (NY, uc, us); Pucaloma, 330-400 m, 12 Jul. 
1950, Ferreyra 7778 (us), 1855-1856, Spruce 
3973 (BM, BR, CGE, G, MG, NY, w), 350 m, Oct. 
1902, Ule 6481 (G, MG, L), 750 m, Dec. 1929, LI. 
Williams 5482 (F), 830 m, 6 Feb. 1947, Woyt- 
kowski 35008 (F, G, MO, uc); entre Bellavista y 
Banos 200-300 m, 5 Sep. 1948, Ferreyra 4760 
(us); Zepelacio near Moyobamba, 1100-1200 m, 
Oct.-Nov. 1933, Klug 3326 (A, F, G, GH, MO, NY, 
s); Roque, 19 Jul. 1925, Melin 186 (s); Saposoa, 
400 m, 14 Oct. 1959, Woytkowski 5525 (F, LE, p, 
us); Rioja, 800 m, 9 Jan. 1961, Woytkowski 6170 
(GH); Boqueron Pass, 92 km from Tingo Maria on 
highway to Pucallpa, ca. 400 m, 16 Dec. 1949-5 
Jan. 1950, Allard 22096 (us); *Prov. San Martin; 
Tarapoto, above Hotel Turista, Plowman 5984 (F). 
Without state: Peru, 1835, Matthews 1320 (CGE, 
K). BRAZIL. Acre: Rio Jurua-Mirim, vicinity of 
Porangaba, 21 May 1971, Maas et al. PI 3228 
(wis); Rio Acre, Xapury, Jan. 1911, Ule 9746 (K, 
MG); Paraguassu, Seringal, Nov. 1911, Ule 9749 
(K, MG); Rio Branco, 26 Feb. 1962, Vasconcelos 
& Coelho s.n. (INPA). Amapa: Rio Oiapoque in 
shade of virgin forest, 5 km SE of Clevelandia, 
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 



81 



Guanai-Tipuani, Bolivian Plateau, Apr.-Jun. 
1892, Bang 1398 (A, BM, F, c, GH, K, M, MICH, MO, 
NY, PH, R, us, w); Mapiri, 550 m, 9 Apr. 1927, 
Buchtien 1299 (us); Mapiri region, San Carlos at 
Sarampiuni, 600 m, 28 Feb. 1927, Buchtien 1300 
(us); Charopampa, 570 m, Nov. 1907, Buchtien 
2029 (us), 5,000 ft, Apr. 1886, Rusby 621 (BM, G, 
GH, K, LE, MO, NY, PH, us, w); Tumu-pasa, 1800 ft, 
7 Jan. 1902, White 1840 (BM, NY, us). Pando: 
bank of Rio Madeira, 6 km above Abuna, 1 1 July 
1968, Prance et al. 5854 (wis); Rio Acre, Cobija, 
Dec. 1911, Ule 9747 (K, MG), Jan. 1912, We 9748 
(G, K, MG). Santa Cruz: Prov. Gutierrez, Canton 
Buenavista, Dolores, 450 m, Steinbach 1805 (A, 
uc); Buenavista, 500 m, 14 Dec. 1920, Steinbach 
5487 (GH), 450 m, 2 Jan. 1926, Steinbach 7399 
(A, K); *Buena Vista, Bella Union, Steinbach 7936 
(A, BM, F, K). 

Brunfelsia grandiflora was first collected dur- 
ing the Ruiz and Pavon expedition in 1798. Ben- 
tham (1846) considered it to be very similar to 
his B. latifolia and perhaps also to its variety. 
Most material that Bentham referred to B. latifolia 
is now considered B. grandiflora subsp. schultesii, 
indicating that he recognized the affinities of the 
plants concerned, although he misinterpreted B. 
latifolia, which is endemic in Rio de Janeiro (Bra- 
zil). 

Brunfelsia grandiflora consists of two subspe- 
cies: subspecies grandiflora and subspecies schul- 
tesii. Subspecies grandiflora is much more limited 
in its distribution than is the second subspecies. It 
occurs in the drainage systems of the Ribs Mar- 
anon, Huallaga, and Ucayali in Peru; at higher 
elevations in eastern Ecuador; and in the upper- 
most tributaries of the Rios Purus and Jurua in 
Brazil. It grows primarily at elevations of 650- 
2000 m in the region known as the montana, hu- 
mid, montane rain forest that stretches from Co- 
lombia to northeasternmost Argentina on the east- 
ern slopes of the Andes. 

Brunfelsia grandiflora subsp. grandiflora is 
characterized by its large flowers (to 50 mm 
across) and fruits (to 22 mm in diameter), as well 
as by its higher altitudinal preference. Subspecies 
schultesii looks very similar but is generally 
smaller in all its parts (see key to subspecies). 
From the relatively few collections available, sub- 
species grandiflora appears to be variable and ex- 
hibits some exceptional forms. For example, col- 
lections from the northern part of the range (Ec- 
uador and northern Peru) often have divaricating, 
not spreading, lateral nerves in the leaves and 



longer pedicels (Pearce 290, Lugo 23). The latter 
feature also appears in high-elevation populations 
of B. uniflora in Bolivia. 

In the 1920s, a Catholic missionary named Tas- 
tevin collected this subspecies in the Rio Tarauaca 
in the state of Acre, Brazil. He stated that it was 
a high-climbing liana (Tastevin, 1922) and con- 
fused the plant with the psychotropic Banisteriop- 
sis. Nevertheless, other collectors have indicated 
that the plants may reach 10 m in height, an ex- 
ceptionally large size. Whether B. grandiflora 
subsp. grandiflora may actually become lianoid in 
habit has yet to be confirmed by field observa- 
tions. 

Brunfelsia grandiflora subsp. grandiflora is 
well-known in cultivation, especially in the New 
World tropics, as an ornamental. In the Amazon 
region, it is also cultivated for its narcotic and 
medicinal properties [see Plowman, 1977; Schul- 
tes & Raffauf, 1990, 1991 Eds.] and is known 
by several common names: sanango, chici-qui- 
sanango, chiricsanango, and sananguillo (Peru); 
and manacd grande (Brazil). 

Brunfelsia grandiflora subsp. schultesii has 
long been recognized in the literature under sev- 
eral misapplied names, including B. bonodora, B. 
latifolia, and B. maritima. This was primarily the 
result of "follow-the-leader" taxonomy in which 
no botanist consulted the type material on which 
the original descriptions were based. It is now 
clear that these other names all apply to species 
that are found only in southeastern Brazil. 

Brunfelsia grandiflora subsp. schultesii is wide- 
ranging and polymorphic, occurring in western 
South America from Venezuela southward to Bo- 
livia. In addition, disjunct populations are known 
from the Territory of Amapa in northern Brazil. 
It grows commonly as an understory shrub in pri- 
mary and secondary forests between 100 and 900 
m elevation. Because this subspecies is widely 
cultivated in South America for medicinal and or- 
namental purposes, some activity of man has un- 
doubtedly influenced its present distribution. 

The two subspecies of B. grandiflora differ pri- 
marily in the size and form of the flowers and in 
the size of the fruits. Subspecies schultesii also 
tends to be smaller in habit and leaf size, but there 
is considerable overlap between the subspecies in 
these characters. 

The two subspecies are also ecologically and, 
to some extent, geographically isolated. The typ- 
ical subspecies is usually found between 900 and 
2000 m elevation in the inter-Andean valleys of 
Peru. Brunfelsia grandiflora subsp. schultesii nor- 



82 



FIELDIANA: BOTANY 



mally occurs at much lower elevations, between 
100 and 900 m, throughout the upper Amazon and 
its tributaries, although isolated populations in 
Bolivia may reach 1800 m. From morphological 
and geographical evidence, it appears that subspe- 
cies grandiflora has arisen from its much more 
widespread and variable counterpart through iso- 
lation in the intermontane valleys. 

Intermediates between B. grandiflora subsp. 
grandiflora and B. grandiflora subsp. schultesii 
are known from areas where the two coexist. This 
occurs in the Brazilian state of Acre and in the 
lower Maranon valley of Peru near Pongo de 
Manseriche (cf. Tessmann 4921, Krukoff 5284, 
Vie 5453). These specimens are intermediate in 
the size of their flowers and fruits, which suggests 
hybridization between the two subspecies in these 
areas. No intermediate forms have been found in 
the higher montane valleys or in the tropical low- 
lands. 

As would be expected from such a wide-rang- 
ing plant, B. grandiflora subsp. schultesii shows 
considerable morphological variation. Some of 
this variation may be due to environmental fac- 
tors such as those that exist between the llanos 
of Venezuela and the cloud forests of Bolivia, 
but it seems that some populations have become 
genetically differentiated as well since morpho- 
logically distinct groups can be readily and con- 
sistently separated. These variants, however, are 
relatively slight and in my judgment should not 
receive formal names. I will instead discuss 
some of the more prevalent variations that occur. 
The most common form of subspecies schultesii, 
represented by the type collection (Schultes et al. 
24108), is found from the Colombian Putumayo 
south to Tarapoto in Peru and eastward to Leticia 
in the Colombian Amazon. These plants have 
large, more or less oblong-lanceolate leaves and 
many flowers in a compact inflorescence. It is 
this form that occurs disjunct in the north Bra- 
zilian state of Amapa (Invin 47359, 48753). This 
is the only known collection east of Leticia, a 
disjunction of some 2300 km. The species is ap- 
parently absent from the lower Amazon, al- 
though additional collecting in this region may 
prove otherwise. 

In the northern part of its range, in the llanos 
of Venezuela and Colombia and south to the Co- 
lombian Putumayo, another form of B. grandiflo- 
ra subsp. schultesii occurs. These populations 
have large, oblong, subcoriaceous leaves and 
more lax, branched inflorescences (cf. Ariste- 
guieta 1669, Perez-Arbelaez 688). The leaves in 



some specimens approach those of B. chiricaspi 
in their large size and ochraceous coloring on 
drying. These plants apparently survive in scat- 
tered patches of woodland in the llanos and in 
gallery forests along the rivers. 

A third form of B. grandiflora subsp. schul- 
tesii may be distinguished among populations 
occurring on the eastern slopes of the Bolivian 
plateau (cf. R. S. Williams 425) in wet, montane 
forests up to 1600 m altitude. These plants have 
rather small, lanceolate leaves and few flowers 
per inflorescence, rarely reduced to three. These 
populations show strong similarities to B. bono- 
dora of the coastal forests of Rio de Janeiro, a 
species still very poorly known. Brunfelsia gran- 
diflora subsp. schultesii differs primarily in hav- 
ing shorter pedicels and a smaller calyx, al- 
though additional characters may be found when 
more material becomes available. The floristic 
affinities between the Yungas regions of Bolivia 
and Rio de Janeiro have been pointed out else- 
where (Plowman, 1979). 

These differing forms of B. grandiflora subsp. 
schultesii intergrade completely with adjacent 
populations and should not be considered taxo- 
nomically significant. They are mentioned here 
to point out the degree of variation that exists in 
this subspecies, which may, in some cases, lead 
to confusion with other species. 

Brunfelsia grandiflora subsp. schultesii grows 
usually as a shrub 1-2 m tall, at times becoming 
a treelet to 5-6 m. It lives in the understory of 
shady, moist forests; at the margins of forest 
clearings; and along rivers. It commonly persists 
after the forest has been cut for lumber or shift- 
ing agriculture, sprouting easily from the roots. 
Considered a valuable ornamental and medicinal 
plant, subspecies schultesii is much planted in 
Indian houseyards and fields and often persists 
after the inhabitants move elsewhere. In this 
way, the plant has probably increased its distri- 
bution under the influence of man. There is some 
indication that its occurrence as far east as Le- 
ticia in the Colombian Amazon may be due to 
man's activities. Populations in this area have so 
far been located only near the major rivers in 
places most likely inhabited at some time in the 
past. 

Brunfelsia grandiflora subsp. schultesii flow- 
ers and fruits sporadically throughout the year 
over most of its range, with two main periods of 
flowering: December to February and July to 
August. The flowers are usually unscented, al- 
though they may occasionally contain the odor 



PLOWMAN: REVISION OF BRUNFELSIA 



83 



of jasmine (Plowman 2019, Ducke 865). This 
plant is known by a number of vernacular names 
that are indicative of its usefulness in native 
medicine [these are given, along with an account 
of the medicinal and narcotic uses, in Plowman, 
1977 Eds.]. 

15. Brunfelsia hydrangeiformis (Pohl) Benth. in 
DC., Prodr., 10: 198. 1846. 

Franciscea hydrangeiformis Pohl, PI. Bras. Ic. 1: 7, 
t. 7. 1826. Type: BRAZIL. Rio de Janeiro: ad 
Olaria et Sumido rio in via ad Rio Parahybuna, 
habitat inter frutices locis umbrosis, floret Sep- 
tembri et Octubri, 1818, Pohl (115) (holotype, 
w; isotypes, F-875036, M, w). 

Shrub 1-2 m tall. Stem erect, sparsely 
branched at intervals. Bark grayish brown. 
Branches few, appearing at intervals, 4-6 mm in 
diameter, stout, naked, glabrous, the epidermis 
splitting longitudinally. Leaves crowded at ends 
of branches, alternate, subverticellate with 5-10 
together, blade 13-32 cm long, 3-15 cm wide, 
oblanceolate to subspathulate, rarely lanceolate to 
oblong, abruptly to long acuminate with straight 
or subfalcate acumen, rarely acute, narrowed-cu- 
neate at base, the upper surface glabrous, often 
nitid, dark green, the lower surface sparsely pu- 
bescent mostly at nerves, or glabrous, pale green, 
membranaceous to subcoriaceous, reticulately ru- 
gose, with sulcate nerves, lateral nerves 7-11, 
spreading or diverging, depressed in sulci; petiole 
2-13 mm long, glabrous. Inflorescence terminal, 
forming in uppermost whorl of leaves, densely 
capituliform to corymbiform, rarely lax, hemi- 
spheric, sessile, the axis 1-3 cm long, persistent. 
Flowers 5-50, purple with white spot at mouth, 
fading to nearly white with age. Bracts paired, 2- 
10 mm long, lanceolate to linear-lanceolate, pu- 
bescent or glabrous, ciliolate at margin. Pedicel 
2-8 mm long, glabrous or rarely pubescent. Calyx 
15-35 mm long, 3-10 mm in diameter, tubular to 
tubular-campanulate, oblong in bud, slightly in- 
flated, variably pubescent, densely pilose, glan- 
dular, or glabrous, light green to ferrugineous, thin 
to firmly membranaceous, teeth 2-10 mm long, 
erect, ovate to linear-lanceolate, acute to acumi- 
nate at apex; calyx in fruit, thin, completely en- 
closing fruit. Corolla tube 20-35(45) mm long, 



1-2 mm in diameter, 1-1 Vi times as long as calyx, 
rarely longer, slender, glabrous or glandular pu- 
bescent; limb 18-32 mm across, spreading, lobes 
7-12 mm long, rounded, overlapping at lateral 
margins. Stamens included in upper part of co- 
rolla tube; filaments slender, upper pair 5 mm 
long, lower pair 3 mm long; anthers 1.5-2 mm 
long, reniform. Ovary 2-2.5 mm long, ovoid; 
style slender, equaling the filaments; stigma brief- 
ly bifid, lobes unequal, about 1 mm long. Fruit 
completely enclosed by calyx, 12-20 mm long, 
9-15 mm in diameter, ovoid-subglobose, apicu- 
late at apex, smooth, greenish brown, pericarp 
thin-walled, dry at maturity, crustaceous, sparing- 
ly dehiscent. Seeds ca. 5-10, 5 mm long, 2 mm 
in diameter, oblong, irregularly angular, dark 
brown, deeply reticulate-pitted. Embryo un- 
known. 

DISTRIBUTION Brazil. See Figure 33. 

Note: According to the International Code of 
Botanical Nomenclature (Article 60.8, Recom- 
mendation 60G) (Greuter et al., 1994), it is rec- 
ommended that in compound names of Latin or- 
igin, the final vowel of the first part be reduced 
to i before a consonant. Thus, correctly, the orig- 
inal orthography of hydrangeaeformis is correct- 
able to hydrangeiformis. 



Key to the Subspecies of Brunfelsia hydrangei- 
formis (Pohl) Benth. 

1. Leaves subspathulate to broadly oblanceolate, 
reticulately rugose with sulcate nerves; pedicel 
2-4 mm long; calyx 3-6 mm in diameter with 
yellowish brown pubescence; calyx teeth 4-10 

mm long, linear to linear-lanceolate 

15a. subsp. hydrangeiformis 

1 . Leaves oblong to oblong-lanceolate, smooth or 
sulcately nerved; pedicel 4-8 mm long; calyx 
5-10 mm in diameter, glabrous or with sparse 
glandular hairs; calyx teeth 2-4 mm long, 

ovate to ovate-lanceolate 

15b. subsp. capitata 

15a. Brunfelsia hydrangeiformis subsp. hydran- 
geiformis. Figure 30 and frontispiece. 



FIG. 30. Brunfelsia hydrangeiformis subsp. hydrangeiformis (originally as Franciscea hydrangeiformis). From 
Pohl, J. E., Plantarum Brasiliae Icones et Descriptiones (1826). 



84 



FIELDIANA: BOTANY 




PLOWMAN: REVISION OF BRUNFELSIA 



85 



Franciscea macrophylla Cham. & Schltdl., Linnaea 
2: 601. 1827. Type: BRAZIL. Sellow sp. nov. 
(holotype destroyed at Berlin; fragment of ho- 
lotype, F-643247; photograph of holotype, 
F-621825). 

Brunfelsia macrophylla (Cham. & Schltdl.) Benth. in 
DC., Prodr. 10: 198. 1846. 

Leaves with blade 18-32 cm long, 6-15 cm 
wide, subspathulate to broadly oblanceolate, 
abruptly acuminate, glabrous on both sides, retic- 
ulately rugose with sulcate nerves depressed from 
above. Pedicel 2-4 mm long. Calyx 18-35 mm 
long, 3-6 mm in diameter, tubular, scarcely in- 
flated, sparsely pubescent to velutinous with yel- 
lowish hairs, otherwise purple to brownish, dry- 
ing ferrugineous, teeth 4-10 mm long, linear to 
linear-lanceolate. Corolla tube briefly exserted 
from calyx. 

DISTRIBUTION Brazil (Bahia, Minas Gerais, 
Rio de Janeiro, Sao Paulo). 

