ILL
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BIOLOGY
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FIELDIANA
Botany
Published by Field Museum of Natural History
New Series, No. 8
FIVE NEW SPECIES OF BRUNFELSIA
FROM SOUTH AMERICA (SOLANACEAE)
TIMOTHY PLOWMAN
DEC 11981 ^
July 27, 1981
Publication 1322
FIVE NEW SPECIES OF BRUNFELSIA
FROM SOUTH AMERICA (SOLANACEAE)
FIELDIANA
Botany
Published by Field Museum of Natural History
New Series, No. 8
FIVE NEW SPECIES OF BRUNFELSIA
FROM SOUTH AMERICA (SOLANACEAE)
TIMOTHY PLOWMAN
Assistant Curator of Vascular Plants
Department of Botany
Field Museum of Natural History
Accepted for publication January 27, 1981
July 27, 1981
Publication 1322
Library of Congress Catalog Card Number: 81-65830
ISSN 0015-0746
PRINTED IN THE UNITED STATES OF AMERICA
CONTENTS
LIST OF ILLUSTRATIONS vi
1. Brunfelsia boliviano Plowman, sp. nov 1
2. Brunfelsia imatacana Plowman, sp. nov. 4
3. Brunfelsia rupestris Plowman, sp. nov 7
4. Brunfelsia burchellii Plowman, sp. nov 9
5. Brunfelsia dandestina Plowman, sp. nov 11
ACKNOWLEDGMENTS 15
LITERATURE CITED . 15
LIST OF ILLUSTRATIONS
1. Brunfelsia boliviano Plowman 3
2. Brunfelsia imatacana Plowman 5
3. Brunfelsia rupestris Plowman 8
4. Brunfelsia burchellii Plowman 10
5. Brunfelsia dandestina Plowman 13
6. Geographical distribution of new taxa described 14
VI
Continuing taxonomic studies in the genus Brunfelsia have revealed the ex-
istence of five previously undescribed species. Although some of these taxa were
recognized earlier (Plowman, 1973, 1979), a full assessment of their status was
not possible due to insufficient materials available for study. Since many
new collections from South America have been received and critical specimens
in European and South American herbaria have been discovered, it is now pos-
sible to circumscribe correctly the new taxa that are presented herein.
The species of Brunfelsia in South America and adjacent eastern Panama, com-
prising sections Franciscea and Guianenses, now number 24. Eleven of these have
been described within the past decade.
1. Brunfelsia boliviana Plowman, sp. nov. Sect. Franciscea. Figure 1.
Frutex. Cortex tenuis, longitudinaliter rimosus. Ramuli plus minusve villosi, glabres-
centes. Folia breviter petiolata, obovata, raro elliptico-obovata, ad apicem rotundata, acu-
mine brevi instructa, margine minute revoluta, basi attenuata, nervis inferioribus villosis
exceptis glabrescentia, ad margines ciliato-villosa, firme membranacea vel subcoriacea,
nervis lateralibus 4-9 subtus prominulis. Inflorescentia corymbiformis, in ramulis annotinis
terminalis, pedunculo breviter ramoso villoso instructa, bracteolata, bracteolis lanceolahs
vel cymbiformibus, floribus 6-15, violaceis, speciosis. Pedicellus brevis, glaber vel sparse
glanduloso-puberulus. Calyx tubulosus, pentagonalis, in siccis plicatus, glaber vel sparse
glanduloso-puberulus, dentibus subaequalibus, ovatis, acuminatis, post anthesin persis-
tens, marcescens. Corollae tubus quam calyx plus minusve duplo longior, limbo paten te,
lobis subaequalibus, rotundatis. Stamina superiora e tubo parum exserta, stamina inferiora
et stigma in tubi parte superiore inclusa. Capsula magna, sicca, subglobosa, pericarpio
tenui crustaceo. Semina oblonga vel subreniformia, reticulato-foveolata, brunnescentia.
A Brunfelsia cuneifolia foliis apice acuminatis et margine ciliatis obovatis et cymis in
ramulis annotinis multifloris et breviter pedunculatis differt.
Shrub 1-2 m tall (fide Cardenas). Bark on branches thin, yellowish to grayish brown,
longitudinally 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, short petiolate, blade obovate, rarely elliptic-obovate, apically rounded with a
short acumen, the acumen itself blunt to acute, 5-10 mm long, somewhat revolute at
margin, basally attenuate, 40-135 mm long, 23-58 mm wide, glabrescent on both surfaces
except at the midrib which bears villous and glandular hairs, ciliate-villous on the adaxial
surface of the revolute margin, firmly membranaceous or somewhat coriaceous, dull green
above, paler, yellowish green beneath, the lateral nerves 4-9, straight, arcuately anasto-
mosing towards the margin, prominulous beneath; petiole 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. Bracteoles small, lanceolate or cymbiform, 1-4 mm long, sparsely to
densely villous especially at margin, caducous. Flowers light violet fading to white (fide
Cardenas). Pedicel short, 2-6 mm long, 1 mm in diameter, scarcely thickened at apex,
glabrous or with sparse glandular hairs. Calyx tubular, truncate at base, five-angled in
cross-section, appearing plicate in pressed specimens, 9-15 mm long, 3-^1 mm in diameter,
glabrous or with scattered glandular hairs, teeth subequal, 2-4 mm long, ovate, apically
short acuminate, the acumen itself blunt and glandular-papillose, fruiting calyx persistent,
withering. Corolla tube about twice as long as calyx, straight, 21-25 mm long, 2-3 mm
in diameter; limb spreading, 18-30 mm in diameter, lobes subequal, rounded, 8-14 mm
2 FIELDIANA: BOTANY
long. Stamens inserted in upper part of corolla tube; filaments strap-shaped, the longer
anterior pair suberect, briefly exserted from mouth of corolla tube, 3 mm long, the shorter
posterior pair included, 2 mm long; anthers orbicular-reniform, subequal or the upper pair
slightly larger, 1-1.2 mm in diameter. Ovary narrowly ovoid, 2-3 mm long, 1.8-2.2 mm
in diameter, with about 30 ovules; style incurved at apex, 19 mm long; stigma briefly bifid,
2 mm long. Capsules 3-6 per infructescence, dry at maturity, subglobose, 20-25 mm long,
20-25 mm in diameter, smooth, green; pericarp thin, 1-1.5 mm thick, endocarp thin-
crustaceous, with 22-28 seeds per capsule. Seeds oblong to subreniform, subterete, 5-8
mm long, 2-3 mm in diameter, reticulate-pitted, dark brown. Embryo straight, 4 mm long;
cotyledons ovate, 1.5 mm long; radicle 2.5 mm long.
Type. — BOLIVIA. Dept. Santa Cruz: Prov. de la Cordillera, region of Lagun-
illas, Cordillera of Incahuasi. Altitude 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. August 1934. M. Cardenas 2813 (holotype, F 756420).
Etymology. — From Latin bolivianus, "Bolivian," referring to the country of origin
where the species appears to be endemic.
Common name. — Bella union, "beautiful union," referring to the occurrence of
both violet and white flowers on the same plant. This name is applied to other
brunfelsias in Bolivia.
Distribution. — The eastern Andes of southern Bolivia.
Additional specimens studied.— BOLIVIA. SANTA CRUZ: Prov. Cordillera,
Nov.-Dec. 1845, Weddell 3621 (P, 2 sheets), 5 km N. of Yatarenda, 63° 32' W, 19°
12' S., 17 April 1977, Krapovickas & Schinini 31476 (CTES, F). CHUQUISACA:
mountain above Bartolo, on road from Monteagudo to Sucre, 20° 00' S., 64° 45'
W., 5,000 ft, 20 Sept. 1949, Brooke 5653 (BM).