SPECIMENS EXAMINED BRAZIL. Bahia: 1857, 
Blanchet s.n. (G). Minas Gerais: Pirange, Oct. 
1840, Gardner 5065 (BM, CGE, F, G, GH, K, NY, p, 
us, w); Sao Francisco de Paula, Rio do Peixe, 
Aug. 1896, Magalhaes 1643 (R); Mun. Serro, Bo- 
cada Mata, 1400 m, 6 Oct. 1945, Williams & As- 
sis 7928 (GH, R, uc, us); Sabara, Nov. 1839, 
Claussen s.n. (BM, BR, G, p); without locality, 
Claussen s.n. (CGE), Claussen 201 (BR), Langs- 
dorffs.n. (us), Sep. 1836, Raben 14 (c), Schiich 
591 (CGE, w), 1858, Weddell 878 (G); *km 30-32 
on the road from Batinga to Jequitinhonha, Bun- 
ting et al. 10351 (F). Rio de Janeiro: Floresta da 
Tijuca, 7 Aug. 1879, Glaziou 11393 (K, p), Nov. 
1894, Ule s.n. (R, us); Serra de Estrella, 1841, 
Claussen s.n. (w); estrada entre a Estrella e a Es- 
trada Rio-Petropolis, 24 Oct. 1932, J.G. Kuhl- 
mann s.n. (RB); Nova Friburgo, Oct. 1947, Leite 
4188 (NY); Serra de Friburgo, Sitio do Gaucho, 
Oct. 1964, Duarte 8445 (HB); Cachoeira de Ma- 
cacu, 22 Oct. 1964, Duarte & Pereira s.n. (GH, 
RB); Serra Tingua, Rio San Antonio, 24 Aug. 
1879, Glaziou 11392 (c, NY, p); Rio Parahyba, 
1887, Moura s.n. (R); Ipiahos, Aug. 1934, O. 
Peckolt s.n. (R); Magdalena, Marreiras, Oct. 1933, 
Santos Lima 200 (RB); without locality, Langs- 
dorff43 (LE), 1816-1821, St. Hilaire A-555 (p); 
*Nova Friburgo, Parada do Gaucho, Pereira 9241 



(F); *Municipio Santa Maria Madalena; Parque do 
Desengano, Santa Clara, Araujo 954 (F); *near 
Mury, Maas et al. 3326 (u). Sao Paulo: *Muni- 
cipio de Banana]; Parque Nacional da Serra Bo- 
caina, Martinelli 1115 (F). Without state: 1814- 
1817, Bowie & Cunningham 100 (BM), Sellow s.n. 
(F, M, MO, s). 

15b. Brunfelsia hydrangeiformis subsp. capitata 

(Benth.) Plowman comb, et slat. nov. Figure 
31. 

Brunfelsia capitata Benth. in DC., Prodr., 10: 198. 
1846. Type: BRAZIL. Rio de Janeiro: Serra dos 
Orgaos, shrub about 4 ft high growing in rather 
moist places in virgin forests reaching an ele- 
vation of about 4500 ft, manaca dobrado, Feb. 
1837, Gardner 563 (holotype, K; isotypes, BM, F 
678955, G, NY, P, w). 

Franciscea capitata (Benth.) Miers, Ann. Mag. Nat. 
Hist. Sen 2, 5:250. 1850. 

Brunfelsia capitata var. angustifolia Benth. in DC., 
Prodr., 10:198. 1846. Type: BRAZIL. Rio de Ja- 
neiro: Organ Mountains, 1841, Gardner 5826 
(holotype, K). 

Brunfelsia hydrangeiformis var. glabriscula J. A. 
Schmidt in Mart., Fl. Bras. 8(1):256. 1864. Lec- 
totype (designated here): BRAZIL, prope Rio, 
Gardner 563 (isotypes, BM, F, G, K, NY, p, w). 
Lectoparatype: BRAZIL. Rio de Janeiro: in 
montibus prope Nova Friburgo, Jul. 1832, Riedel 
85 (LE). 

Leaves with blade 13-26 cm long, 3-11 cm 
wide, oblong to oblanceolate, rarely lanceolate, 
acute to acuminate at apex, glabrous on upper sur- 
face, puberulent beneath at nerves or glabrous, 
smooth to sulcately nerved, rarely reticulate ru- 
gose. Pedicel 4-8 mm long. Calyx 14-35 mm 
long, 5-10 mm in diameter, tubular to tubular- 
campanulate, often somewhat inflated, glabrous or 
sparsely glandular pubescent, light green, teeth 2- 
4(7) mm long, ovate to ovate-lanceolate. Corolla 
tube rarely exserted from calyx. 

DISTRIBUTION Brazil (Espirito Santo, Minas 
Gerais, Rio de Janeiro, Sao Paulo). 

SPECIMENS EXAMINED BRAZIL. Espirito 
Santo: Santa Barbara de Caparao, 4 Dec. 1929. 
Mexia 4089-a (uc, us). Minas Gerais: Estasoao 
Experimental de Cafe, Coronel Pacheco, 29 Nov. 
1940, Heringer 460 (RB); without locality, Ker- 



FIG. 31. Brunfelsia hydrangeiformis subsp. capitata comb, et stat. nov. Isotype specimen in the herbarium of The 
Natural History Museum, London (BM). 



86 



FIELDIANA: BOTANY 



Del. bl T. PlowiMB Wl - <V & . "JI 







, /</; ; 



PLOWMAN: REVISION OF BRUNFELSIA 



87 



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- 
per surface. 

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- 
geiformis. 

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- 



FIELDIANA: BOTANY 




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 
Hydrangea hortensis. 

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 



89 




FIG. 33. Distribution of Brunfelsia hydrangeiformis (solid triangle), B. imatacana (solid square), and B. mire 
(solid circle). 



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- 
ted. 

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 



90 



FIELDIANA: BOTANY 



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- 
mark, 1979). 

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- 
type, w). 

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 



91 




./? 




92 



FIELDIANA: BOTANY 



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 
Figure 37. 

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. 
latifolia. 

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 
ornamental. 

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 
Descriptiones (1826). 



PLOWMAN: REVISION OF BRUNFELSIA 



93 



BRUNFELSIA macrocarpa Plowman 




FIG. 35. Brunfelsia macrocarpa. Reproduced courtesy of the Botanical Museum of Harvard University. 



94 



FIELDIANA: BOTANY 



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, 
broadly ovate. 

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 



95 



BRUNFELSIA mire 

Monachino 




FIG. 36. Brunfelsia mire. Reproduced courtesy of the Botanical Museum of Harvard University. 



96 



FIELDIANA: BOTANY 



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 
Figure 32. 

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 



97 



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 
1067018, w). 

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- 
bryo unknown. 

DISTRIBUTION Brazil. See Figure 37. 



Key to the Varieties of Brunfelsia obovata 
Benth. 

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- 



98 



FIELDIANA: BOTANY 




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 
38. 

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, 
Sao Paulo). 

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 



99 




BRIJNFELSLA 



100 



FIELDIANA: BOTANY 



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 
Figure 39. 

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, 
Gois, Piaui). 

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- 
man's. Eds. 



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. 
calycina. 

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 
(Peckolt, 1909). 

Brunfelsia obovata var. coriacea corresponds 



FIG. 38. Brunfelsia obovata var. obovata. From Martius, Flora Brasiliensis (1846). 



PLOWMAN: REVISION OF BRUNFELSIA 



101 




w 

..1812226 

FIELD MUSEUM 

OF 
NATURAL HISTORY 



THE NEW YORK BOTANICAL GARDEN 
Plants of the Planalto do Brasil 

Distrito Federal 
No. 851U 

< Isin nbov.i*,a ssn. c -i-iaca 
.dot. w<*n la 

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sriKTRsr* sssMts^esai 




102 



FIELDIANA: BOTANY 



well to B. silvicola published in 1896 by Taubert, 
which is here placed in synonymy. The type of B. 
silvicola was destroyed in Berlin in 1945, but an- 
other collection (Ule 77) from the same locality 
is extant and may serve as a representative col- 
lection and may possibly be a topotype. 

Brunfelsia obovata var. coriacea flowers in the 
month of September, somewhat earlier than var. 
obovata; fruits have been collected in March. Va- 
riety coriacea appears to grow under the same 
ecological conditions as var. obovata (swampy ar- 
eas) but is found in a generally much drier region 
of cerrado vegetation, probably being confined to 
localized wet areas. 

21. Brunfelsia pauciflora (Cham. & Schltdl.) 
Benth. in DC., Prodr. 10: 199. 1846. Figure 
40. 

Franciscea pauciflora Cham. & Schltdl., Linnaea 2: 
600. 1827. Type: BRAZIL. E Brasilia meridion- 
ali, without locality or date, Sellow s.n. (holotype 
destroyed at Berlin; fragment, F 621699; photo- 
graphs, F, NY; isotypes, M). 

Besleria inodora Veil., Fl. Flum.: 261. 1829 
("1825"), Ic. 6, t. 81. 1831 ("1827"). Lectotype 
(designated here): plate 81 of Vellozo. 

Brunfelsia calycina Benth. in DC., Prodr. 10: 199. 
1846. Lectotype (designated here): BRAZIL. 
Santa Catarina: Insula Santa Catarina, 1832, 
Baron de Colchester s.n. (K). Lectoparatype: 
BRAZIL. Santa Catarina: Insula Santa Catarina, 
Bade s.n. (specimen not located). Lund 755, also 
cited by Bentham, is excluded as a type. It is B. 
obovata Benth. var. obovata. 

Franciscea calycina (Benth.) Miers, Ann. Mag. Nat. 
Hist. ser. 2, 5: 250. 1850. 

Brunfelsia pauciflora var. calycina (Benth.) J. A. 
Schmidt in Mart., Fl. Bras. 8(1): 257. 1864. 

Franciscea eximia Scheidw. ex Moore & Ayres, 
Card. Mag. Bot. 1: 16. 1850. Lectotype (desig- 
nated here): Moore & Ayres, Gard. Mag. Bot. 2: 
177 + t. 1850. Representative collection: culti- 
vated at Royal Botanic Gardens, Kew, Herbari- 
um Hookerianum, 1 867, without collector (K). 

Brunfelsia eximia (Scheidw.) Bosse, Handb. Blu- 
meng. 524. 1859. 

Brunfelsia calycina var. eximia (Scheidw.) L. H. Bai- 
ley & Raffill, Stand. Cyclop. Hort. 1: 581. 1914. 

Franciscea macrantha Lem., Jard. Fleur. 3: 349, t. 
249. 1853. Lectotype (designated here): Lemai- 
re, Jard. Fleur. 3: t. 249. 1853. Representative 
collection: Royal Botanic Gardens, Kew, culti- 
vated in temperate house, 1 1 Nov. 1952, Souster 
s.n. (K). 

Brunfelsia calycina var. macrantha (Lem.) L. H. Bai- 



ley & Raffill, Standard Cyclop. Hort. 1: 581. 
1914. 

Franciscea lindeniana Planch., Belgique. Horticole. 
15: 100, t. 16. 1865. Type: Planchon, Belgique. 
Horticole. 15: 100, t. 16. 1865. Representative 
collection: Royal Botanic Gardens, Kew, culti- 
vated in temperate house, 16 Jun. 1953, Souster 
s.n. (K) (see Fig. 42). 

Brunfelsia lindeniana (Planch.) G. Nicholson, 111. 
Diet. Gard. 1: 216. 1884. 

Brunfelsia calycina var. lindeniana (Planch.) Raffill, 
Gard. Chron. ser. 3, 83: 370. 26 May 1928. 

Brunfelsia calycina var. floribunda L. H. Bailey & 
Raffill, Standard Cyclop. Hort. 1: 581. 1914. 
Representative collection: New Jersey, Ruther- 
ford, cultivated at Julius Roehrs Co., 29 Mar. 
1918, Bailey s.n. (BH). 

Shrub 1-3(5) m tall. Bark somewhat smooth 
or longitudinally cracked, grayish brown. Branches 
sparse, erect or spreading. Bra nth lets stout, gla- 
brous, rarely pubescent or glandular, smooth, dark 
green. Leaves scattered or crowded near branch 
tips, blade 6.5-16 cm long, 2-6.5 cm wide, ob- 
long to oblong-lanceolate, more rarely elliptic-ob- 
long or obovate-oblong, apically acute to short 
acuminate, sometimes blunt or emarginate, cune- 
ate to attentuate at base, glabrous, punctate or 
rarely glandular pubescent beneath, chiefly at 
midrib, upper surface dark green, dull to nitid, 
lower surface pale green, firmly membranaceous 
to subcoriaceous, lateral nerves 5-11, spreading, 
straight or broadly arching; petiole 5-12 mm long, 
glabrous or pubescent. Inflorescence terminal, 
sessile or subsessile, with 1-11 flowers. Flowers 
showy, deep, reddish purple fading to pale lav- 
ender or white with age, with white spot at mouth 
sometimes edged with violet. Bracts small, 1-3 
per flower, 1-8 mm long, linear-lanceolate, con- 
cave, narrowly acute apically, glabrous or glan- 
dular ciliolate, caducous. Pedicel 1 1-25(35) mm 
long, stout, broader toward apex, glabrous or 
sparsely glandular, in fruit becoming thicker and 
corky-verrucose. Calyx 18-32 mm long, 6-10 
mm in diameter, tubular, rarely tubular-campanu- 
late or inflated, punctate to glandular pubescent or 
completely glabrous, firmly membranaceous to 
subcoriaceous, pale to dark green, teeth 3-8 mm 
long, ovate to ovate-lanceolate, incumbent, acute 
to acuminate at apex; calyx in fruit campanulate- 
urceolate, completely enclosing fruit, becoming 
slightly larger and thick-coriaceous, conspicuous- 



FlG. 39. Brunfelsia obovata var. coriacea. 



PLOWMAN: REVISION OF BRUNFELSIA 



103 



3 cm 




BRUNFELSIA 



pauci flora (Cham.K Schlecht] Benth. 

FIG. 40. Brunfelsia pauciflora. Reproduced courtesy of the Linnean Society, London. 



104 



FIELDIANA: BOTANY 




FIG. 41. Distribution of Brunfelsia pauciflora (solid circle) and B. rupestris (solid square). 



ly striate-nerved with light brown, lenticular-ver- 
rucose outgrowths. Corolla tube 28-36 mm long, 
1.5-3 mm in diameter, 1-2 times as long as calyx, 
reddish purple above, paler beneath, ring at mouth 
4-5 mm long, elliptic, white, sparsely glandular; 
lobes 15-30 mm long, broadly rounded to subel- 
liptic, rounded-truncate or blunt at apex, some- 
what accrescent with age, more or less overlap- 
ping laterally. Stamens included in upper part of 
tube; filaments subterete, anteriorly canaliculate, 
white, upper pair 4-6 mm long, lower pair 2-4 
mm long; anthers 1.5-2 mm long, orbicular-reni- 
form, greenish brown. Ovary 2-3 mm long, 1.5 
mm in diameter, conical-ovoid, pale green; style 
25-30 mm long, filamentous, lavender; stigma 1 
mm long, briefly bifid, lobes subequal, white. 
Capsule included in calyx, 20-22 mm long, 15- 
18 mm in diameter, subglobose to ovoid, smooth, 



light green, pericarp thin, dry at maturity, spar- 
ingly dehiscent. Seeds 12-30, 5-6 mm long, 2.5- 
3 mm in diameter, ovoid or oblong, angular, dark, 
reddish brown, reticulate-pitted. Embryo ca. 5 
mm long, straight; cotyledons 2-3 mm long, 
ovate, flat. 

DISTRIBUTION Brazil (Parana, Rio de Janeiro, 
Santa Catarina, Sao Paulo). See Figure 41. 

SPECIMENS EXAMINED BRAZIL. Parana: 
Ponta Grossa, 16 Oct. 1910, Dusen 10305 (s), 16 
Oct. 1910, Dusen 10354 (F, GH, MICH, NY, s), 19 
Oct. 1911, Dusen 8679 (s); Carvalho, 18 Nov. 
1909, Dusen 8961 (F, GH, K, L, MICH, MO, NY, p, 
s), 5 Nov. 1911, Dusen 13298 (F, GH, K, L, MICH, 
MO, NY, P, s), 7 Nov. 1911, Dusen 13327 (F, GH, 
NY, s); Jaguariahyba, 10 Oct. 1911, Dusen 13180 
(GH, s); Sena do Mar, Itupava, 450 m, 4 Nov. 



PLOWMAN: REVISION OF BRUNFELSIA 



105 



1915, Dusen 17301 (GH, NY, s); Mun. Sao Jose 
dos Pinhais, Campo Largo da Roseira, 13 Oct. 
1946, Hatschbach 515 (G, us); Mun. Piraguara, 
Bauhado, 2 Nov. 1948, Hatschbach 1058 (us); 
Mun. Campina Grande do Sul: Rio Pardinho, Ver- 
tente Altantico, 29 Oct. 1965, Hatschbach 13055 
(F, P, NY, us); Pico Caratuva, 1500-1600 m, 5 Oct. 

1967, Hatschbach 17308 (c, uc); Mun. Morretes, 
Rio Ipiranga, 29 Nov. 1966, Hatschbach 15300 
(NY, us); Mun. Quatro Barras, Rio Taquari, 8 Oct. 