Brunfelsia boliviana is known from only four collections, 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 Chuquisaca
Department. His collection was probably made near the border between the two
departments.
Brunfelsia boliviana is most closely related to B. cuneifolia J. A. Schmidt, 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 summarized in the
following:
Character B. boliviana B. cuneifolia
Leaf shape Obovate to elliptic- Oblong-obovate to
obovate elliptic-lanceolate
Leaf apex Rounded with a short Cuneate to blunt
acumen
Leaf indument Villous-ciliate at Glabrous at margin
margin
Location of On last season's On current season's
inflorescence growth growth
Number of flowers per 6-16 1-3
inflorescence
Distribution Southern Bolivia Southern Brazil
BRUNFELSIA boliviana Plowman
FIG. 1. Brunfelsia boliviana Plowman. 1, flowering branch; 2, inflorescence in bud; 3,
excised apical portion of corolla tube showing anthers and stigma; 4, leaf margin showing
indument. (From Cardenas 2813.)
4 FIELDIANA: BOTANY
The geographical distribution of these two closely related species deserves
some comment because it exemplifies further a pattern observed in other brun-
felsias in which vicarious species pairs are found in southeastern Brazil and the
eastern Bolivian Andes, respectively. These include B. bonodora (Veil.) Macbr.-B.
grandiflora subsp. schultesii Plowman and B. hydrangeiformis (Pohl) Benth.-B. mire
Monachino. In addition, disjunct and somewhat distinct populations of B. uni-
flora (Pohl) D. Don, 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 over 2,000 km in which no intervening populations are
known to occur. This phytogeographic pattern has been discussed by Smith
(1962) and by Plowman (1979).
Interestingly, the leaves of B. boliviano are reputed to be lethal to cattle, a
feature that has been reported for other Brunfelsia species (Plowman, 1977).
2. Brunfelsia imatacana Plowman, sp. nov. Sect. Franciscea. Figure 2.
Frutex vel arbor parva. Cortex transversaliter et longitudinaliter rimosus. Ramuli tine-
rascentes, pubescentes vel glabrati. Folia in ramulis dispersa, breviter petiolata, oblongo-
elliptica vel elliptico-obovata, apice longe vel breviter acuminata, apice ipso acuto vel
obtuso, basi cuneata vel angustata, utrinque glabra vel subtus sparsim glanduloso-pu-
berula, firme chartacea vel subcoriacea, nervis lateralibus 5-7, rectis, patentibus. Inflores-
centia terminalis, uni- vel biflora, subsessilis, bracteolata, bracteolis caducis, floribus fra-
grantibus, violaceis ad orificium macula alba instruchs, demum albis, pedicellis glabris
post anthesin verrucosis. Calyx tubulosus, teres, glaber vel sparsim glanduloso-puberulus,
dentibus subaequalibus, ovatis, acuminatis, post anthesin paulo accrescens, campanula-
tus, coriaceus, lenticellis instructus. Corollae tubus quam calyx duplo longior, limbo pa-
tente, lobis subaequalibus, rotundatis. Capsula in calyce accrescente omnino inclusa, sicca,
ovoidea vel subglobosa, pericarpio tenui crustaceo. Semina oblongo-ellipsoidea, aliquan-
tum angulata, reticulato-foveolata, brunnescentia.
A Brunfelsia pauciflora foliis elliptico-oblongis vel elliphco-obovatis, nervis lateralibus
5-7, floribus fragrantibus 1-2, pedicellis brevioribus, capsula breviori et seminibus 6-10
majoribus differt.