1968, Hatschbach 19937 (MICH, MO); km 40 da 
estrada Curitiba-Paranagua, 17 Oct. 1961, Pabst 
5393/Pereira 6066 (F, w); *Quatro Barras, Morro 
Mae Catiro, Hatschbach 15071 (MBM); *Campina 
Grande do Sul, Serra Virgem Maria, Hatschbach 
20304 (MBM); *Municipio Sao Mateus do Sul; Te- 
zoura, Hatschbach 22278 (MBM); *Municipio 
Campina Grande do Sul; Serra Capivari Grande, 
Hatschbach 22771 (MBM); *Municipio Cerro 
Asul; Barra do Turvo, Hatschbach 32637 (MBM); 
*Municipio Sao Jose dos Pinhaes; Guaricana, 
Hatschbach 34918 (MBM, MO); *Municipio Mor- 
retes; Estrada da Graciosa, Grota Funda, Hatsch- 
bach 40004 (F); *Quatro Barras, Arredores, 
Hatschbach 43273 (F); *Municipio Sao Jose dos 
Pinhaes; Purgatorio, Hatschbach 45295 (F); 
*Municipio Curitiba; Capao do Centre Politecni- 
co, Hatschbach & Cervi 46844 (F); *Municipio 
Quatro Barras; Morro do Anhangara, Kummrow 
1575 (F); *Municipio Guaratuba; Serra do Ara- 
catuba, Kummrow 2420 (F). Rio de Janeiro: Alto 
Macahe de Nova Friburgo, 30 Oct. 1857, Glaziou 
17168 (c, P, R); Serra do Mocoto, Santa Magda- 
lena, 1400 m, Aug. 1940, Santos Lima s.n. (RB); 
Rio de Janeiro, 1867, Glaziou 811 (BR, c), 1842, 
Raben 746 (BR). Santa Catarina: Mun. Brusque: 
Mata de Hoffman, 50 m, 19 Oct. 1950, Klein 319 
(us); Azambuja, 41 m, 25 Nov. 1947, Reitz 2768 
(uc, s); Brusque, 35 m, 25 Nov. 1947, Reitz C 
1952 (s, us); Mato do Malucher, 40-50 m, 23 
Feb. 1952, L. B. Smith 5786 (us). Mun. Itajai: 
Itajai, Morro da Fazenda, 150 m, 3 Nov. 1955, 
Klein 1751 (M, us), 17 May 1969, Plowman 2739 
(GH); Itajai, Sep. 1870, Mueller 435 (K); Bra? 
Joaquim, Luis Alves, 300 m, 30 Sep. 1954, Reitz 
& Klein 2160 (G, us); Una de Santa Catarina, Saco 
Grande, 300 m, 23 Nov. 1966, Klein & Bresolin 
6864 (us). Mun. Bom Retire, Riozinho, 1000 m, 
23 Dec. 1948, Reitz 2743 (us); pinheiral, 25 Nov. 
1956, Smith, Reitz & Klein 7936 (K, NY, R, us); 
Mun. Ibirama, Horto Florestal I.N.P., 350 m, 2 
Nov. 1953, Reitz & Klein 1158 (us), 450-700 m, 
12 Nov. 1956, Smith & Klein 7532 (R, us); Blu- 
menau, Mata da Companhia Hering, 300 m, 22 



Oct. 1959, Reitz & Klein 9205 (us). Mun. Rio do 
Sul, Serra do Matador, 550 m, 17 Oct. 1958, Reitz 
& Klein 7329 (us). Joinville, Estrada Dona Fran- 
ciscea, 450 m, 26 May 1957, Reitz & Klein 4238 
(us), 600 m, 6 Nov. 1957, Reitz & Klein 5601 
(NY, uc, us). Sao Paulo: Moaca, 22 Sep. 1912, 
Brade 6036 (s, SP); Mun. Iguape, Morro das Ped- 
ras, Sep. 1920, Brade 8088 (RB); Iguape, 20-100 
m, Sep. 1901, Wettstein & Schiffner s.n. (w); cam- 
inho do Porto da Ribeira, 4 Oct. 1894, Lofgren & 
Edwall 2705 (SP); Bananal, Serra da Bocaina, Ser- 
tao do Rio Vermelho, 1200 m, 7 Oct. 1949, Brade 
& Duarte 20119 (GH, RB); Campos do Jordao, Oct. 
1937, Campos Porto 3400 (GH, R), Oct. 1945, Lei- 
te 3660 (A); Mata da Boa Vista, 25 Nov. 1949, 
M. Kuhlmann 2176 (SP); Esta9ao Pilar do Sul, 28 
Sep. 1898, Edwall 4329 (SP); Sao Paulo, Agua 
Funda, Mata do Institute de Botanica, 25 Mar. 
1965, Rubens Faria 4 (SP); Jabaquara, 27 Sep. 
1945, Handro s.n. (SP); Campinas, Heiner 209 (s), 
Sep. 1900, Campos Novaes 214 (us), 18 Aug. 
1894, Campos Novaes 2960 (SP); Butantan, 25 
Sep. 1917, Hoehne 582 (M, SP). Bosque de Saude, 
9 Sep. 1922, Hoehne 7953 (SP, us); Hansa, 24 
Oct. 1928, Hoehne 23169 (SP, us); Jaragua, 10 
Oct. 1929, Hoehne 24405 (SP); Limeira, 30 Aug. 
1951, M. Kuhlmann 2714 (SP); Cordeiropolis, 19 
Nov. 1948, Levy 12 (SP); Araraquara, 28 Sep. 
1888, Lofgren 948 (SP); Piruibe, 1 Nov. 1891, Lof- 
gren & Cowan 1660 (c, SP); Sao Bernardo, 18 
Sep. 1908, Puttemans 6135 (SP); Semin. Espirito 
Santo-Santo Amaro, 18 Oct. 1942, Roth 356 (SP); 
Santa Branca, 21 Oct. 1901, Vert 6152 (SP). 

In his Flora Fluminensis of 1829, 15 Vellozo in- 
cluded two species of Brunfelsia under the genus 
Besleria. One of these, Besleria inodora, was de- 
scribed 2 years later as Franciscea pauciflora by 
Chamisso and Schlechtendal from a form of the 
plant with a glandular-pubescent calyx. This name 
was transferred to Brunfelsia in 1846 by Bentham, 
who made the combination B. pauciflora. The ear- 
lier epithet inodora of Vellozo cannot be used for 
this species since it is preoccupied by B. inodora 
Mart. (1847) (= B. americana). 

In 1846 Bentham published B. calycina, which 
is essentially a glabrous form of B. pauciflora. He 
cited three syntypes. One of these (Lund 755) be- 



15 As mentioned in the discussion of B. bonodora, 
there is evidence indicating that Vellozo's work was not 
offered for sale in 1825, which is the date that appears 
on the title page, but rather in September-November 
1829, which becomes the effective date of publication 
(Carauta, 1973). 



106 



FIELDIANA: BOTANY 



longs to B. obovata; another, collected by Baron 
de Colchester in Ilha Santa Catarina, has been 
chosen as the lectotype. The third collection (Ba- 
de s.n.\ also from Santa Catarina, has not been 
located. Schmidt (1864) reduced B. calycina to a 
variety of B. pauciflora in recognition of their 
similarity. 

Owing to its elegant beauty as a conservatory 
ornamental, B. pauciflora was introduced at an 
early date into European horticulture and become 
very popular in the mid- 1800s. Several new "spe- 
cies" under the genus name Franciscea were 
meagerly described from cultivated forms, includ- 
ing F. eximia, F. macrantha, and F. lindeniana. 
These names continue to persist in the horticul- 
tural literature in various combinations and have 
created a great deal of confusion in nomenclature. 
These forms are here considered as cultivars. 

Brunfelsia pauciflora is relatively uniform in 
morphological characters, perhaps reflecting the 
uniformity of its habitat in coastal, mostly low- 
elevation, rain forest. Some variation does exist, 
however, in the number and size of the flowers 
and in the size of the leaves. 

The presence and nature of the indument in B. 
pauciflora has been used to distinguish varieties 
and to segregate species. The type specimen of B. 
pauciflora has a glandular pubescence in the calyx 
and pedicels. This form is found primarily inland 
in the states of Sao Paulo and Parana, where pre- 
cipitation is less than on the coast. Brunfelsia ca- 
lycina and B. pauciflora var. calycina were based 
on the more widespread form with a glabrous and 
perhaps more coriaceous calyx, occurring in 
coastal forests from Rio de Janeiro south to Santa 
Catarina. 

Closer examination of the nature of the indu- 
ment shows a continuous range of intermediate 
forms of pubescence and glaberrity. The calyx 
may bear small, scattered, punctate dots; very 
short, scattered, glandular hairs; a well-developed, 
glandular villosity; or may be completely gla- 
brous. Indeed, some variability in the develop- 
ment and density of the hairs can be found within 
a single population. Furthermore, the hairs may 
be shed from the calyx as the fruit develops. Be- 
cause of the instability of the calyx indument as 
a taxonomic character in this species, I have not 
segregated the populations with glabrous calyces 
as distinct forms or subspecies, as has been done 
in the past. 

Smith and Downs (1966) pointed out two col- 
lections (Reitz 2743; Smith, Reitz & Klein 7936) 
from the mixed Araucaria forests of Santa Catar- 



ina as "possible hybrids" (parents unknown). 
These specimens differ in having a nonglandular 
villosity on the lower surface of the leaves at the 
midrib, whereas the calyces are completely gla- 
brous. These forms suggest possible introgression 
of genes from other species, such as B. cuneifolia 
or B. brasiliensis. Both of these species contain a 
similar, nonglandular pubescence and occur in the 
planalto region of Santa Catarina. 

Brunfelsia pauciflora occurs primarily on the 
Atlantic-facing slope of the Serra do Mar of 
southeastern Brazil from the State of Rio de Ja- 
neiro to Santa Catarina, including the Serra dos 
Orgaos, Serra de Mantiqueira, Serra de Parana- 
piacaba, and several lesser planaltos and morros. 
Altitudinally, it ranges from near sea level on the 
coastal plain to about 1500 m at Pico Caratuvain, 
Parana. This is exclusively the zone of the mata 
pluvial, the wet tropical rain forest with up to 
1600 mm of rainfall annually (Eiten, 1969). A few 
populations, as mentioned above, occur to the 
west in the mixed Araucaria forests of the plan- 
alto of Parana and Santa Catarina. 

This species is often found in shade along small 
streams and ravines, in humid primary and sec- 
ondary forests, and in other places where the soil 
is moist but well-drained. It apparently does not 
survive well when the forest is clear-cut. Brunfel- 
sia pauciflora flowers from September to Decem- 
ber, with the fruits appearing between January and 
March. Fruiting specimens are very scarce be- 
cause the large showy flowers are most often no- 
ticed by collectors. 

As an ornamental plant, B. pauciflora is one of 
the most important brunfelsias. It is commonly 
grown in conservatories in the temperate zone and 
frequently planted in tropical gardens. In limb di- 
ameter, the flowers are the largest of the genus, 
reaching 8 cm across. There is considerable vari- 
ation in color from deep, reddish purple to deep 
purple with a white spot or "eye" at the mouth. 
This spot is often surrounded by a blue-violet 
band that fades into the purple. The flowers fade 
rapidly with age to pale lavender or nearly white 
(but never pure white), with a concomitant in- 
crease of up to 1 cm in the diameter of the corolla. 

Several cultivars may now be recognized in 
cultivation. Cultivar "eximia" (B. eximia 
(Scheidw.) Bosse) has medium-sized leaves with 
an undulate margin and a densely glandular calyx 
with sharply acuminate teeth. Cultivar "macran- 
tha" (B. calycina var. macrantha (Lem.) L. H. 
Bailey and Raffill) is distinguished by large, leath- 
ery, plane leaves and by a large, glabrous calyx 



PLOWMAN: REVISION OF BRUNFELSIA 



107 



and a large corolla up to 8 cm across (Fig. 41). 
Cultivar "Lindeniana" (B. lindeniana (Planch.) 
G. Nichols) is a form with rosy-lilac flowers and 
slightly bullate leaves, according to Raffill (1928). 
Cultivar "floribunda" [B. calycina var. floribunda 
L. H. Bailey & Raffill] has smaller, violet flowers 
(to 3.7 cm across), shiny leaves, and is very free- 
flowering. 

Several of these cultivars represent very old 
clones, persisting in horticulture since their intro- 
duction in the last century. In an attempt to sta- 
bilize these names, I have designated as "repre- 
sentative collections" [for style of citation see 
Fig. 42 Eds.] material at the Royal Botanic Gar- 
dens at Kew that agrees with the original descrip- 
tions of the horticultural forms. A living collec- 
tion of Brunfelsia cultivars is maintained in the 
Temperate House of Kew, some of which proba- 
bly represent the original clones of these forms. 

22. Brunfelsia pilosa Plowman, Bot. Mus. Leafl. 
24(2): 42. 1974. Type: BRAZIL. Santa Ca- 
tarina: Mun. Sao Miguel d'Oeste, forest 
above Rio Reperiguacu, Peperi, ca. 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 

Gardens, Kew. 



108 



FIELDIANA: BOTANY 



- 




Revision of lirunl.-1-i.i (SoUnncne) 

Brunfelsia pauciflora (Cham. & Schlecht.) Benth. in 
DC. Prodr. JO: 199. 1846. 

cv* 'Lindeniana' 

Del. bv Timotlir 1'lowm.n I~ 1974 ""-"^^""i"' 



r.*i HERB. HORT. EOT. REG. KEW. 



'<**.' 

- 



Franciscea Lindeniana Planch. 
Belg. Hortio. 15:100. 1. 16. 1865. 





PLOWMAN: REVISION OF BRUNFELSIA 



109 



BRUNFELSIA pilosa Bowman 




FIG. 43. Brunfelsia pilosa. Reproduced courtesy of the Botanical Museum of Harvard University. 



110 



FIELDIANA: BOTANY 



Hatschbach 32869 (GH); *Municipio Adrianopol- 
is; Rio Pardo, prox. a Barra, Hatschbach 37876 
(MBM); *Municipio Balsa Nova; Campina da Cas- 
cavel, Hatschbach 39164 (GH); *Municipio Tiju- 
cas do Sul; Rincao, Hatschbach 40470 (F); *Mun- 
icipio Curitiba; Orto Guabirotuba, Hatschbach 
45415 (F); *Municipio Campina Grande do Sul; 
Belizario, Hatschbach & Guimardes 15286 
(MBM); *Municipio Jaguariaiava; Rio Samambaia, 
Hatschbach & Guimardes 25473 (MBM); *Muni- 
cipio Curitiba; Capanema, Kummrow 94 (MBM); 
*near Campo Largo, Lindeman & Haas 3248 (u, 
GH, MBM); *Parque Municipio do Iguacu (Muni- 
cipio Curitiba), Oliveira 100 (F). Rio Grande do 
Sul: Sao Leopoldo, 10 Sep. 1946, Henz 27246 
(MO, NY), 20 m, 17 Sep. 1946, Henz 35305 (s); 
Mun. Rio Pardo, Fazenda Horticola, 70 m, Oct. 
1923, Jurgens 19 (B); Colonia Santo Angelo, 4 
Feb. 1893, C. Lindman A-597-b (s), Sep. 1900, 
Schwarzer 50 (L, s); Silveira Martins, 20 Mar. 
1893, C. Lindman A-597-c (s); Rio dos Sinos, 8 
Nov. 1949, Rambo 44295 (BR, L); Butterberg pro- 
pe Montenegro, 13 Nov. 1950, Rambo 49144 (GH, 
p, w); without locality, 1833, Gaudichaud 705 (P), 
1835, Isabelle 6 (G); *Municipio Sta. Barbara; 
Cruz Alta, Lindeman et al. 8233 (u). Santa Ca- 
tarina: Mun. 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- 
eter. 

Brunfelsia pilosa ranges from the Brazilian 
state of Sao Paulo south to Rio Grande do Sul and 
extreme northeastern Argentina (Misiones). It has 
a fairly wide altitudinal range, occurring from 
near sea level to 1100 m elevation. In the central 
part of its range, where it occurs most commonly, 
this species seems to be associated with the for- 
merly extensive Araucaria forests found in the 
south Brazilian planalto. 

In contrast to B. australis, which may become 
a small tree, B. pilosa grows as a weak shrub with 
slender, horizontal, arching branches. It occurs 
primarily in the understory of primary and sec- 
ondary woods, as well as in thickets and along 
watercourses. Flowering takes place from October 
to December with fruits appearing from February 
to May. 

Owing to its attractive flowers, this shrub is cul- 
tivated in tropical gardens, although not as exten- 
sively as B. australis. Brunfelsia pilosa appeared 
in the Florida nursery trade in the mid-1970s and 
is increasing in importance as an ornamental. 

In its native area, B. pilosa is known by a va- 
riety of common names: flor de primavera, pri- 
mavera, sempre triste, and manacd (Brazil); azuc- 
ena silvestre, azucena del monte, and Jazmin del 
Paraguay (Argentina). 

23. Brunfelsia rupestris Plowman, Fieldiana, 
Bot. n. s. 8: 7. 1981. Type: BRAZIL. Minas 
Gerais: Estrada Diamantina a Corinto ate 20 
km. Campo rupestre e cerrado. Arbusto ate 2 
m, flores roxas. 12 Jan. 1976, Shepherd, de 



PLOWMAN: REVISION OF BRUNFELSIA 



111 



BRUNFELSIA 

rupest ris 

Plowman 




5 cm. 



FIG. 44. Brunfelsia rupestris. Reproduced courtesy of the Field Museum of Natural History. 



Andrade, Konoshita & Tamashiro 3935 (ho- 
lotype, UEC 11912; isotypes, F 1877095, NY, 
UEC 11912A). Figure 44. 

Much-branched shrub 1-2 m tall. Bark on 
branches yellowish to reddish brown, cracked lon- 
gitudinally and transversely, exfoliating in thin 



flakes. Branch lets about 2 mm in diameter, gla- 
brous or puberulent. Leaves usually congested 
near branch tips, short petiolate, blade 10-40 mm 
long, 6-17 mm wide, lanceolate to oblong, api- 
cally acute, sometimes blunt or emarginate, 
strongly revolute at margin, basally cuneate, gla- 
brous above, puberulent beneath with short 



112 



FIELDIANA: BOTANY 



curved glandular or eglandular hairs, shiny, me- 
dium green above, dull beneath, drying ochra- 
ceous brown, thickly coriaceous, the lateral 
nerves 4-5, straight, scarcely distinct; petiole 1- 
3 mm long, glabrous. Inflorescence terminal on 
current season's branchlets, short pedunculate, 
with 2-5(8) flowers. Flowers deep violet with a 
white "eye" at orifice, fading to lavender with 
age. Peduncle 1-5 mm long, short-branched, the 
branches articulating with the pedicels, persistent, 
puberulent. Bracts 16 mm long, linear-lanceolate 
or cymbiform, sparsely villous, ciliolate at mar- 
gin, caducous. Pedicel 4-12 mm long, 1-1.5 mm 
in diameter, glabrous. Calyx 10-15 mm long, 3- 
6 mm in diameter, tubular-campanulate, terete, 
glabrous, light green, teeth 2-3 mm long, sub- 
equal, ovate to apically acute, the tip itself some- 
times blunt or truncate. Corolla tube orifice 2-3 
mm in diameter; limb 15-22 mm in diameter, 
spreading, slightly thickened at mouth of tube, 
lobes 5-10 mm long, subequal, rounded. Stamens 
inserted in upper part of corolla tube; filaments 
strap-shaped, the anterior pair 2-3 mm long, api- 
cally incurved, included, the posterior pair 1-2 
mm long; anthers 1.5 mm in diameter, orbicular- 
reniform. Ovary 1.5 mm long, ovoid; stigma 1 
mm long, briefly bifid. Capsule enclosed by per- 
sistent calyx, 16 mm long, 14 mm in diameter, 
ovoid to globose, apiculate at apex, smooth, peri- 
carp thin, crustaceous, dry at maturity. Seeds 5 
mm long, 2.5-3 mm in diameter, oblong-ellipsoid, 
somewhat angular, brown, reticulate-pitted. Em- 
bryo 4 mm long, straight; cotyledons 2 mm long, 
ovate. 

DISTRIBUTION Restricted to the campo rupes- 
tre community in the Serra do Espinhaso, Minas 
Gerais, Brazil. See Figure 41. 

SPECIMENS EXAMINED BRAZIL. Minas Ger- 
ais: Serra do Espinhaso, Municipio Diamantina: 
Guinda, 5 Nov. 1937, Mello Barreto 9479 (F, RB), 
1300 m, 14 Nov. 1971, Hatschbach & Pelanda 
27941 (GH); ca. 25 km SW of Diamantina on road 
to Gouveia, 1300 m, 16 Jan. 1969, Irwin et al. 
22077 (F, MO, NY, UB); ca. 12 km SW of Diaman- 
tina, 1350 m, 23 Jan. 1969, Irwin et al. 22465 
(NY, UB); Serra do Cruzeiro, 5 Nov. 1979, V.F. 
Ferreira et al. 887 (RB); *Municipio Gouveia; Da- 
tas, Hatschbach and Zelma 50296 (F). 

Brunfelsia rupestris is a very distinct species 
that is restricted to the compos rupestres at higher 
elevations in the Serra do Espinha?o of central 
Minas Gerais. It is most closely related to, and 



probably derived from, the widespread and poly- 
morphic species B. brasiliensis. Brunfelsia rupes- 
tris differs in having the leaves more congested, 
thickly coriaceous, strongly revolute at the mar- 
gins, sparsely pubescent, and with only four to 
five lateral nerves and in having glabrous pedicels 
and calyces. 

24. Brunfelsia uniflora (Pohl) D. Don, Edin- 
burgh New Philos. J. 86. July. 1829. Figure 
45. 

Franciscea uniflora Pohl, PI. Bras. Ic. 1: 2, t. 1. 1826. 