Shrub or treelet 3-5 m tall. Bark reddish or grayish brown, longitudinally and trans-
versely cracked, shedding in thin flakes. Branchlets about 2 mm in diameter, grayish
brown, longitudinally cracked, pubescent or glabra te. Leaves scattered along branchlets,
short petiolate, blade oblong-elliptic or elliptic-obovate, short to long acuminate, the apex
itself acute to blunt, cuneate to narrowed at base, 65-150 mm long, 20-65 mm wide,
glabrous or sparsely glandular-pubescent beneath, especially at midrib, firmly chartaceous
to subcoriaceous, dull dark green above, dull paler green beneath, the lateral nerves 6-10,
spreading, straight to somewhat arching, indistinct; petiole 5-10 mm long, 1.5 mm in
diameter, pubescent. Inflorescence terminal, 1- or 2-flowered, subsessile. Bracteoles linear-
lanceolate or squamiform, 1-3 mm long, pubescent, caducous. Flowers light violet with
a white spot at orifice, fading to pure white with age, fragrant with odor resembling
jasmine. Pedicel 8-9 mm long, 1 mm in diameter, glabrous, becoming thickened, warty
in fruit. Calyx tubular, terete, 15-21 mm long, 7-10 mm in diameter, glabrous or with
scattered glandular hairs, teeth subequal, ovate, acuminate, 4-5 mm long, calyx in fruit
18-25 mm long, campanulate, thickly coriaceous, dotted with lenticels. Corolla tube twice
as long as calyx, straight, 30-38 mm long, 1-2 mm in diameter, with scattered glandular
hairs, orifice 3 mm across, lobes subequal, rounded, 13-23 mm long. Stamens inserted
in upper part of corolla tube; filaments strap-shaped, 1 mm wide; the longer anterior pair
suberect, included in mouth of corolla tube, 5 mm long, the shorter posterior pair 3 mm
long; anthers orbicular- reniform, upper pair slightly smaller, 1.5 mm across; lower pair
1.8 mm across. Ovary broadly ovoid, 1.8 mm long, 2 mm in diameter, with 7-8 ovules per
locule; style 22-23 mm long; stigma briefly bifid, 1.5-2 mm long. Capsule dry at maturity
and completely enclosed by coriaceous calyx, ovoid to subglobose, 12-17 mm long, 14-16
mm in diameter, smooth, light green, pericarp 0.5 mm thick, crustaceous, containing 6-10
seeds. Seeds oblong-ellipsoid, somewhat angular, 6-9 mm long, 2-5 mm in diameter,
reticulate-pitted, reddish brown.
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6 FIELDIANA: BOTANY
Type. — VENEZUELA. State of Bolivar: About 20 km northeast of Guasipati on
road to El Miamo along side road just before Rio Cabeza Mala, altitude 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. Jazmin del monte, Juan de la Calle. Used
for banos by healers. 25 Sept. 1968. T. Plowman 1918 (holotype, F 1813259; iso-
types, F 1746565, GH, K, NY, P, UC, US, YEN).
Etymology. — From Latin imatacanus, referring to the Serrania de Imataca to
which the species apparently is confined.
Common names. — Jazmin del monte, "wild jasmine"; Juan de la Calle, "John of
the street." The former refers to the jasmine-like fragrance of the flowers.
Distribution. — Restricted to the region of the Serrania de Imataca and Serrania
de Nuria in the northeastern part of the State of Bolivar, Venezuela.
Additional specimens studied.— VENEZUELA. BOLIVAR: Reserva Forestal de
Imataca, along logging road to Campamento Paraiso, 28 km N.E. of Upata,
altitude ca. 100 m, 26 Sept. 1968, Plowman 1919 (ECON, F, GH, K, S, US, VEN);
low, flat woodland east of Rio Cabeza Mala, 15 km N.E. of Guasipati, on road
to Miamo, 2 June 1960, Steyermark 86244 (US, VEN); Altiplanicie del Nuria, 5 km
from Hato de Nuria, east of Miamo, altitude 400 m, 10 Jan. 1961, Steyermark
88290 (NY, VEN); more or less level forest along pica 105, 40 km south of Tu-
meremo, east of highway between Tumeremo and El Dorado, 29 km north of
El Dorado, altitude 220 m, 23 July 1960, Steyermark 86574 (NY, US, VEN); Reserva
Forestal "La Paragua," margenes del no Asa, June 1970, Blanco 826 (F, VEN); El
Dorado, cultivated, Aug. 1957, Trujillo 3488 (MY).
Previously, I considered Brunfelsia imatacana to be a subspecies of the related
species B. pauciflora (Cham. & Schlecht.) Benth. of southeastern Brazil (Plowman,
1973, 1979). Although the two species are superficially similar, B. imatacana can
be readily distinguished by the leaf shape, number of lateral nerves, fragrant
flowers, shorter pedicel, calyx and capsule, and by the smaller number and
larger size of the seeds. In addition the two species are widely disjunct by a
distance of 3,800 km.
Brunfelsia imatacana is endemic in the northeasternmost 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 phytogeographic 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 (Stey-
ermark, 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 Imataca region has been
documented as a major center of both plant and animal species endemism and
has been proposed as a Pleistocene forest refuge area (Prance, 1973; Brown,
1979; Steyermark, 1979).
During the collection of the type material, local guides informed the author
that the foliage of Brunfelsia imatacana was used by healers (brujos) for herbal
baths. Although this use of the plant was not observed, 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 for testing. Dr. Leary (personal communication, 1973) reported that
PLOWMAN: FIVE NEW SPECIES OF BRUNFELSIA 7
extracts of this material yielded alkaloid-positive precipitates based on six dif-
ferent precipitating agents. In addition he obtained a positive test for phyto-
steroids 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 Karolinska Institute!, Stockholm, Sweden. Mr. Lindgren (per-
sonal communication, 1973) prepared methanol extracts of the samples, which
were examined by thin-layer chromatography. He also performed a complete
alkaloid extraction on the material, followed by gas and thin-layer chromatog-
raphy on the extracts. Mr. Lindgren was unable to detect the presence 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 pharmacologic activity have
proved to be similarly difficult (Plowman, 1977).
3. Brunfelsia rupestris Plowman, sp. nov. Sect. Franciscea. Figure 3.
Frutex dense ramosus. Folia ad apicem ramulorum subcongesta, breviter petiolata, lan-
ceolata vel oblonga, apice acuta, plerumque obtusa vel emarginata, margine valde revoluta,
basr cuneata, superne glabra, nitida, subtus puberula, admodum coriacea, nervis later-
alibus 4-5 indistinctis. Inflorescentia in ramulis hornotinis terminalis, pedunculo brevi
pubescente instructa, floribus violaceis 2-5 (8). Pedicellus glaber. Calyx tubuloso-campa-
nulatus, teres, glaber, dentibus subaequalibus, ovatis, acutis. Corollae tubus glaber, limbo
patente, lobis subaequalibus, rotundatis. Capsula in calyce inclusa, ovoidea-globosa, api-
culata, laevis.
A Brunfelsia brasiliensi foliis congestis, coriaceis, valde revolutis, sparse pubescentibus,
nervis lateralibus 4-5 indistinctis, et pedicellis calycibusque glaberrimis differt.
Much-branched shrub 1-2 m tall. Bark on branches yellowish to reddish brown, cracked
longitudinally and transversely, exfoliating in thin flakes. Branchlets about 2 mm in di-
ameter, glabrous or puberulent. Leaves usually congested near branch tips, short petiolate,
blade lanceolate to oblong, apically acute, sometimes blunt or emarginate, strongly revolute
at margin, basally cuneate, 10-40 mm long, 6-17 mm wide, glabrous above, puberulent
beneath with short curved glandular or eglandular hairs, thickly coriaceous, shiny, medium
green above, dull beneath, drying ochraceous brown, 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. Peduncle 1-5 mm long, short-
branched, the branches articulating with the pedicels, persistent, puberulent. Bracteoles
linear-lanceolate or cymbiform, 1-6 mm long, sparsely villous, ciliolate at margin, ca-
ducous. Flowers deep violet with a white "eye" at orifice, fading to lavender with age.