Type: BRAZIL. Rio de Janeiro: habitat in um- 

brosis inter frutices in via a Sumidorio ad Gov- 

erno et ad Rio Parahybuna, Sep.-Oct. 1818, Pohl 

[9] (holotype, w; isotype, PR). 
Franciscea hopeana Hook., Bot. Mag. 55: t. 2829. 

1828. Type: Hooker, Bot. Mag. 55: t. 2829. 

1828. 
Brunfelsia hopeana (Hook.) Benth. in DC., Prodr. 10: 

200. 1846. 
Brunfelsia hopeana var. pubescens Benth. in DC., 

Prodr. 10: 200. 1846. Type: TRINIDAD. In in- 

sula Trinitatis montibus Monos Bocas, without 

date, Lockhart 197 (holotype, K; isotype, GOET). 
Brunfelsia uniflora var. pubescens (Benth.) R. E. D. 

Baker & N. W. Simmonds in R. O. Williams, Fl. 

Trinidad and Tobago 2(4): 270. 1953. 
Franciscea mutabilis H. Jacq., Ann. Fl. Pomone: 285. 

Jun. 1842. Type: Jacq., Ann. Fl. Pomone: 285. 

Jun. 1842. 
Brunfelsia mutabilis (Jacq.) Vilm., Rev. Hort. ser. 2, 

1: 261. Aug. 1842. 

Shrub 0.5-3 m tall with diffuse, spreading 
branches, often branched from near base. Branch- 
es slender, terete, becoming somewhat knobby, 
naked beneath, glabrous. Bark grayish brown, 
shiny. Branchlets slender, glabrous or pubescent. 
Leaves scattered along the branchlets, highly 
variable in size and shape often in the same in- 
dividual, blade 2.5-8.0 cm long, 1.0-4.0 cm wide, 
oblong-elliptic, lanceolate or obovate, acute to 
acuminate at apex, narrowed or cuneate at base, 
sparsely glandular or variably pubescent beneath 
at the nerves, or glabrous, dark to light green 
above, paler green beneath, unusually dull, mem- 
branaceous to subcoriaceous, deciduous, lateral 
nerves 5-8, spreading, straight or arching; petiole 
short, 1-5 mm long, pubescent or glabrous. Inflo- 
rescence 1 -flowered, terminal, produced at tips of 
new shoots, sessile, rarely short pedunculate. Pe- 
duncle 1-6 mm long, slender, terete, glabrous or 
glandular pubescent, rarely villous, persistent. 
Bracts 1-3, 1-5 mm long, linear-lanceolate, cili- 
olate, pubescent at midrib, light green, caducous. 
Pedicel 1-6 mm long, erect, glabrous. Calyx 8- 



PLOWMAN: REVISION OF BRUNFELSIA 



113 




114 



FIELDIANA: BOTANY 



20 mm long, 3-7 mm in diameter, tubular to tu- 
bular-campanulate, 16 narrowed at base, glabrous, 
rarely with sparse glandular hairs, pale green, 
membranaceous, teeth 1-3 mm, erect or incum- 
bent, triangular-ovate, acute to acuminate; calyx 
in fruit coriaceous, striately veined, dark green. 
Corolla tube 13-25 mm long, 1-2 mm diameter, 
1-2 times as long as calyx, pentagonal in section, 
glabrous or sparsely glandular; limb 15-30 mm in 
diameter, spreading, lobes equal, broadly rounded, 
at times emarginate, overlapping at the lateral 
margins, narrowed at base. Stamens included in 
upper part of corolla tube; filaments slender, upper 
pair 3-5 mm long, lower pair 2-3 mm long; an- 
thers about 1 mm in diameter, orbicular-reniform. 
Ovary 2-3 mm long, 1-1.5 mm in diameter, nar- 
row conical to ovoid; style 15-23 mm long, slen- 
der, glabrous; stigma briefly bifid, lobes unequal, 
obtuse. Capsule nearly enclosed by persistent ca- 
lyx, 7-8 mm long, 8-18 mm in diameter, subglo- 
bose to ovoid, smooth, shiny, dark green, pericarp 
0.5 mm thick, tardily dehiscent. Seeds 10-20, 3- 
5 mm long, 2-3 mm in diameter, variable in 
shape, ovoid-ellipsoid, somewhat angulate, dark 
reddish brown to almost black, reticulate-pitted. 
Embryo 2-4 mm long, curved slightly; cotyle- 
dons ovate. 

DISTRIBUTION Venezuela, Trinidad, Guyana, 
Brazil, Bolivia, Argentina. See Figure 46. 

SPECIMENS EXAMINED VENEZUELA. Boli- 
var: Upata, 1839-1841, Otto 1003 (w). Cara- 
bobo: *Entre Trincheras y El Cambur, Guevara 
2276 (F, NY). Isla Margarita: El Valle, 300 m, 3 
Aug. 1903, Johnston 71 (GH), 2 Aug. 1901, John- 
ston & Miller 265 (GH, NY); without locality, *Fa- 
cultad Ciencias Forestales s.n. (NY). Guarico: 
Selvas de Tamanaco, cerca del Cementerio de 
Paso Real, 11-12 Jun. 1966, Aristeguieta 6153 
(NY, YEN), 21 Sep. 1968, Plowman 1911 (ECON). 
Without state: Los Cedros, Aug. 1953, Gines 
3605 (us). TRINIDAD. Monos Island, 100 ft, 
Apr. 1936, Graf 10 (NY). GUYANA. Western ex- 



16 After completing his thesis, Powman described B. 
clandestine based on materials he had originally includ- 
ed in B. uniflora. The reference to the tubular-campan- 
ulate calyx in this description may derive from "fl. clan- 
destina." See discussion of B. clandestine! N.H. 



tremity of Kanuku Mountains, in drainage of Tak- 
utu River, 200 m, 4-22 Mar. 1938, AC. Smith 
3163 (A, B, F, G, LE, MO, NY, p, us, w). BRAZIL. 
Bahia: Espigao Mestre, 6 km S of Cocos, 520 m, 
16 Mar. 1972, Anderson, Steiber & Kirkbride 
37048 (wis); Itambe, Salobinho, basin of Rio Par- 
do, 14 Nov. 1942, Froes 12681/46; Morro do 
Chapeu, ca. 7 km S of town of Morro do Chapeu, 
1,125 m, 17 Feb. 1971, Irwin, Harley & Smith 
32406 (wis); Caatinga bei Tamburi, Oct. 1906, 
Ule 7061 (K, L); Serra Jacobina, St. Thome, Ponco 
d'Areia, 1843, Blanchet 3862 (BR, G, P); Blanchet 
2616 (P); 1843, Blanchet s.n. (c, G, w); without 
locality, Jun. 1844, Blanchet s.n. (G), 1847, Blan- 
chet s.n. (G), Martius s.n. (L); *Monte Santo, Har- 
ley et al. 16432 (F, K); *Feira de Santana, Serra 
Sao Jose, Noblick 2038 (F); *Barra da Choca, 
Santos, T. Soares dos 2511 (F). Ceara: Horto Flo- 
restal de Ubajara, "Sitio Buriti," 5 Jun. 1942, Ai- 
res do Nascimento 19 (RB); Vi9osa de Ceara, 
1860, Allemao 1249 (P, R); Mun. Sao Gonsalo, 
near Paracuru, 10 m, 7 Jan. 1945, Cutler 8271 
(MO, us); Guaramiranga, Sao Salvador, 24 Aug. 
1908, Ducke 1588 (MG); Fortaleza, tabuleiro alem 
de Aldeota, 5 Feb. 1957, Ducke 2954 (MG, R, RB); 
Varzea de Vaca, Feb. 1839, Gardner 2432 (BM, 
K); Serra do Bezouro, 20 Jan. 1958, Guedes 472 
(MG); without locality, Allemao & Cysneiro 1250 
(R), 1840, Gardner 1798 (CGE, F, G, L, NY, p, us, 
w); *Municipio de Sao Benedito; Sitio Castelo, 
Serra da Ipiapaba, Bezerra 334 (F); *Sitio Sao An- 
tonio, Serra de Maruoca, Fernandes, A., s.n. (F); 
*Camocim, Cafundo, S. J. Filho 8 (F). Espirito 
Santo: Bananal, 1924, Freire 14 (R, us); Gorto- 
cazes, 10 Nov. 1943, J. G. Kuhlmann s.n. (RB). 
Goias: Luziana, 18 Sep. 1967, Heringer 11549 
(us). Minas Gerais: Vi9osa, state agricultural 
school, 20 Feb. 1959, Irwin 2674 (F, NY, R, us), 
22 Feb. 1959, Irwin 2701 (F, NY, R, uc); 19 Nov. 
1928, J. G. Kuhlmann s.n. (RB), 2 Apr. 1935, /. 
G. Kuhlmann s.n. (RB); oxcart road to Sao Miguel, 
about km 4, 685 m, 2 Apr. 1930, Mexia 4551 (GH, 
us); road to Barrosa, about km 6, 700 m, 15 Oct. 
1930, Mexia 5182 (A, BH, BM, F, c, GB, GH, MICH, 
MO, NY, PH, s, uc, us); 1827, Pr. von Nevwied s.n. 
(BR). Caldas, 18 Oct. 1854, Lindberg 203 (BR, s); 
25 Sep. 1873, Mosen 673 (c, LD, p, R, s), 1844- 
1865, Regnell 1-374 (BR, c, GH, GOET, LE, M, NY, 



FIG. 45. Brunfelsia uniflora (originally as Franciscea uniflora). From Pohl, J. E., Plantarum Brasiliae Icones et 

Descriptiones (1826). 



PLOWMAN: REVISION OF BRUNFELSIA 



115 




FIG. 46. Distribution of Brunfelsia uniflora (solid circles). 



p, R, s, us); Cabo Verde, Oct. 1876, Barbosa Ro- 
drigues 261 (RB); Parahybuna, Oct. 1840, Gard- 
ner 5062 (EM, CGE, K, p, us, w); without locality, 
1858, Weddell s.n. (G), 1845, Widgren s.n. (BR, 
LD, s). Pernambuco: Isla de Itamarica, Dec. 
1839, Gardner 1095 (BM, CGE, G, NY, p, w); Per- 
nambuco, May 1838, Gardner s.n. (BM, K); Cabo 
Agostinho, 22 May 1952, D. de A. Lima 52-1039 
(R); Olinda, 20 Apr. 1924, Picket 670 (SP); Tapera, 
18 Dec. 1929, Pickel 2200 (us). Rio Grande do 
Norte: Monte Alegre, 26 Jan. 1961, Castellanos 
23004 (R); near Natal, Mar. 1914, Dawe s.n. (K). 
Rio de Janeiro: Corcovado, Sep. 1833, Lus- 
chnath s.n. (LE), Nov. 1833, Luschnath s.n. (BR, 
LE), Oct.-Nov. 1832, Riedel 1112 (LE, NY, p, us, 
w); Restinga do Leblon, 5 Feb. 1948, Macedo s.n. 
(RB); Parahyba, Oct. 1840, Gardner 5566 (CGE, 
GH, K, w); Sobral prope Parahyba do Sul, Nov. 



1881, Schwacke 3311 (GOET, R, us); Estrada de 
Minas, Miers 4721 (us); Serra d'Estrella, 1844, 
Weddell s.n. (NY, p); without locality, 1867, Gla- 
ziou 357 (BR); *Municipio Sao Sabastiao do Alto; 
Fazenda Bahia, Araujo 968 (F); *Municipio Silva 
Jardim; Reserva Biologica 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); 



116 



FIELDIANA: BOTANY 



TABLE 7. Flowering and fruiting times of Brunfelsia uniflora in different parts of its geographical range. 



Region 



Flowering 



Fruiting 



Venezuela 
Northeastern Brazil 
Southeastern Brazil 
Bolivia 



April-August 
December- April 
October-November 
September-December 



August 

February-May 
February-April 
Unknown 



20 km N of Boa Vista, 1 Feb. 1969, Prance et al. 
9553 (wis). Sao Paulo: Botucatu, Edwall 6133 
(SP); Campinas, Heiner s.n. (s); 2 May 1918, 
Campos Novaes 7756 (us); Sao Jose do Rio Par- 
do, 1 Oct. 1889, Lofgren & Edwall 1435 (SP, us); 
Xiririca, 16 Oct. 1894, Lofgren & Edwall 2742 
(SP, us); BR-2a km 10 antes de Jacupiranga, 15 
Oct. 1961, Pabst & Pereira 6023 (F, R, us); Ban- 
anal, Oct. 1833, Riedel s.n. (LE, us); pr. Sao Car- 
los, Jan. 1834, Riedel s.n. (LE); *SW de Jundiai, 
Serra de Japi, H. F. Leitdo Filho et al. 3219 (F). 
Without state: Pohl 264 (PR), Pohl s.n. (M). BO- 
LIVIA. Cochabamba: Prov. Ayopaya, Sailapata, 
2,700 m, Dec. 1935, Cardenas 3380 (GH); Prov. 
Cochabamba, Siberia, 3,300 m, Jul. 1955, Car- 
denas 5216 (L, us); Carrasco, Jatum-Pino, 3300 
m, Sep. 1961, Cardenas 5952 (K, us). Santa 
Cruz: Comarapa, 2800 m, 20 Oct. 1928, Stein- 
bach 8345 (A, BM, GH, K, MO, NY, s, us); *W of 
Comarapa on Carretera Fundamental 4, Davidson 
3846 (F). Without department: Padcaya-Motovi, 
2400 m, 24 Sep. 1927, Troll 240 (B, M). ARGEN- 
TINA. Jujuy: Dep. Ledesma, Serrama de Cali- 
legua, 1500 m, 18 Oct. 1963, Fabris 4502 (M). 
Salta: Oran, cerca del Achiral mas abajo de San 
Andres, 17-24 Sep. 1873, Lorentz & Hieronymus 
258 (BR, G, LE, p). 

Brunfelsia uniflora was first described by Pohl 
in the genus Franciscea. Pohl collected the type 
in the state of Rio de Janeiro in Brazil. The ho- 
lotype, bearing the number 9, is preserved at Vi- 
enna (w); a probable isotype with the number 256 
is found in Prague (PR). In 1828, Hooker de- 
scribed another species, F. hopeana, based on a 
living plant sent from Brazil, which, according to 
Hooker, differed from F. uniflora in having a 
shorter corolla tube. D. Don united Franciscea 
with the older genus Brunfelsia in 1829, making 
the correct combination B. uniflora. However, 
Bentham, in his monograph of 1846, used Hook- 
er's specific epithet in the combination B. ho- 
peana. This name is still widely, though incor- 
rectly, used for this species, particularly in the 



horticultural and pharmaceutical literature [see 
Plowman, 1977 Eds.]. 

Brunfelsia uniflora is the most widespread and 
variable of the South American species of the ge- 
nus. It encompasses a heterogeneous complex of 
forms found throughout eastern Brazil with dis- 
junct populations in Venezuela and Bolivia. While 
showing some differentiation in response to dif- 
ferent habitats and as a result of geographical iso- 
lation, the variation is considered to fall within 
the normal range of a single polymorphic species. 
The characters and nature of the variability will 
be discussed according to geographical regions. 

Brunfelsia uniflora is distinguished from related 
species by its smaller, solitary flowers (tube 13- 
25 mm long), the more or less tubular calyx, and 
the short (1-3 mm) calyx teeth. It is highly vari- 
able in the shape and size of the leaves, in the 
length of the calyx, and in degree and location of 
pubescence. 

This species grows under diverse environmen- 
tal conditions throughout its range, which is, in 
part, responsible for the different phenotypes that 
are encountered in different areas. It thrives in 
wet, montane rain forests in Rio de Janeiro and 
in the high Bolivian yungas. It has also become 
adapted to the drier regions of Minas Gerais, Sao 
Paulo, and Ceara, as well as the cerrado forma- 
tions of Goias and northeastern Brazil. It occurs 
sporadically as a component of the understory in 
primary and secondary forests and persists in cut- 
over areas (capoeiras). It grows in different types 
of soils, including red clay, sand, and moist hu- 
mus. Altitudinally, B. uniflora ranges from sea 
level to 3300 m in the Andes. 

Flowering and fruiting in B. uniflora occur at 
different times of the year in different parts of its 
range. These are indicated in Table 7. 

In the coastal rain forest of Rio de Janeiro and 
adjacent states, B. uniflora assumes its "typical" 
form, exemplified by the type collection and orig- 
inal description. These plants are very glabrous 
with obovate-elliptic leaves. The leaves tend to be 
larger and shinier than in other areas. 



PLOWMAN: REVISION OF BRUNFELSIA 



117 



Further inland, in the drier highlands of Minas 
Gerais, there is a tendency toward smaller and 
more pubescent leaves. In addition the flowers 
may be short-stalked and the calyx longer and 
narrower in relation to the corolla tube. Popula- 
tions showing this trend extend southward in the 
interior of Sao Paulo and northward to central 
Brazil and along the coast to the dry northeast of 
the country, stopping abruptly in the state of 
Ceara. Here the plant may become deciduous, 
flowering at the beginning of the wet season. 

Forms similar to those in drier areas of Brazil 
appear again along the coast of Venezuela, in Isla 
Margarita and Monos Island (Trinidad). In these 
northern populations, the young twigs and pedun- 
cle, if present, are frequently densely pubescent 
[perhaps because of environmental conditions; see 
Knapp, 1989 Eds.]. Bentham tentatively named 
these plants B. hopeana var. pubescens pending 
observation of the corolla, which was missing in 
his specimen. 

When pubescence and calyx length in this form 
are considered across its entire area, the variation 
appears more or less continuous, often showing 
greater differences in adjacent populations than in 
those separated by thousands of kilometers. The 
development of pubescence, which may be glan- 
dular or nonglandular, is not a good character in 
this species. The relative length of the calyx is 
also variable, sometimes appearing tubular-cam- 
panulate, sometimes long tubular, with many in- 
termediates. [Later, Plowman differentiated B. 
claudestina, with tubular-campanulate calyx, from 
B. uniflora, with "narrowly tubular calyx." See 
discussion of B. claudestina. N.H.] In contrast 
to past treatments of B. uniflora, these characters 
are not considered here for segregating varieties 
or subspecies. 

In southeastern Guyana and adjacent Brazil 
(Roraima) and southern Venezuela, there occurs 
another pubescent form of B. uniflora but with 
oblong-ovate leaves, slightly curved, tubular caly- 
ces, and hispidulous branchlets. Known only from 
four collections (A. C. Smith 3163, Ule 7715, Otto 
1003, Prance et al. 9553), these distinctive pop- 
ulations skirt the eastern part of Guyana High- 
lands. Their uniformity suggests that they have 
been isolated from other populations of B. uniflo- 
ra for considerable time. 

A final component of B. uniflora occurs along 
the eastern edge of the Bolivian altiplano at about 
2500 m altitude, extending south into northwest- 
ern Argentina. These populations closely resem- 
ble plants growing in the coastal rain forest of Rio 



de Janeiro. They differ in having shiny, coria- 
ceous leaves and longer pedicels. The Bolivian 
populations grow in very wet, montane cloud for- 
ests with Podocarpus and Alnus and are morpho- 
logically homogenous. 