Pedicel 4-12 mm long, 1-1.5 mm in diameter, glabrous. Calyx tubular-campanulate, terete,
10-15 mm long, 3-6 mm in diameter, glabrous, light green, teeth subequal, 2-3 mm long,
ovate to triangular, apically acute, the tip itself sometimes blunt or truncate. Corolla tube
about IVz times as long as calyx, straight, 15-18 mm long, 1.5-2 mm in diameter, glabrous,
orifice 2-3 mm in diameter; limb spreading, slightly thickened at mouth of tube, 15-22
mm in diameter, lobes subequal, rounded, 5-10 mm long. Stamens inserted in upper part
of corolla tube; filaments strap-shaped, the longer anterior pair apically incurved, included,
2-3 mm long, the shorter posterior pair 1-2 mm long; anthers orbicular-reniform, 1.5 mm
in diameter. Ovary ovoid, 1.5 mm long; stigma briefly bifid, 1 mm long. Capsule enclosed
by persistent calyx, ovoid to globose, apiculate at apex, 16 mm long, 14 mm in diameter,
smooth, pericarp thin, crustaceous, dry at maturity. Seeds oblong-ellipsoid, somewhat
angular, 5 mm long, 2.5-3 mm in diameter, reticulate-pitted, brown. Embryo straight, 4
mm long; cotyledons ovate, 2 mm long.
Type. — BRAZIL. State of Minas Gerais: Estrada Diamantina a Corinto ate 20
km. Campo rupestre e cerrado. Arbusto ate 2 m, flores roxas. 12 Jan. 1976. G. /.
Shepherd, J. B. de Andrade, L. S. Konoshita & J. Y. Tamashiro 3935 (holotype, UEC
11912; isotypes, F 1877095, NY, UEC 11912A).
FIELDIANA: BOTANY
BRUNFELSIA
rupest ris
Plowman
5 cm.
FIG. 3. Brunfelsia rupestris Plowman. 1, flowering branch; 2, fruiting branch; 3, excised
apical portion of corolla tube showing anthers and stigma. (1 and 3 drawn from Shepherd
et al. 3935, 2 from Ferreyra 887.)
Etymology. — From Latin rupestris, "rock-dwelling, of rocks," referring to the
habitat preference of the species for rocky outcrop areas.
Distribution. — Restricted to the "campo rupestre" community in the Serra do
Espinhago, Minas Gerais, Brazil.
Additional specimens studied.— BRAZIL. MINAS GERAIS: Serra do Espinhago,
Municipio Diamantina: Guinda, 5 Nov. 1937, Mello Barreto 9479 (F, RB), altitude
1,300 m, 14 Nov. 1971, Hatschbach & Pelanda 27941 (GH); ca. 25 km S.W. of
Diamantina on road to Gouveia, altitude 1,300 m, 16 Jan. 1969, Irwin et al. 22077
(F, MO, NY, UB); ca. 12 km S.W. of Diamantina, altitude 1,350 m, 23 Jan. 1969,
PLOWMAN: FIVE NEW SPECIES OF BRUNFELSIA 9
Irwin et al. 22465 (NY, UB); Serra do Cruzeiro, 5 Nov. 1979, V. F. Ferrrira et al.
887 (RB).
Brunfelsia rupestris is a very distinct species that is restricted to the campos
rupestres at higher elevations in the Serra do Espinhac.o of central Minas Gerais.
It is most closely related to and probably derived from the widespread and
polymorphic species B. brasiliensis (Spreng.) Smith & Downs. Brunfelsia rupestris
differs in having the leaves more congested, thickly coriaceous, strongly revolute
at the margins, sparsely pubescent and with only four to five lateral nerves, and
in having glabrous pedicels and calyces.