In southern Brazil, from the state of Sao Paulo 
south to northeastern Argentina, several species 
occur that may superficially resemble, and have 
been confused with, B. uniflora. Two of these, B. 
australis and B. pilosa, are closely related to B. 
uniflora and probably have been derived from it 
through geographical isolation in the past. They 
form, in fact, a southern component of what may 
be referred to as the B. uniflora complex. 

Both B. pilosa and B. australis are distin- 
guished from B. uniflora by a number of constant 
morphological characters that are listed under 
those species. Few or no intermediates or putative 
hybrids have been found, although their ranges 
overlap somewhat in Sao Paulo (B. uniflora with 
B. pilosa) and in Rio Grande do Sul (B. pilosa 
with B. australis). Schmidt (1864) considered B. 
australis a variety of B. uniflora, but he did not 
see a full range of specimens of either species and 
included material of B. guianensis and B. pilosa. 

In the states of Parana and Santa Catarina, two 
additional species occur that may be confused 
with the B. uniflora complex. Smith and Downs 
(1966), in the Flora Ilustrada Catarinense, as- 
signed specimens of both B. pilosa and B. bras- 
iliensis subsp. macrocalyx to what they consid- 
ered B. uniflora. They further confused B. cunei- 
folia with B. brasiliensis . These workers appar- 
ently did not examine more typical material of B. 
uniflora and B. brasiliensis from farther north. 

The breakdown of sharp specific boundaries in 
these species (especially B. pilosa, B. cuneifolia, 
and B. brasiliensis) suggests a certain amount of 
hybridization among them in the past as they 
came together in the planalto of southern Brazil. 
Subsequent convergence in several characters 
may also be attributed to a relatively uniform en- 
vironment in the zone of the Araucaria forests. 
Similarities among these species are particularly 
noticeable in the size and shape of the leaves, the 
form and location of pubescence, and in the re- 
duction in the number (often to 1) of flowers per 
inflorescence. Field studies in which additional 
collections are correlated with local ecological 
conditions should prove very useful in under- 
standing their specific relationships. 

Brunfelsia uniflora is the source of manaca 
root, which is widely used in Brazil in the treat- 
ment of rheumatism and syphilis. It is known by 



118 



FIELDIANA: BOTANY 



a number of common names that, along with an 
account of the uses of the drug, are discussed in 
Plowman (1977). 



Dubious and Excluded Species 

Brunfelsia bahiensis Benth. in DC., Prodr. 10: 
590. 1846. Type: BRAZIL. Bahia: without 
locality, 1834, Blanchet 1509 (holotype, c- 
DC, non vidi; photographs of holotype, 
F-645866, NY; isotypes, BM, G, p). 

Schmidt considered B. bahiensis a dubious spe- 
cies of Brunfelsia, and I am inclined to follow 
suit. It is known only from the type collection. 
One specimen from Minas Gerais (Pr. von Nev- 
wied s.n., 1816, BR) resembles the type but in fact 
seems better placed in B. bonodora. Another 
specimen from Bahia (Blanchet s.n., 1852, G) is 
also similar but is considered a form of B. hy- 
drangeiformis. Until additional material becomes 
available, the true affinities of this species must 
remain in doubt. 

Brunfelsia nyctaginoides Standl., Field Mus. Nat. 
Hist. Bot. ser. 22 (1): 47. 1940. Type: MEX- 
ICO. Chiapas: Monte Tacana, 1000-2000 m, 
Aug. 1938, Matuda 2411 (holotype, MICH; 
isotypes, G, K, MICH, MO, NY, us). 

This anomalous plant is known only from a few 
collections on high-elevation, volcanic peaks near 
the Guatemala-Mexico border. In the original de- 
scription, Standley (1940) expressed doubt as to 
the generic placement of this plant. He later rec- 
ognized its intermediate position between Bro- 
wallia, Streptosolen, and Brunfelsia (Standley and 
Steyermark, 1940). 

It must, however, be excluded from Brunfelsia 
on the basis of having a broadly lamellate stigma 
and plicate corolla lobes that are bright scarlet in 
color, resembling the corolla of Streptosolen ja- 
mesonii. Streptosolen has a distinctly twisted co- 
rolla tube that is lacking in the present plant. Its 
proper generic disposition must await a thorough 
evaluation of related genera of the Salpiglossi- 
deae. [It has now been placed in the monotypic 
genus Plowmania Hunz. (Hunziker and Subils, 
1986). Eds.] 

Nomina Nuda 

The following names, mostly of horticultural 
origin, have no taxonomic standing. 



Franciscea angusta Heynh., Nom. Bot. Hort. ed. 

3. 551. 1850. 
Brunfelsia angusta Gentil., PI. Cult. Serres Jard. 

Bot. Brux. 40. 1907. 
Brunfelsia falcata Regel, Cat. PI. Hort. Aksakov. 

21. 1860. 
Brunfelsia gracilis Heynh., Nom. Bot. Hort. ed. 

2. 279. 1846. 
Franciscea gracilis W. Baxt. in Loud., Hort. Brit. 

Suppl. 3. 551. 1850. 
Brunfelsia latifolia Steud., Nom. Bot. ed. 2. 231. 

1840. Non B. latifolia (Pohl) Benth. 
Brunfelsia lockhartii Heynh., Nom. Bot. Hort. ed. 

2. 79. 1846. 
Brunfelsia magnifica Gentil., PI. Cult. Serres Jard. 

Bot. Brux. 40. 1907. 
Brunfelsia montana Lodd., Cat. PI. ed. 12. 4. 

1820. 

Brunfelsia multiflora Regel, Cat. PI. Hort. Aksa- 
kov. 21. 1860. 
Brunfelsia schomburgkiana Klotsch ex Schom- 

burgk, Versuch. Fauna Fl. Br. Guy. 1155. 

1848. 
Brunfelsia sieberi Regel., Cat. PI. Hort. Aksakov. 

21. 1860. 
Brunfelsia spinosa Jacq., Enum. Syst. PI. 14. 

1760. 
Brunfelsia spruceana J.A. Schmidt in Mart., Fl. 

Bras. 8(1): 258. 1864. A nomen nudum 

placed in the synonymy of B. maritima 

Benth. by Schmidt. 
Franciscea pohliana Heynh., Nom. Bot. Hort. ed. 

2. 1846. 
Mania opifera Lacerda ex J. A. Schmidt in Mart., 

Fl. Bras. 8(1): 261. 1864. A nomen nudum 

placed in the synonymy of B. hopeana 

(Hook.) Benth. by Schmidt. 



XI. Editors' Acknowledgments 

It is impossible for us to acknowledge with cer- 
tainty all those whom Timothy Plowman would 
have wished to thank for their help. Because this 
work is based primarily on his Ph.D. dissertation, 
the original acknowledgments to individuals are 
reprinted here. We extend our apologies to anyone 
who should be mentioned but is not. 

As editors, we would like to add our own ac- 
knowledgments, especially to Dr. Michael Dillon 
of the Field Museum of Natural History, Chicago, 
who originally suggested taking on the task of 
preparing this revision for publication and who 



PLOWMAN: REVISION OF BRUNFELSIA 



119 



patiently answered all our queries and requests for 
advice. Nancy Hensold and Christine Niezgoda, 
also of the Field Museum, gave us invaluable aid 
in searching Plowman's notes and other archival 
material and provided additional data. Several in- 
dividuals and organizations kindly gave us per- 
mission to reproduce previously published illus- 
trations: Curtis' s Botanical Magazine; the Field 
Museum of Natural History, Chicago; the Botan- 
ical Museum, Harvard University; Christabel 
King; and the Linnean Society, London. The 
drawing of Brunfelsia amazonica was prepared by 
Margaret Tebbs, and all the distribution maps 
were prepared by Malcolm Penn. Kai Vollesen of 
the Royal Botanic Gardens, Kew, provided the 
photograph of B. obovata cv. Lindeniana, and Dr. 
Jim Mallet of University College, London, pro- 
vided the photograph of Methona larvae. The 
Photographic Unit at the Natural History Museum 
produced several of the figures published here, 
and Marian Short checked the manuscript. 



XII. Plowman's Acknowledgments 

The following acknowledgments are from Tim- 
othy Plowman 's dissertation: 

Many persons have generously given to me 
their time and assistance during the course of this 
study. I gratefully acknowledge their help, which 
has both enhanced the quality and facilitated the 
completion of this work. 

I would especially like to express my sincere 
gratitude to Prof. Richard Evans Schultes, who 
first suggested the genus Brunfelsia as a problem 
in ethnopharmacology and provided continued 
support and encouragement throughout the study. 
His profound interest in the plants and the people 
of South America demonstrated to me the impor- 
tance of field studies in ethnobotanical and taxo- 
nomic research. 

During the course of my field work, many in- 
dividuals graciously offered their facilities and 
hospitality, food, and lodging, and generally made 
my travels more pleasant and scientifically re- 
warding. I am particularly indebted to the follow- 
ing persons: Dr. Leandro Aristeguieta in Venezue- 
la; Drs. Jesus Idrobo, Eduardo Mora Osejo, and 
Alvaro Fernandez, Sr. Rafael Wandurraga, and Sr. 
Pedro Juajibioy Chindoy in Colombia; Dr. Ramon 
Ferreyra and Sr. Fernando Tina in Peru; Dr. Julian 
Camara Hernandez and Drs. Antonio and Carmen 



Krapovickas in Argentina; Dr. Roberto Klein and 
Sr. Dimitri Sucre in Brazil; Drs. George R. Proc- 
tor and C. D. Adams in Jamaica; and Dr. Roy 
Woodbury in Puerto Rico. 

The following made many useful suggestions 
and criticisms and assisted in solving taxonomic 
problems: Dr. Cleofe Calderon, Dr. Jose Cuatre- 
casas, Dr. William T Gillis, Dr. Leslie A. Garay, 
Mr. William Grime, Prof. Richard A. Howard, Mr. 
Tom E. Lockwood, Prof. Robert F. Raffauf, Prof. 
Reed C. Rollins, Dr. Velva E. Rudd, Dr. Elizabeth 
A. Shaw, Dr. Bernice G. Schubert, Dr. Lyman B. 
Smith, Dr. Thomas R. Soderstrom, Dr. William T. 
Stearn, Prof. Rolla M. Tryon, Prof. Carroll E. 
Wood, and Dr. John J. Wurdack. 

Technical assistance was provided by Drs. 
Alice F. Tryon, Umesh Banerjee, and Ihsan 
Al-Shehbaz and Mr. Gilbert Pierce. 

Dr. Fred Barkley and Mr. Eugene Courtney at 
the Botanical Research Station in Woburn, Mas- 
sachusetts, Mr. Al Fordham of the Arnold Arbo- 
retum, and Mr. Conrad Smith of Harvard Univer- 
sity provided much help with the care of living 
material. 

Bibliographic services and many kindnesses 
were provided by the staff of the libraries of the 
Gray Herbarium and Arnold Arboretum, particu- 
larly Mrs. Patricia Hall, and by Mrs. Mary Gau- 
det, Mrs. Lilian Hanscom, and Miss Esther Reyn- 
olds of the Botanical Museum. 

I would also like to thank Ms. Lynda Bates and 
Judith Gronim for the line drawings that appear 
in the text, and Mr. Coburn Bennett, Ms. Amina 
Anderson, and Mr. John Lupo, who were of in- 
valuable assistance with the photographic work. 

This study was supported in part by grants from 
the National Defense Education Act Title IV Fel- 
lowship and National Institutes of Health Training 
Grant (T-01 GM 00036). Field work was partly 
funded through the Evolutionary Biology Com- 
mittee, Harvard University (National Science 
Foundation Training Grant GB-7346), and the 
generosity of Mrs. Edward C. Sweeney. 

I am grateful to the curators of the herbaria 
mentioned in the text, who lent specimens of 
Brunfelsia used in this study. 

Lastly I would like to express my deep grati- 
tude to my parents, Dr. and Mrs. J. Wesley Plow- 
man, who have continuously supported and en- 
couraged me. 

XIII. Literature Cited 

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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- 
diflora) 

Allard 21176 (grandiflora subsp. grandiflora), 
22096 (grandiflora subsp. schultesii) 

Allemao 1249 (uniflora} 

Allemao & Cysneiro 1250 (uniflora) 

Almeida 1286 (brasiliensis subsp. brasiliensis) 

Alston & Lutz 138 (latifolid) 

Amaral, A. s.n. (brasiliensis subsp. brasiliensis) 

Amaral, I. L., et al. 803 (mire) 

Anderson 11129 (mire), 6751 (obovata var. cori- 
acea), 8949 (brasiliensis subsp. brasiliensis) 

Anderson et al. 37048 (uniflora) 

Andrade 1024 (brasiliensis subsp. brasiliensis) 

Angeli 250 (brasiliensis subsp. brasiliensis) 

Aparicio & Edmundo 868 (hydrangeiformis 
subsp. capitata) 

Araujo 3405 (uniflora), 954 (hydrangeiformis 
subsp. hydrangeiformis), 968 (uniflora) 

Araujo & Kennedy 24 Mar. 1971 (latifolia) 

Araujo et al. 3202 (uniflora) 



Aristeguieta 1669 (grandiflora subsp. schultesii), 
6153 (uniflora) 

Assis 102 (brasiliensis subsp. brasiliensis), 137 
(brasiliensis subsp. brasiliensis), 142 (bras- 
iliensis subsp. brasiliensis), 181 (brasiliensis 
subsp. brasiliensis) 

Atamp. (?) 6560 (brasiliensis subsp. brasiliensis) 

Bailey & Bailey 758 (brasiliensis subsp. bras- 
iliensis) 

Baker 61 (grandiflora subsp. grandiflora) 

Balansa 2240 (australis) 

Balgooy 1592 (australis) 

Bang 1398 (grandiflora subsp. schultesii), 2352 
(mire) 

Barbosa Rodrigues 261 (uniflora) 

Barclay 911 (macrocarpa) 

Barreto 7787 (brasiliensis), 7790 (brasiliensis), 
9749 (brasiliensis subsp. macrocalyx) 

Beck 13335 (grandiflora), 4937 (mire) 

Beck & Liberman 9379 (boliviano) 

Bedi s.n. (uniflora) 

Belshaw 3130 (grandiflora subsp. schultesii) 

Bernardi 18138 (australis), 20495 (australis), 
3310 (grandiflora subsp. schultesii) 

Bertoni 2939 (australis), 2974 (pilosa), 3434 
(australis) 

Besse et al. 513 (mire) 

Beyrich s.n. (brasiliensis subsp. brasiliensis) 

Bezerra 334 (uniflora) 

Black 48-2905 (amazonica) 

Blanchet 1455 (clandestina), 2616 (uniflora), 
3352 (clandestina), 3354 (clandestina), 3862 
(uniflora), Jun. 1844 (uniflora), s.n. (clandes- 
tina), s.n. (clandestina), s.n. 1843 (clandestina), 
s.n. 1847 (uniflora), s.n. 1857 (hydrangeiformis 
subsp. hydrangeiformis) 

Blanco 826 (imatacana) 

Boeke 1224 (grandiflora subsp. grandiflora), 
1477 (mire) 

Boer 1047 (guianensis) 

Bonpland 528 (australis) 

Bornmuller 219 (australis) 

Bowie & Cunningham 100 (hydrangeiformis 
subsp. hydrangeiformis), 228 (brasiliensis 
subsp. brasiliensis) 

Brade 10569 (brasiliensis subsp. brasiliensis), 
10583 (brasiliensis subsp. brasiliensis), 14074 
(brasiliensis subsp. macrocalyx), 18643 (bon- 
odora), 18785 (brasiliensis subsp. brasiliensis), 
20953 (brasiliensis subsp. macrocalyx), 21218 
(brasiliensis subsp. brasiliensis), 6036 (pauci- 
flora), 6037 (brasiliensis subsp. macrocalyx), 
7625 (brasiliensis subsp. macrocalyx), 8088 



124 



FIELDIANA: BOTANY 



(pauciflord), Oct. 1928 (hydrangeiformis subsp. 
capitatd) 

Brade & Duarte 20119 (pauciflord) 

Brade & Tamandare 6576 (brasiliensis subsp. 
macrocalyx) 

Braga 1169 (brasiliensis subsp. brasiliensis), 182 
(brasiliensis subsp. brasiliensis), 203 (bras- 
iliensis subsp. brasiliensis) 

Branddo 2066 (brasiliensis subsp. brasiliensis) 

Bristan 1220 (chocoensis), 566 (chocoensis), 569 
(chocoensis) 

British Guiana Forest Department 4422 (marti- 
ana) 

Brooke 5653 (boliviano) 

Brunei sp. nov. 1888 (hydrangeiformis subsp. 
capitatd) 

Buchtien 1299 (grandiflora subsp. schultesii), 
1300 (grandiflora subsp. schultesii), 2029 
(grandiflora subsp. schultesii), 5640 (mire), 
7615 (mire) 

Bunting et al. 10351 (hydrangeiformis subsp. hy- 
drangeiformis) 

Burchell 1372 (brasiliensis subsp. brasiliensis), 
8494 (burchellii), 8653 (burchellii), 8752 (bur- 
chellii) 

Buttura 157 (australis) 

Camargo 2123 (cuneifolia), 3 Oct. 1957 (cunei- 
folia) 

Campos Novaes 214 (pauciflord), 216 (pilosa), 
2960 (pauciflord), 7756 (uniflora) 

Campos Porto 173 (brasiliensis subsp. macroca- 
lyx), 1879 (hydrangeiformis subsp. capitatd), 
3381 (brasiliensis subsp. brasiliensis), 3400 
(pauciflord), s.n. (brasiliensis subsp. bras- 
iliensis) 

Capanema s.n. (brasiliensis subsp. brasiliensis) 

Caprilioni 1659 (pilosa) 

Carauta 2420 (brasiliensis subsp. brasiliensis) 

Carauta & de Oliveira 1523 (brasiliensis) 

Cardenas 2813 (mire), 3380 (uniflora), 5216 (un- 
iflora), 5952 (uniflora) 

Carvalho 83 (pilosa) 

Carvalho & Gatti 795 (clandestina) 

Castellanos 23004 (uniflora), 23294 (obovata var. 
coriacea), 23973 (brasiliensis subsp. bras- 
iliensis) 

Cavalcante 2518 (guianensis) 

Chiar s.n. (australis) 

Claussen 146 (brasiliensis subsp. brasiliensis), 
147 (brasiliensis subsp. brasiliensis), 157 
(brasiliensis subsp. brasiliensis), 201 (hydran- 
geiformis subsp. hydrangeiformis), 334 (bras- 
iliensis subsp. brasiliensis), Nov. 1839 (hydran- 
geiformis subsp. hydrangeiformis), Nov. 1842 



(hydrangeiformis subsp. capitatd), s.n. (bras- 
iliensis subsp. brasiliensis), s.n. (hydrangeifor- 
mis subsp. hydrangeiformis), s.n. 1841 (hy- 
drangeiformis subsp. hydrangeiformis) 

Constantino Nov. 1931 (brasiliensis subsp. bras- 
iliensis) 

Correa & Dressier 471 (dwyeri) 

Croat 13072 (dwyeri), 36856 (australis) 

Cuatrecasas 10611 (grandiflora subsp. schulte- 
sii), 10795 (grandiflora subsp. schultesii), 
11275 (grandiflora subsp. schultesii) 

Curran 619 (brasiliensis subsp. brasiliensis), 623 
(brasiliensis subsp. brasiliensis), s.n. (australis) 

Cutler 8271 (uniflora) 

D'Arcy 3959 (dwyeri) 

D'Arcy et al. 3948 (dwyeri) 

Davidse & Ramamoorthy 10709A (brasiliensis 
subsp. brasiliensis) 

Davidse et al. 11611 (brasiliensis) 

Davidson 3846 (uniflora) 

Davis & Marshall 1106 (mire) 

Davis & Yost 958 (grandiflora subsp. schultesii) 

Dawe 241 (grandiflora subsp. schultesii), Mar. 
1914 (uniflora) 

Dobereiner & Tokarnia 732 (cuneifolia), 841 
(australis) 

Dodson 2822 (grandiflora subsp. schultesii) 

Drummond 7330 (grandiflora subsp. schultesii) 

Duarte 2029 (brasiliensis subsp. brasiliensis), 
5477 (bonodora), 631 (brasiliensis subsp. bras- 
iliensis), 8445 (hydrangeiformis subsp. hydran- 
geiformis), 8741 (brasiliensis subsp. bras- 
iliensis) 

Duarte & Pereira 22 Oct. 1964 (hydrangeiformis 
subsp. hydrangeiformis), 9 Dec. 1948 (hydran- 
geiformis subsp. capitatd) 

Duarte & Santos 102 (obovata var. coriacea) 

Duarte de Barros 1195 (brasiliensis subsp. bras- 
iliensis) 

Ducke 2954 (uniflora), 430 (amazonica), 865 
(grandiflora subsp. schultesii), 29 Nov. 1928 
(hydrangeiformis subsp. capitatd), 10 Jan. 
1930 (mire), s.n. (amazonica), s.n. (guianensis), 
s.n. (martiana) 

Ducke & Kuhlmann s.n. (brasiliensis subsp. bras- 
iliensis) 

Duke 11351 (chocoensis), 13331 (chocoensis), 
16047 (chiricaspi), 5300 (chocoensis), 9429 
(dwyeri) 

Dupre s.n. 1842 (brasiliensis subsp. brasiliensis) 

Dusen 10305 (pauciflord), 10354 (pauciflord), 
10443 (pilosa), 13180 (pauciflord), 13298 
(pauciflord), 13327 (pauciflord), 13562 (bras- 
iliensis subsp. macrocalyx), 17301 (pauciflord), 



PLOWMAN: REVISION OF BRUNFELSIA 



125 



2023 (brasiliensis subsp. macrocalyx), 3388 
(brasiliensis subsp. macrocalyx), 569 (bras- 
iliensis subsp. macrocalyx), 7077 (pilosa), 8679 
(pauciflora), 8961 (pauciflora) 

Dwyer 7280 (dwyeri) 

Dwyer & Ganger 7349 (dwyeri) 

Dwyer et al. 5051 (dwyeri) 

Edwall, 15183 (hydrangeiformis subsp. capitata), 
1981 p.p. (brasiliensis subsp. macrocalyx), 
1981 p.p. (obovata var. obovatd), 4329 (pau- 
ciflora), 6133 (uniflora) 

Elburg 9819 (guianensis) 

Emmerich 985 (brasiliensis subsp. brasiliensis) 

Engler 1006 (brasiliensis subsp. brasiliensis), 
1009 (brasiliensis subsp. brasiliensis), s.n. 
(brasiliensis subsp. brasiliensis) 

Escobar 359 (dwyeri) 

Fabris 4502 (uniflora) 

Facultad Ciencias For estates s.n. (uniflora) 

Farney et al. 584 (brasiliensis subsp. bras- 
iliensis), 597 (brasiliensis) 

Fernandes, A., s.n. (uniflora) 

Fernandez 365 (macrocarpa) 

Ferreira & Costa 16 (grandiflora subsp. grandi- 
flora) 

Ferreira, V.F., et al. 887 (rupestris) 

Ferreyra 4760 (grandiflora subsp. schultesii), 
7778 (grandiflora subsp. schultesii), 985 (gran- 
diflora subsp. grandiflora) 

Fiebrig 49 (australis), 5460 (pilosa) 

Filho S. J. 8 (uniflora) 

Filho et al. 3219 (uniflora) 

Plaster 56 (hydrangeiformis subsp. capitata) 

Florschiitz & Maas 2924 (guianensis), 3094 (gui- 
anensis) 

Forestry Department of British Guiana 7495 (gui- 
anensis) 

Franklin & Glaziou 5319 (hydrangeiformis subsp. 
capitata) 

Frenzel 666 (cuneifolia) 

Froes 11620 (burchellii), 12681\46 (uniflora), 
20067 (clandestina), 20210 (clandestina), 
20535 (amazonica), 23068 (uniflora), 23974 
(grandiflora subsp. schultesii), 24035 (grandi- 
flora subsp. schultesii), 25650 (martiana), 
28393 (martiana) 

Garcia Barriga 4586 (grandiflora subsp. schul- 
tesii) 

Gardner, G. 1095 (uniflora), 1798 (uniflora), 
2432 (uniflora), 248 (brasiliensis subsp. bras- 
iliensis), 5062 (uniflora), 5063 (brasiliensis 
subsp. brasiliensis) 5065 (hydrangeiformis 
subsp. hydrangeiformis), 5566 (uniflora), 563 



(hydrangeiformis subsp. capitata), 564 (bras- 
iliensis subsp. brasiliensis) May 1838 (uniflora) 

Gaudichaud 442 (brasiliensis subsp. brasiliensis), 
705 (pilosa) 

Gehrt 3667 (brasiliensis subsp. brasiliensis) 

Gentry 16249 (grandiflora subsp. schultesii) 

Gentry & Clewell 7138 (chocoensis) 

Gentry et al. 15707 (grandiflora subsp. schulte- 
sii), 18573 (grandiflora subsp. schultesii), 
18664 (grandiflora subsp. schultesii) 

Germain 171 (brasiliensis subsp. brasiliensis) 

Gill 45 (grandiflora subsp. schultesii) 

Gines 3605 (uniflora) 

Ginzberger 93 (brasiliensis subsp. macrocalyx) 

Glaziou 11392 (hydrangeiformis subsp. hydran- 
geiformis), 11393 (hydrangeiformis subsp. hy- 
drangeiformis), 11394 (brasiliensis subsp. 
brasiliensis), 11395 (brasiliensis subsp. macro- 
calyx), 12110 (australis), 13478 (brasiliensis 
subsp. macrocalyx), 14171 (brasiliensis subsp. 
brasiliensis), 14172 (brasiliensis subsp. macro- 
calyx), 14173 (hydrangeiformis subsp. capita- 
ta), 14174 (hydrangeiformis subsp. capitata), 
15132 (brasiliensis subsp. brasiliensis), 15311 
(brasiliensis subsp. brasiliensis), 17168 (pau- 
ciflora), 17169 (brasiliensis subsp. macroca- 
lyx), 17170 (brasiliensis subsp. brasiliensis), 
21920 (obovata var. coriacea), 357 (uniflora), 
4909 (hydrangeiformis subsp. capitata), 5970 
(latifolia), 6908 (brasiliensis subsp. macroca- 
lyx), 811 (pauciflora), 812 (brasiliensis subsp. 
brasiliensis), 9558 (hydrangeiformis subsp. 
capitata) 

Goes & Constantino 543 (brasiliensis subsp. 
brasiliensis), 883 (brasiliensis subsp. bras- 
iliensis) 

Graf 10 (uniflora) 

Granville 1378 (guianensis), 2204 (guianensis), 
2224 (guianensis), B-4854 (guianensis) 

Guedes 265 (mire), 472 (uniflora) 

Guevara 2276 (uniflora) 

Guillemin 41 (brasiliensis subsp. brasiliensis), 
171 (brasiliensis subsp. brasiliensis), 825 
(brasiliensis subsp. brasiliensis), 954 (bras- 
iliensis subsp. brasiliensis) 

Gurgel 27 Feb. 1929 (pilosa) 

Handro 27 Sep. 1945 (pauciflora), 984 (obovata 
var. obovata) 

Harley 21898 (obovata var. coriacea) 

Harley et al. 16432 (uniflora) 

Harling & Andersson 11724 (grandiflora subsp. 
schultesii), 11771 (chiricaspi), 11804 (chiricas- 
pi), 13906 (grandiflora subsp. grandiflora), 
13934 (grandiflora subsp. grandiflora) 



126 



FIELDIANA: BOTANY 



Marling et al. 7021 (chiricaspi), 7151 (chiricas- 
pi), 7474 (grandiflora subsp. schultesii), 7700 
(grandiflora subsp. schultesii), 7772 (chiricas- 

pi) 

Hashimoto 1 (brasiliensis subsp. brasiliensis) 

Hassler 12559 (australis), 725 (australis) 

Hatschbach 1058 (pauciflora), 11878 (bras- 
iliensis subsp. macrocalyx), 13055 (pauciflora), 
15071 (pauciflora), 15159 (cuneifolia), 15300 
(pauciflora), 17308 (pauciflora), 17610 (pilo- 
sa), 18747 (brasiliensis subsp. macrocalyx), 
19447 (brasiliensis subsp. macrocalyx), 19937 
(pauciflora), 20304 (pauciflora), 20997 (bras- 
iliensis subsp. macrocalyx), 22278 (pauciflora), 
22383 (pilosa), 22771 (pauciflora), 25613 (pi- 
losa), 26532 (brasiliensis subsp. macrocalyx), 
28320 (pilosa), 32637 (pauciflora), 32869 (pi- 
losa), 34918 (pauciflora), 37876 (pilosa), 
39164 (pilosa), 39209 (brasiliensis subsp. mac- 
rocalyx), 40004 (pauciflora), 40470 (pilosa), 
42939 (brasiliensis subsp. brasiliensis), 43192 
(australis), 43273 (pauciflora), 43398 (bras- 
iliensis subsp. brasiliensis), 43843 (obovata 
var. coriacea), 45295 (pauciflora), 45415 (pi- 
losa), 515 (pauciflora) 

Hatschbach & Ahumada 31333 (brasiliensis 
subsp. brasiliensis) 

Hatschbach & Cervi 46844 (pauciflora) 

Hatschbach & Guimaraes 15286 (pilosa), 25473 
(pilosa), 30302 (cuneifolia), 45004 (brasiliensis 
subsp. brasiliensis) 

Hatschbach & Kummrow 37257 (obovata var. 
obovata), 45569 (brasiliensis subsp. bras- 
iliensis) 

Hatschbach & Pelanda 27941 (uniflora), 27993 
(brasiliensis subsp. brasiliensis) 

Hatschbach & Zelma 50296 (rupestris) 

Heiner 209 (pauciflora), s.n. (uniflora) 

Heinrichs 496 (grandiflora subsp. schultesii) 

Henz 27246 (pilosa), 35305 (pilosa) 

Heringer 10556 (obovata var. coriacea), 10719 
(obovata var. coriacea), 11549 (uniflora), 460 
(hydrangeiformis subsp. capitata), 8662 (obo- 
vata var. coriacea), 8993/1197 (obovata var. 
coriacea), 9096/1290 (obovata var. coriacea) 

Heringer et al. 4281 (obovata var. coriacea), 
6148 (brasiliensis subsp. brasiliensis) 

Hermann 11237 (grandiflora subsp. schultesii) 

Heyligers 438 (guianensis) 

Hoehne, F. C. 582 (pauciflora), 23169 (pauciflo- 
ra), 23309 (pilosa), 24405 (pauciflora), 4595 
(brasiliensis subsp. macrocalyx), 4920 (bras- 
iliensis subsp. brasiliensis), 5481 (brasiliensis 
subsp. brasiliensis), 6129 (brasiliensis subsp. 



brasiliensis), 6846 (brasiliensis subsp. bras- 
iliensis), 7953 (pauciflora), 8250 (brasiliensis 
subsp. macrocalyx) 

Hoehne, W., 6154 (brasiliensis subsp. bras- 
iliensis), 6155 (brasiliensis subsp. brasiliensis) 

Huber 14990 (grandiflora subsp. grandiflora), 
1556 (grandiflora subsp. grandiflora), 1562 
(grandiflora subsp. grandiflora), 4204 (gran- 
diflora subsp. schultesii), 4377 (grandiflora 
subsp. schultesii), 4575 (grandiflora subsp. 
grandiflora), 4589 (grandiflora subsp. grandi- 
flora), 887 (guianensis) 

I.N.T.A. 4435 (pilosa) 

Irwin 2269 (brasiliensis subsp. brasiliensis), 2674 
(uniflora), 2701 (uniflora), 48753 (grandiflora 
subsp. schultesii) 

Irwin et al. 13126 (obovata var. coriacea), 13194 
(brasiliensis subsp. brasiliensis), 15799 (bras- 
iliensis subsp. brasiliensis), 19549 (brasiliensis 
subsp. brasiliensis), 20150 (brasiliensis subsp. 
brasiliensis), 22077 (rupestris), 22465 (rupes- 
tris), 26556 (brasiliensis subsp. brasiliensis), 
29189 (brasiliensis subsp. brasiliensis), 29407 
(brasiliensis subsp. brasiliensis), 30628 (bras- 
iliensis subsp. brasiliensis), 32406 (uniflora), 
47359 (grandiflora subsp. schultesii), 47900 
(guianensis), 55978 (guianensis), 8152 (obov- 
ata var. coriacea), 8514 (obovata var. coriacea) 

Isabelle 6 (pilosa) 

Jarenkow 132 (cuneifolia) 

Johnston 71 (uniflora) 

Johnston & Miller 265 (uniflora) 

Joly 1133 (brasiliensis subsp. brasiliensis) 

Jons son, G. 1054a (pilosa) 

Jorgensen 2098 (australis), 3661 (australis), 4 
Oct. 1909 (pilosa), Nov. 1910 (pilosa) 

Juajibioy Chindoy 277 (grandiflora subsp. schul- 
tesii) 

Jurgen 19 (pilosa) 

Kennedy 1386 (chiricaspi), 316 (dwyeri) 

Kermann 270 (hydrangeiformis subsp. capitata) 

Kermes 1363 (australis) 

Killip & Smith 26844 (grandiflora subsp. schul- 
tesii), 27667 (grandiflora subsp. schultesii), 
28644 (grandiflora subsp. schultesii) 

Klein 1751 (pauciflora), 319 (pauciflora), 3518 
(pilosa), 4102 (pilosa), 4343 (cuneifolia) 

Klein & Bresolin 6864 (pauciflora) 

Klug 741 (grandiflora subsp. schultesii), 2880 
(grandiflora subsp. schultesii), 3326 (grandiflo- 
ra subsp. schultesii) 

Krapovickas 29261 (australis) 

Krapovickas & Cristobal 11706 (australis), 
15385 (australis) 



PLOWMAN: REVISION OF BRUNFELSIA 



127 



Krapovickas & Schinini 31476 (boliviano), 35155 
(grandiflora subsp. schultesii) 

Krapovickas et al. 26092 (australis) 

Krukoff 1532 (mire), 5284 (grandiflora subsp. 
grandiflora) 

Kuhlmann, J.G., 1326 (mire), 1327-k (mire), 141 
(obovata var. obovata), 1722 (guianensis), 
2287 (mire), 809 (mire), 943 (amazonica), 8 
Nov. 1922 (latifolia), 19 Nov. 1928 (uniflora), 
3 Feb. 1930 (brasiliensis subsp. brasiliensis), 
24 Oct. 1932 (hydrangeiformis subsp. hydran- 
geiformis), 2 Apr. 1935 (uniflora), 10 Nov. 
1943 (uniflora), 8 Oct. 1946 (pilosa) 

Kuhlmann, M., 2176 (pauciflora), 2714 (pauciflo- 
ra), 2926 (obovata var. obovata), 4309 (bras- 
iliensis subsp. macrocalyx), 4359 (brasiliensis 
subsp. macrocalyx) 

Kummrow 1575 (pauciflora), 2420 (pauciflora), 
2432 (cuneifolia), 94 (pilosa), 970 (brasiliensis 
subsp. macrocalyx) 

Lalandes s.n. (brasiliensis subsp. brasiliensis) 

Landrum et al. 4150 (brasiliensis subsp. bras- 
iliensis) 

Langsdorff 43 (hydrangeiformis subsp. hydran- 
geiformis), s.n. (brasiliensis subsp. bras- 
iliensis), s.n. (hydrangeiformis subsp. hydran- 
geiformis) 

Lanjouw 345 (guianensis) 

Lanjouw & Lindeman 2137 (guianensis), 2564 
(guianensis), 2888 (guianensis) 

Leite 3660 (pauciflora), 4188 (hydrangeiformis 
subsp. hydrangeiformis) 

Lemme s.n. (guianensis) 

Levy 12 (pauciflora) 

Lima, A. S. 7859 (mire) 

Lima, D. de A., 52-1039 (uniflora) 

Lima, H. C. de, and Ramamoorthy 1224 (bras- 
iliensis subsp. brasiliensis) 

Lima, H. C. de, et al. 1637 (uniflora) 

Lima, H. C. de, et al. 2122 (grandiflora subsp. 
grandiflora) 

Lindberg 203 (uniflora) 

Lindeman 5889 (guianensis), 6069 (guianensis), 
6162 (guianensis) 

Lindeman & Haas 1174 (brasiliensis subsp. mac- 
rocalyx), 3248 (pilosa), 5157 (brasiliensis 
subsp. macrocalyx) 

Lindeman et al. 8233 (pilosa) 

Lindeman, C. A-597-a (australis), A-597-b (pilo- 
sa), A-597-c (pilosa) 

Llamas 1530 (pilosa) 

Llamas sp. nov. (australis) 

Lobb 65 (grandiflora subsp. grandiflora), s.n. 



(brasiliensis subsp. brasiliensis), s.n. (hydran- 
geiformis subsp. capitata) 

Lofgren 86 (brasiliensis subsp. brasiliensis), 389 
(pilosa), 3986 (brasiliensis subsp. brasiliensis), 
948 (pauciflora) 

Lofgren & Cowan 1660 (pauciflora) 

Lofgren & Edwall 1435 (uniflora), 2705 (pauci- 
flora) 

Longfield 367 (macrocarpa) 

Lorentz & Hieronymus 258 (uniflora) 

Luderwaldt 1086 (brasiliensis subsp. bras- 
iliensis), 2117 (pilosa) 

Lugo 23 (grandiflora subsp. grandiflora), 2724 
(chiricaspi), 2942 (chiricaspi), 2981 (chiricas- 
pi), 3476 (chiricaspi) 

Lund 754 (brasiliensis subsp. brasiliensis), 755 
(obovata var. obovata), 756 (brasiliensis subsp. 
brasiliensis) 

Luschnath Sep. 1833 (uniflora), Nov. 1833 (uni- 
flora), s.n. 1832 (brasiliensis subsp. bras- 
iliensis), s.n. 1834 (brasiliensis subsp. bras- 
iliensis), s.n. 1834 (latifolia) 

Luteyn et al. 4769 (grandiflora subsp. grandiflo- 
ra) 

Lutz 1551 (hydrangeiformis subsp. capitata), Jan. 
1925 (brasiliensis subsp. brasiliensis), Boc 14 
(hydrangeiformis subsp. capitata) 

Liitzelburg 4019 (obovata var. coriacea), 6187 
(brasiliensis subsp. macrocalyx) 6495 (hydran- 
geiformis subsp. capitata), 6559 (brasiliensis 
subsp. macrocalyx) 

Maas et al. 12963 (grandiflora subsp. grandiflo- 
ra), 3326 (hydrangeiformis subsp. hydrangei- 
formis), PI 2709 (grandiflora subsp. grandiflo- 
ra), P13228 (grandiflora subsp. schultesii) 

Macedo 3131 (brasiliensis subsp. brasiliensis), 5 
Feb. 1948 (uniflora) 

Madison 986 (mire) 

Madison et al. 5390 (grandiflora subsp. schulte- 
sii) 

Magalhaes 1643 (hydrangeiformis subsp. hydran- 
geiformis) 

Maguire 23894 (guianensis) 

Markgraf 3495 (brasiliensis subsp. brasiliensis) 

Markgraf & Apparicio 10495 (hydrangeiformis 
subsp. capitata) 

Martinelli 1115 (hydrangeiformis subsp. hydran- 
geiformis), 5135 (uniflora) 

Martinelli et al. 5665 (uniflora) 

Martius 1292 (brasiliensis subsp. brasiliensis), 
3300 (martiana), 541 (bonodora), Dec. 1817 
(brasiliensis subsp. brasiliensis), s.n. (guianen- 
sis), s.n. (martiana), s.n. (uniflora) 

Matthews 1320 (grandiflora subsp. schultesii) 



128 



FIELDIANA: BOTANY 



Mattos Silva et al. 357 (clandestina) 

Mattos, J., & Mattos, N., 14284 (brasiliensis 
subsp. brasiliensis) 

McDaniel 15328 (grandiflora subsp. schultesii) 

Melin 186 (grandiflora subsp. schultesii) 

Mello Barreto 10530 (brasiliensis subsp. bras- 
iliensis), 7767 (brasiliensis subsp. brasiliensis), 
7784 (brasiliensis subsp. brasiliensis), 7786 
(brasiliensis subsp. brasiliensis), 7788 (bras- 
iliensis subsp. brasiliensis), 7789 (brasiliensis 
subsp. brasiliensis), 7791 (brasiliensis subsp. 
brasiliensis), 7793 (brasiliensis subsp. bras- 
iliensis), 7794 (brasiliensis subsp. brasiliensis), 
9063 (brasiliensis subsp. brasiliensis), 9479 
(rupestris), 9833 (brasiliensis subsp. bras- 
iliensis) 

Mello Filho 793 (pilosa) 

Mexia 4089-a (hydrangeiformis subsp. capitata), 
4551 (uniflora), 5182 (uniflora), 5467 (bras- 
iliensis subsp. brasiliensis), 6444 (grandiflora 
subsp. schultesii), 7275 (grandiflora subsp. 
grandiflora) 

Meyer 12621 (australis), 166 (australis), 3076 
(australis) 

Miers 4495 (brasiliensis subsp. brasiliensis), 
4721 (uniflora), Dec. 1837 (hydrangeiformis 
subsp. capitata), Feb. 1838 (brasiliensis subsp. 
macrocalyx), s.n. 1838 (hydrangeiformis subsp. 
capitata) 

Mikan s.n. (latifolia) 

Mimura 53 (brasiliensis subsp. brasiliensis) 

Monies 27471 (australis), 4703 (australis) 

Mora 1040 (chiricaspi) 

Moreira 5775 (brasiliensis subsp. brasiliensis), 
s.n. (brasiliensis subsp. macrocalyx) 

Morel 3783 (australis), 4144 (australis), 8921 
(australis) 

Mori 6828 (chocoensis) 

Mori & Bolten 8394 (guianensis), 8575 (guianen- 
sis) 

Mori & Boom 14726 (guianensis), 14834 (gui- 
anensis) 

Mori et al. 12082 (clandestina), 8767 (guianen- 
sis), 9775 (clandestina) 

Moura s.n., 1887 (hydrangeiformis subsp. hy- 
drangeiformis) 

Mosen 673 (uniflora) 

Mueller 435 (pauciflora) 

Muniez 99 (australis) 

Neill 6546 (grandiflora subsp. grandiflora) 

Neill & Palacios 7101 (grandiflora subsp. gran- 
diflora) 

Netto s.n. (brasiliensis subsp. brasiliensis) 

Netto et al. s.n. (brasiliensis subsp. brasiliensis) 



Neuwied, Pr. von s.n. 1827 (uniflora) 

Noblick 2038 (uniflora) 

Occhioni 1131 (brasiliensis subsp. macrocalyx), 
9174 (brasiliensis subsp. brasiliensis) 

Oldeman B-1896 (guianensis) 

Oliveira 21 (brasiliensis subsp. brasiliensis) 

Oliveira 100 (pilosa) 

Otto 1003 (uniflora) 

Pabst 5393 (pauciflora), 7411 (brasiliensis subsp. 
brasiliensis), 8340 (brasiliensis subsp. bras- 
iliensis), 9115 (brasiliensis subsp. brasiliensis) 

Pabst & Brade 10283 (brasiliensis subsp. macro- 
calyx) 

Pabst & Pereira 6023 (uniflora) 

Paiva & Stehmann s.n. (australis) 

Palacios et al. 3467 (brasiliensis subsp. bras- 
iliensis) 

Palmer 155 (australis) 

Pearce 290 (grandiflora subsp. grandiflora), 295 
(grandiflora subsp. grandiflora), Mar. 1868 
(grandiflora subsp. grandiflora) 

Peckolt, O., Aug. 1934 (hydrangeiformis subsp. 
hydrangeiformis) 

Peckolt, T., 71 (hydrangeiformis subsp. capitata) 

Pedersen 418 (australis), 418a (australis) 

Pereira 6066 (pauciflora), 9241 (hydrangeiformis 
subsp. hydrangeiformis), 9451 (brasiliensis 
subsp. brasiliensis) 

Perez Arbelaez 688 (grandiflora subsp. schultesii) 

Perrottet s.n. (guianensis) 

Picket 2200 (uniflora), 670 (uniflora) 

Pierroti 6563 (australis) 

Pinheiro 1980 (clandestina), 2086 (clandestina) 

Pinkley 202 (grandiflora subsp. grandiflora), 420 
(chiricaspi), 43 (grandiflora subsp. grandiflo- 
ra), 444 (grandiflora subsp. schultesii), 457 
(grandiflora subsp. schultesii), 460 (grandiflora 
subsp. grandiflora), 563 (chiricaspi), 575 
(grandiflora subsp. schultesii) 

Pires 51899 (guianensis) 

Pires & Silva 10218 (guianensis), 11436 (gui- 
anensis), 11811 (mire) 

Plaumann 164 (pilosa) 

Plowmann 10020 (obovata var. coriacea), 1911 
(uniflora), 1919 (imatacana), 2039 (grandiflora 
subsp. schultesii), 2080 (chiricaspi), 2081 
(chiricaspi), 2090 (grandiflora subsp. schulte- 
sii), 2092 (chiricaspi), 2183 (grandiflora subsp. 
schultesii), 2494 (grandiflora subsp. schultesii), 
2713 (australis), 2723 (australis), 2735 (pilo- 
sa), 2739 (pauciflora), 3160 (dwyeri), 5825 
(grandiflora subsp. schultesii), 5984 (grandiflo- 
ra subsp. schultesii) 



PLOWMAN: REVISION OF BRUNFELSIA 



129 



Plowman & Davis 4852 (mire}, 5000 (mire), 5036 
(mire), 5172 (mire) 

Plowman & de Lima, H.C., 12901 (hydrangeifor- 
mis subsp. capitata) 

Plowman & Kennedy 2310 (grandiflora subsp. 
schultesii) 

Plowman & Sucre 2786 (brasiliensis subsp. bras- 
iliensis), 2891 (brasiliensis subsp. macrocalyx), 
2906 (brasiliensis subsp. macrocalyx) 

Plowman et al. 10066 (clandestina), 10087 (clan- 
destina), 2404 (grandiflora subsp. schultesii), 
2407 (grandiflora subsp. schultesii), 6455 
(grandiflora subsp. schultesii), 6737 (grandiflo- 
ra subsp. schultesii), 6939 (grandiflora subsp. 
schultesii) 

Poeppig 2206 (grandiflora subsp. schultesii) 

Pohl 263 (brasiliensis subsp. brasiliensis), 264 
(uniflord), 265 (brasiliensis subsp. bras- 
iliensis), 266 (brasiliensis subsp. brasiliensis), 
s.n. (uniflord) 

Poiteau s.n. (guianensis) 

Prance & Silva 59030 (obovata var. coriaced) 

Prance et al. 11611 (martiana), 11998 (grandiflo- 
ra subsp. grandiflora), 13471 (grandiflora 
subsp. grandiflora), 14551 (grandiflora subsp. 
schultesii), 3880 (amazonica), 4704 (martiana), 
5854 (grandiflora subsp. schultesii), 6570 
(grandiflora subsp. schultesii), 6933 (cuneifol- 
ia), 8265 (mire), 9553 (uniflord) 

Profice et al. 16 (uniflord) 

Puttemans 4328 (pilosa), 5890 (brasiliensis 
subsp. macrocalyx), 6135 (pauciflord) 

Raben 14 (hydrangeiformis subsp. hydrangeifor- 
mis), 703 (brasiliensis subsp. brasiliensis), 746 
(pauciflord) 

Ragonese 3646 (australis) 

Rambo 29147 (australis}, 321 (australis), 37819 
(australis), 44295 (pilosa), 49144 (pilosa) 

Regnell 173 (brasiliensis subsp. brasiliensis), I- 
374 (uniflord), 1-375 (brasiliensis subsp. bras- 
iliensis) 

Reiss 99 (pilosa) 

Reitz 2743 (pauciflord), 2768 (pauciflord), 3985 
(brasiliensis subsp. macrocalyx), 4375 (bras- 
iliensis subsp. macrocalyx), C-171 (pilosa), C- 
1952 (pauciflord) 

Reitz & Klein 11250 (australis), 1158 (pauciflo- 
rd), 14136 (pilosa), 16178 (cuneifolid), 2160 
(pauciflord), 4076 (cuneifolid), 4238 (pauciflo- 
rd), 5601 (pauciflord), 7329 (pauciflord), 7601 
(brasiliensis subsp. macrocalyx), 9205 (pauci- 
flord) 

Revilla 1724 (grandiflora subsp. schultesii), 304 
(grandiflora subsp. schultesii) 



Riedel 112 (bonodord), 18 (bonodora), 59 (bon- 
odora), s.n. (bonodora) 1112 (uniflord), 113 
(hydrangeiformis subsp. capitata), 1190 (bras- 
iliensis subsp. brasiliensis), 1249 (brasiliensis 
subsp. brasiliensis), 1308 (latifolid), 1465 
(brasiliensis subsp. brasiliensis), 1466 (bras- 
iliensis subsp. brasiliensis), 1467 (obovata var. 
obovata), 467 (brasiliensis subsp. brasiliensis), 
11 Dec. 1823 (hydrangeiformis subsp. capita- 
ta), Oct. 1833 (uniflord), Jan. 1834 (uniflord), 
s.n. (brasiliensis subsp. brasiliensis), s.n. (lati- 
folid) 

Rizzini 7 Nov. 1972 (latifolid) 

Rodriguez 484 (australis) 

Rojas 12133 (australis) 

Rojas Acosta, N., 13004 (australis) 

Rosa 622 (grandiflora) 

Rose & Russel 20221 (brasiliensis subsp. bras- 
iliensis) 

Roth 356 (pauciflord) 

Rubens Faria 4 (pauciflord) 

Rusby 1030 (mire), 2122 (grandiflora subsp. 
schultesii), 2611 (mire), 621 (grandiflora 
subsp. schultesii), 862 (mire) 

Saldanha 8587 (brasiliensis subsp. brasiliensis) 

Sampaio 68 (brasiliensis subsp. brasiliensis), 74 
(brasiliensis subsp. brasiliensis) 

Santos, N. 5790 (latifolid) 

Santos, T. Soares dos, 2511 (uniflord), 1964 
(clandestina), 2015 (clandestina), 433 (clan- 
destina), 689 (clandestina) 

Santos, J. U. et al. 48 (grandiflora subsp. gran- 
diflora) 

Santos Lima Aug. 1940 (pauciflord) 

Santos-Lima 200 (hydrangeiformis subsp. hydran- 
geiformis) 

Sastre 1649 (guianensis), 4751 (guianensis) 

Schinini et al. 18580 (australis) 

Schott 5361 (martiana), s.n. (martiana) 

Schubert 2150 (guianensis) 

Schuch 591 (hydrangeiformis subsp. hydrangei- 
formis), s.n. (brasiliensis subsp. brasiliensis) 

Schultes 3340 (grandiflora subsp. schultesii), 
3422 (chiricaspi) 

Schultes & Cabrera 19115 (grandiflora subsp. 
schultesii) 

Schultes & Rodrigues 26132A (martiana) 

Schulz 85 (australis), 8645 (guianensis) 

Schunke V., J. 8 (grandiflora subsp. schultesii), 
1480 (grandiflora subsp. schultesii), 3480 
(grandiflora subsp. grandiflora), 4034 (gran- 
diflora subsp. grandiflora), 4051 (grandiflora 
subsp. grandiflora) Schwacke 3311 (uniflord), 
s.n. (brasiliensis subsp. brasiliensis) 



130 



FIELDIANA: BOTANY 



Schwarz 139 (australis), 36 (australis), 48 (aus- 
tralis), 93 (australis), 8114 (australis), 8547 
(australis), 8661 (australis), 8972 (australis) 

Schwarzer 50 (pilosa) 

Schwindt 2270 (pilosa) 

Scolnik 1495 (grandiflora subsp. grandiflora) 

Sellow 1573 (pilosa), 719 (hydrangeiformis 
subsp. capitata), s.n. (brasiliensis subsp. bras- 
iliensis), s.n. (hydrangeiformis subsp. capitata), 
s.n. (hydrangeiformis subsp. hydrangeiformis) 

Silva, N. T. & Santos, M.R., 4631 (guianensis) 

Silva, N. T. da 336 (guianensis) 

Silva, M. N. et al. 17 (mire) 

Smith, A.C., 3163 (uniflora) 

Smith, L.B., 1788 (brasiliensis subsp. macroca- 
lyx), 1989 (brasiliensis subsp. macrocalyx), 
5786 (pauciflora) 

Smith, L.B., & Klein 7532 (pauciflora), 13044 
(cuneifolia), 8567 (cuneifolia) 

Smith, L.B., & Reitz 12541 (cuneifolia), 12757 
(pilosa), 12870 (pilosa) 

Smith, L.B., et al. 7936 (pauciflora) 

Solomon 7609 (grandiflora subsp. grandiflora), 
7905 (grandiflora subsp. grandiflora) 

Solomon et al. 12022 (mire), 12071 (mire) 

Soukup 3041 (grandiflora subsp. schultesii), 446 
(mire) 

Souza et al. 521 (brasiliensis subsp. brasiliensis) 

Spruce 1495 (amazonica), 3973 (grandiflora 
subsp. schultesii) 

St. Hilaire A-555 (hydrangeiformis subsp. hy- 
drangeiformis), A-746 (brasiliensis subsp. 
brasiliensis), B-2051 (obovata var. obovata), B- 
58 (hydrangeiformis subsp. capitata), D-80 
(brasiliensis subsp. brasiliensis), s.n. (bras- 
iliensis subsp. brasiliensis) 

Steinbach 1805 (grandiflora subsp. schultesii), 
5487 (grandiflora subsp. schultesii), 7399 
(grandiflora subsp. schultesii), 7505 (mire), 
7936 (grandiflora subsp. schultesii), 8345 (un- 
iflora) 

Stephan s.n. (brasiliensis subsp. brasiliensis) 

Steyermark 86244 (imatacana), 86574 (imata- 
cana), 88290 (imatacana) 

Stork et al. 10461 (mire) 

Strang 172 (brasiliensis subsp. brasiliensis) 

Stuckert 16307 (australis) 

Sucre 2339 (brasiliensis subsp. brasiliensis), 2360 
(brasiliensis subsp. brasiliensis), 4213 (bras- 
iliensis subsp. brasiliensis), 5314 (latifolia), 
5806 (brasiliensis subsp. macrocalyx), 7446 
(brasiliensis subsp. brasiliensis), 8002 (bras- 
iliensis) 

Sucre & Plowman 5086 (brasiliensis subsp. bras- 



iliensis), 5191 (brasiliensis subsp. macrocalyx), 
5206 (brasiliensis subsp. macrocalyx) 

Sucre et al. 6428 (latifolia) 

Tamandare 244 (pilosa) 

Tastevin s.n. (grandiflora subsp. grandiflora) 

Taylor et al. 1084 (martiana) 

Tessman 3243 (grandiflora subsp. schultesii), 
3541 (grandiflora subsp. grandiflora), 4921 
(grandiflora subsp. schultesii), 5005 (grandiflo- 
ra subsp. schultesii), 5541 (grandiflora subsp. 
grandiflora) 

Torres 293 (grandiflora subsp. grandiflora) 

Traill 599 (grandiflora subsp. schultesii), 600 
(grandiflora subsp. schultesii) 

Troll 240 (uniflora) 

Trujillo 3488 (imatacana) 

Tyson 5315 (d\vyeri) 

Tyson et al. 4335 (dwyeri) 

U.S. Exploring Expedition s.n. (hydrangeiformis 
subsp. capitata) 

U.S. South Pacific Exploring Expedition s.n. 
(brasiliensis subsp. brasiliensis) 

Ule 1521 (pilosa), 5453 (grandiflora subsp. schul- 
tesii), 6481 (grandiflora subsp. schultesii), 
7061 (uniflora), 7715 (uniflora), 9746 (gran- 
diflora subsp. schultesii), 9747 (grandiflora 
subsp. schultesii), 9748 (grandiflora subsp. 
schultesii), 9749 (grandiflora subsp. schultesii), 
9750 (mire), Nov. 1894 (hydrangeiformis 
subsp. hydrangeiformis), 23 Aug. 1896 (bras- 
iliensis subsp. brasiliensis), 10 Nov. 1896 
(brasiliensis subsp. brasiliensis) 

Uribe, P., Dec. 1930 (grandiflora subsp. schulte- 
sii) 

Uribe Uribe 1353 (grandiflora subsp. schultesii) 

Usteri 23/306 (brasiliensis subsp. brasiliensis), 23 
Nov. 1906 (pilosa) 

van Donselaar 3114 (guianensis) 

Vargas 15767 (mire), 16301 (grandiflora subsp. 
schultesii), 6311 (mire) 

Vasconcelos & Coelho 26 Feb. 1962 (grandiflora 
subsp. schultesii) 

Vauthier 42 (brasiliensis subsp. brasiliensis), 541 
(brasiliensis subsp. brasiliensis) 

Venturi 4408 (australis) 

Vert 6152 (pauciflora) 

Vianna et al. 1574 (brasiliensis subsp. bras- 
iliensis) 

Vidal 311 (latifolia), 11-5435 (hydrangeiformis 
subsp. capitata), 11-5784 (brasiliensis subsp. 
brasiliensis), s.n. (brasiliensis subsp. bras- 
iliensis) 

Vieira et al. 641 (mire) 



PLOWMAN: REVISION OF BRUNFELSIA 



131 



Vincent s.n. (australis), s.n. (brasiliensis subsp. 
brasiliensis) 

Vogel 677 (brasiliensis subsp. macrocalyx) 

Warming 103 (brasiliensis subsp. brasiliensis) 

Wawra & Maly 426 (brasiliensis subsp. bras- 
iliensis) 

Weberbauer 6151 (grandiflora subsp. grandiflo- 
ra) 

Weddell 1648 (brasiliensis subsp. brasiliensis), 
3621 (boliviano), 818 (hydrangeiformis subsp. 
capitatd), 878 (hydrangeiformis subsp. hydran- 
geiformis), s.n. 1844 (uniflord), s.n. 1858 (un- 
iflord) 

Weir 23 (brasiliensis subsp. brasiliensis) 

Wessels Boer 1195 (guianensis) 

Wettstein & Schiffner Sep. 1901 (pauciflord), s.n. 
(brasiliensis subsp. brasiliensis) 

White 1072 (mire], 1840 (grandiflora subsp. 
schultesii), 2386 (grandiflora subsp. schultesii) 

Widgren 486 (brasiliensis subsp. brasiliensis), 
596 (hydrangeiformis subsp. capitatd), s.n. 

1844 (hydrangeiformis subsp. capitatd), s.n. 

1845 (uniflord) 

Wilbur & Weaver 11358 (dwyeri) 

Williams & Assis 6088 (brasiliensis subsp. bras- 
iliensis), 6859 (brasiliensis subsp. brasiliensis), 
7928 (hydrangeiformis subsp. hydrangeifor- 
mis), 12 Feb. 45 (brasiliensis subsp. bras- 
iliensis) 

Williams, L. 5482 (grandiflora subsp. schultesii) 

Woolston 571 (australis) 

Woytkowksi 34536 (grandiflora subsp. grandiflo- 
ra), 35008 (grandiflora subsp. schultesii), 5026 
(grandiflora subsp. grandiflora), 514 (grandi- 
flora subsp. grandiflora) 5525 (grandiflora 
subsp. schultesii), 6170 (grandiflora subsp. 
schultesii) 

Wullschlagel 1505 (guianensis) 

Ybarrola 2963 (australis), 3270 (australis), 3430 
(australis) 

Zarucchi & Andrade 2316 (grandiflora subsp. 
grandiflora) 

Zaruma et al. 56 (chiricaspi) 

Zernys s.n. (brasiliensis subsp. macrocalyx) 



Appendix II: Brunfelsia sect. 
Brunfelsia 

Description 

Brunfelsia L. sect. Brunfelsia. Type species: B. 
americana L. 



Brunfelsia L., Sp. PL 191. 1753. Type species: B. 

americana L. 
Brunfelsia L. sect. Eubrunfelsia Griseb, Fl. Brit. 

W. Ind. 432. 1861. Type species: B. americana 

L. 
Brunfelsia L. subg. Brunfelsiopsis Urb., Symb. 

Ant. 1: 402. 1899. Type species: B. densifolia 

Krug & Urb. 
Brunfelsiopsis (Urb.) Kuntze in Post & Kuntze, 

Lex. 81. Dec. 1903 ("1904"). Type species: B. 

densifolia Krug & Urb. 
Brunfelsia sect. Brunfelsiopsis (Urb.) Dalla Torre 

& Harms, Gen. Siphon. 453. 1904. Type spe- 
cies: B. densifolia Krug & Urb. 

Shrubs or small trees to 10 m tall. Inflores- 
cence terminal or axillary. Flowers large, 1-many 
per inflorescence. Corolla white, turning yellow 
with age, or reddish purple; tube 6-24 times as 
long as calyx, gradually dilated toward apex, 
open, not constricted at mouth. Anthers oblong, 
weakly bilobate. Stigma subcapitate, convex, sub- 
entire or shallowly bilobate. West Indies (see Fig. 
9). About 20 species. 17 



Names of Brunfelsia Sect. Brunfelsia 

Brunfelsia acunae Hadac, Fol. Geobot. Phytotax. 
5: 430. 1970. 

Brunfelsia americana L., Sp. PI. 191. 1753. (B. 
terminalis Salisb., Prodr. Stirp. 109. 1796. B. 
violacea Lodd, Bot. Cab., t. 792. 1823. B. in- 
odora Mart., Del. Sem. Hort. Monac. 1847; 
Linnaea 24: 180. 1851. B. fallax Duchass. ex 
Griseb., Abh. Konigl. Ges. Wiss. Gottingen. 7: 
242. 1857. B. americana var. pubescens Gri- 
seb., Fl. Brit. W. Ind. 432. 1864. B. abbottii 
Leonard, J. Wash. Acad. Sci. 17(3): 71. 1927.) 
Lesser Antilles, Puerto Rico, Dominican Re- 
public. 

Brunfelsia cestroides A. Rich., Fl. Cub. Fan. 3: 
151. 1853. (B. vinciflora Griseb., Mem. Am. 
Acad. Arts 8(1): 523. 1860.) Cuba. 

Brunfelsia clarensis Britton & P. Wilson, Mem. 
Torrey Bot. Club 16(2): 102. 1920. Cuba. 

Brunfelsia densifolia Krug & Urb., Notizbl. Kon- 
igl. Bot. Gart. Berlin 1(10): 324. 1897. Puerto 
Rico. 

Brunfelsia grisebachii Amshoff, Contrib. Ocas. 



17 This Caribbean group of brunfelsias is being revised 
at present by Victor Fuentes Fiallo of INIFAT, Havana, 
Cuba. Eds. 



132 



FIELDIANA: BOTANY 



Mus. Hist. Nat. Col. "De la salle," Habana 15: 

6. 1956. Cuba. 
Brunfelsia jamaicensis (Benth.) Griseb., Fl. Brit. 

W. Ind. 432. 1864. (B. nitida var. jamaicensis 

Benth. in DC., Prodr. 10: 201. 1846. B. harrisii 

Urb., Symb. Ant. 3: 373. 1903.) Jamaica. 
Brunfelsia lactea Krug & Urb., Notizbl. Bot. 

Gart. Berlin 1(10): 323. 1897. Puerto Rico. 
Brunfelsia linearis Ekman, Feddes Repert. Spec. 

Nov. Regni Veg. 21: 223. 1925. Cuba. 
Brunfelsia macroloba Urb., Symb. Ant. 9: 252. 

1924. Cuba. 
Brunfelsia maliformis Urb., Symb. Ant. 3: 372. 

1903. (B. fawcettii Urb, Symb. Ant. 3: 371. 

1903.) Jamaica. 
Brunfelsia membranacea Urb., Symb. Ant. 5: 

491. 1908. Jamaica. 
Brunfelsia nitida Benth. in DC., Prodr. 10: 201. 

1846. (B. parvifolia A. Rich., Fl. Cub. Fan. 3: 

151. 1853. B. longituba Lem., Jard. Fleur. 4: 

61. 1854.) Cuba. 
Brunfelsia picardae Krug & Urb., Notizbl. Bot. 

Gart. Berlin 1(10): 321. 1897. Haiti. 
Brunfelsia plicata Urb., Symb. Ant. 6: 39. 1909. 

Jamaica. 
Brunfelsia pluriflora Urb., Symb. Ant. 9: 252. 

1924. Cuba. 
Brunfelsia portoricensis Krug & Urb., Notizbl. 

Bot. Gart. Berlin 1(10): 322. 1897. Puerto Rico. 
Brunfelsia purpurea Griseb., Mem. Ann. Acad. 

Arts n. s. 8(1): 523. 1869. Cuba. 
Brunfelsia shaferi Britton & P. Wilson, Mem. 

Torrey Bot. Club 16(2): 102. 1920. Cuba. 
Brunfelsia sinuata A. Rich., Fl. Cub. Fan. 3: 151, 

t. 66. 1853. Cuba. 
Brunfelsia splendida Urb., Symb. Ant. 5: 491. 

1908. Jamaica. 
Brunfelsia undulata Sw., Nov. Gen. Spec. PL 90. 

1788. Jamaica. 



Appendix III: Additional References 

The following papers and accounts relevant to 
Brunfelsia were published after preparation of the 
original manuscript by Plowman. The list is by no 
means exhaustive. Eds. 



BENNETT, B. C. 1992. Hallucinogenic plants of the Shuar 
and related indigenous groups in Amazonian Ecuador 
and Peru. Brittonia, 44(4): 483-493. 

BOAVENTURA, Y. M. S., AND L. H. PiEDADE. 1993. Floral 
biology and cytological analysis of Brunfelsia nitida 
Benth. Revista Brasileira de Genetica, 16(3): 785- 
792. 

BRANTJES, N. B. M. 1978. Sensory responses to flowers 
in night-flying moths, pp. 13-19. In Richards, A. J., 
ed., The Pollination of Flowers by Insects. 

CARLQUIST, S. 1992. Wood anatomy of the Solanaceae: 
A survey. Allertonia, 6(4): 279-326. 

DAULATABAD, C. D., AND K. M. HOSAMANI. 1991. Un- 
usual fatty acids in Brunfelsia americana seed oil: A 
rich source of oil. Journal of the American Oil Chem- 
ist's Society, 68(8): 608-609. 

GASTIAZORO, M. T. C. DE. 1993. Estudio morfoanatomico 
de organos vegetativos en Cestroideae (Solanaceae) II: 
Tribu Salpiglossideae. Kurtziana, 22: 47-72. 

GUYOT, M. 1987. Les stomates du genre Brunfelsia (So- 
lanaceae). Bull. Mus. natn. Hist, nat., Paris, ser. 4, 9, 
section B, Adansonia, 3: 289-315. 

IYER, R. P., J. K. BROWN, M. G. CHAUBAL, AND M. H. 
MALONE. 1977. Brunfelsia hopeana I: Hippocratic 
screening and antiinflammatory evaluation. Lloydia, 
40: 356-360. 

NYAWUAME, H. G. K., AND L. S. GILL. 1994. Epidermal 
studies of some species of family Solanaceae used in 
traditional medicine in West Africa. Feddes Reperto- 
rium, 105(1-2): 49-60. 

RUPPELT, B. M., E. F. R. PEREIRA, L. C. GON^ALVES, AND 
N. A. PEREIRA. 1991. Pharmacological screening of 
plants recommended by folk mesdicine as anti-snake 
venom 1. Analgesic and anti-inflammatory activi- 
ties. Memorias do Institute Oswaldo Cruz, 86(suppl. 
II): 203-205. 

TANNER, E. V. J., AND V. KAPOS. 1982. Leaf structure of 
Jamaican upper montane rain-forest trees. Biotropica, 
14(1): 16-24. 

THYR, B. D., M. J. SAMUEL, AND P. G. BROWN. 1975. 
New solanaceous host records for Corynebacterium 
michiganense. Plant Disease Reporter, 59(7): 595- 
598. 

TOKARNIA, C. H., A. GAVA, L. STOLF, AND P. V. PEIXOTO. 
1991. Experimental poisoning in cattle by Brunfelsia 
pauciflora (Solanaceae). Pesquisa Veterinaria Brasi- 
leira, ll(l-2):9-12. 

TRIGO, M. M. 1992. Contribution to the study of pollen 
in ornamental species. Solanaceae, Convolvulaceae 
and Hydrophyllaceae. Acta Botanica Malacitana, 17: 
209-222. 

VICKERS, W. T, AND T. PLOWMAN. 1984. Useful plants 
of the Siona and Secoya Indians of Eastern Ecuador. 
Fieldiana, Botany, 15: 1-63. 



PLOWMAN: REVISION OF BRUNFELSIA 



133 



Index 

Synonyms and non-English common names are italicized. New names or combinations are in bold- 
face. Page numbers for illustrations are also in boldface. 



Anthocercideae 6 
Atropaceae 4 



Banisteriopsis caapi vii 
Besleria bonodora 59, 60 
Besleria inodora 103 
biogeography 29-34 
Browallia 5 
Brunfelsia 4, 34 
Brunfelsia abbottii 132 
Brunfelsia acuminata 62, 68 
Brunfelsia acunae 132 
Brunfelsia amazonica 38, 39 
Brunfelsia americana 34, 132 

var. pubescens 132 
Brunfelsia angusta 1 19 
Brunfelsia australis 53, 55 
Brunfelsia bahiensis 119 
Brunfelsia boliviana 57, 58 
Brunfelsia bonodora 59, 60 
Brunfelsia brasiliensis 61, 68 

ssp. brasiliensis 62, 64 

ssp. macrocalyx 66, 69 

var. acuminata 62 
Brunfelsia burchellii 40, 41 
Brunfelsia calycina 103 

f. grandiflora 78 

var. eximia 103 

var. floribunda 103 

var. lindeniana 103 

var. macrantha 103 
Brunfelsia capitata 86 

var. angustifolia 86 
Brunfelsia cestroides 132 
Brunfelsia chiricaspi vii, 70, 71 
Brunfelsia chocoensis 40, 42 
Brunfelsia clandestina 43, 44 
Brunfelsia clarensis 132 
Brunfelsia confertiflora 62 
Brunfelsia cuneifolia 72, 73 
Brunfelsia densiflora 34 
Brunfelsia densifolia 132 
Brunfelsia dwyeri 74, 75 
Brunfelsia eximia 103 
Brunfelsia falcata 1 1 9 
Brunfelsia fallax 132 
Brunfelsia fawcettii 133 
Brunfelsia gracilis 119 
Brunfelsia grandiflora 77 

ssp. grandiflora 78, 79 

ssp. schultesii vii, 53, 79, 80 
Brunfelsia grisebachii 132 
Brunfelsia guianensis 38, 46, 47 
Brunfelsia harrisii 133 
Brunfelsia hopeana 1 1 3 

var. australis 53 

var. macrocalyx 66 

var. pubescens \ 1 3 



Brunfelsia hydrangeiformis 84, 85 

ssp. capitata 86, 87 

ssp. hydrangeiformis 86, 85 

var. glabriscula 86 
Brunfelsia imatacana 89 
Brunfelsia inodora 132 
Brunfelsia jamaicensis 133 
Brunfelsia lactea 133 
Brunfelsia latifolia 91, 92, 119 
Brunfelsia lindeniana 103 
Brunfelsia linearis 133 
Brunfelsia lockhartii 119 
Brunfelsia longituba 133 
Brunfelsia macrocarpa 93, 94 
Brunfelsia macroloba 119 
Brunfelsia macrophylla 86 
Brunfelsia magnifica 119 
Brunfelsia maliformis 133 
Brunfelsia maritima 91 
Brunfelsia martiana 49, 50 
Brunfelsia membranacea 133 
Brunfelsia mire 95, 96 
Brunfelsia montana 119 
Brunfelsia multiflora 119 
Brunfelsia mutabilis 1 1 3 
Brunfelsia nitida 133 

\ar.jamaicensis 133 
Brunfelsia nyctaginoides 1 1 9 
Brunfelsia obovata 98, 100 

var. coriacea 101 

var. obovata 99 
Brunfelsia paraguayensis 53 
Brunfelsia parvifolia 133 
Brunfelsia pauciflora 103, 104, 109 

var. calycina 103 
Brunfelsia picardae 133 
Brunfelsia pilosa 108, 110 
Brunfelsia plicata 133 
Brunfelsia plurifolia 133 
Brunfelsia portoricensis 133 
Brunfelsia purpurea 133 
Brunfelsia ramosissima 62, 68 

var. confertiflora 62 

var. laxiflora 62 

var. parcifolia 62 
Brunfelsia rupestris 111, 112 
Brunfelsia schomburgkiana 119 
Brunfelsia sieberi 1 1 9 
Brunfelsia shaferi 133 
Brunfelsia silvicola 101, 102 
Brunfelsia sinuata 133 
Brunfelsia spinosa 119 
Brunfelsia splendida 133 
Brunfelsia spruceana 38, 119 
Brunfelsia tastevinii 78 
Brunfelsia terminalis 132 
Brunfelsia undulata 133 
Brunfelsia uniflora 113, 114 

f. intermedia 53 



f. obovatifolia 53 

f. typica 53 

var. pubescens 1 1 3 
Brunfelsia vinciflora 132 
Brunfelsia violacea 132 
Brunfelsiopsis 34 



chiricaspi 72 
Conejo, village vii 
cultivars 107 
cytology 16 



dispersal 23 



ecology 20 
edible fruits 43, 94 
erva de macaco 69 
evolutionary trends 26-30 
Erythroxylum viii 



flor de Trovoada 74 
flower morphology 10-15 
Franciscea 4, 34, 51 
Franciscea acuminata 62 
Franciscea angusta 1 1 9 
Franciscea australis 53 
Franciscea calycina 103 
Franciscea capitata 86 
Franciscea confertiflora 62, 64 
Franciscea divaricata 62 
Franciscea eximia 103 
Franciscea gracilis 119 
Franciscea grandiflora 77 
Franciscea hopeana 1 1 3 
Franciscea hydrangeiformis 84 
Franciscea latifolia 91 
Franciscea lindeniana 103 
Franciscea macrantha 103 
Franciscea macrophylla 86 
Franciscea maritima 91 
Franciscea mutabilis 1 1 3 
Franciscea obovata 98 
Franciscea pauciflora 103 
Franciscea pohliana 119 
Franciscea ramosissima 62 
Franciscea uniflora 34, 52 



geographical distributions 29-34 
Gerardia brasiliensis 61 
guayabilla 95 
Guianenses, section 38 



hybridization 19-20, 83, 88, 98 



134 



FIELDIANA: BOTANY 



insect-plant relationships 23-26 
Juajibioy, Pedro vii 

key to sections and species 35-37 
Kofan customs vii, 72 

lectotypes (new) 46, 51, 53, 59, 72, 
86, 103 

manacd 40, 49, 69 
manacd assu 69 
manacd da serra 69 
manacd grande 82 
manacd opifera 119 
Mania opifera 119 



medicinal use 77, 82, 84, 98, 118 

mire 95 

morphology 6-16, 26-30 

ornamental 57, 82, 107, 111 

Pinkley, Homer vii 
Plowman, Timothy vii-ix, xi 
Plowmania 6, 1 19 
poisonous 69, 82 
pollination biology 22 

refugium hypothesis 33 

Salpiglossideae 4, 5 
sanango 82 



sananguillo 82 
Schultes, R. E. vii 
Schultesianthus megalandrus 76 
section Franciscea 51 
section Guianenses 38 
Solanaceae 4 

specialization, trends 26-30 
Streptosolen 6 
su-tim-ba-che vii 



thunderstorm flower 74 
tsontinba"k"d vii 
tsontinb"k'o vii 
yage vii 



INDEX 



135 



UNIVERSITY OF ILLINOIS-URBANA