4. Brunfelsia burchellii Plowman, sp. nov. Sect. Guianenses. Figure 4.
Frutex. Ramuli glabri, nitidi, ochracei vel brunnei. Folia breviter petiolata, ovata vel
ovato-lanceolata, apice acuminata, acumine longo acutissimo saepe falcato instructa, mar-
gine ciliolata, basi rotundata vel obtusa, plerumque abrupte acuminata, glabra, nervis
lateralibus 5-9, valde arcuatis. Inflorescentia in ramulis hornotinis terminalis, pedunculo
brevi instructa, floribus 1-3. Pedicellus glaber vel sparse glanduloso-puberulus. Calyx
tubulosus vel tubuloso-campanulatus, sparse glanduloso-puberulus vel glaber, dentibus
subaequalibus triangulari-ovahs acutis vel acuminatis. Corollae tubus glaber, limbo pa-
tente, lobis subaequalibus, late rotundatis. Fructus seminaque ignota.
A Brunfelsia guianensi foliis ovatis acutissime acuminatis basi rotundatis, calycibus
tubuloso-campanulatis, pedicellis et calycibus longioribus differt.
Shrub. Mature branchlets spreading, 2-3 mm in diameter, with shiny yellowish to dark
brownish, longitudinally cracked bark. Leaves scattered along stem, short petiolate, blade
broadly to narrowly ovate, apically acuminate with a long, pointed, often falcate acumen,
minutely ciliolate at margin, basally rounded to obtuse, sometimes abruptly acuminate,
70-180 mm long, 30-75 mm wide, glabrous on both sides, firmly membranaceous to
chartaceous, medium green above, paler green beneath, dull or somewhat shiny above,
dull beneath, 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. Inflorescence terminal on current year's branchlets, very briefly pedunculate,
with 1-3 flowers. Peduncle 1-4 mm long, persistent, sparsely glandular-pubescent, gla-
brescent. Bracteoles 1-3 per flower, linear to lanceolate, apically acuminate, 4-20 mm long,
glabrous or villous at nerves and margin, caducous. Flower color unknown. Pedicel 4-8
mm long, 1 mm in diameter, sparsely glandular-pubescent, glabrescent. Calyx tubular or
tubular-campanulate, terete, 11-20 mm long, 5-9 mm in diameter, with scattered glandular
hairs, glabrescent, striately nerved, teeth unequal, 2-6 mm long, triangular-ovate, apically
acute to acuminate, minutely gland-tipped. Corolla tube ll/2 to 2 times as long as calyx,
straight, 24-28 mm long, 1.5-2 mm in diameter, glabrous or bearing few scattered glandular
hairs, orifice 5 mm in diameter; limb spreading, 18-30 mm in diameter, lobes subequal,
rounded, overlapping at the lateral margins. Stamens inserted in upper part of corolla
tube; filaments straplike, 1 mm wide, the longer anterior pair 3.5-6 mm long, apically
slightly incurved or suberect, the shorter posterior pair 3-4 mm long; anthers orbicular-
reniform, 1 mm in diameter. Ovary conical-ovoid, 1.5 mm long; style broadened and
incurved at apex, 20-21 mm long; stigma included between pairs of anthers, briefly bifid,
1 mm long, upper lobe slightly larger. Fruit and seed unknown.
Type. — BRAZIL. State of Goias: Porto Real (now Porto Nacional), ford of Igar-
ape, 1828-1830, W. /. Burchell 8527 (holotype, P; isotypes, K, L).1
Etymology. — Named in honor of the collector William John Burchell, British
botanist who collected extensively in Brazil in the early 19th century.
Distribution. — Basin of the upper Rio Tocantins (Goias State) and adjacent Ma-
ranhao State, Brazil.
!A11 Burchell localities are taken from his itinerary published by L. B. Smith and R. B.
Smith, Phytologia, 14: 492-506. 1967.
10
FIELDIANA: BOTANY
BRUNFELSIA
bu rebel Hi Plowman
FIG. 4. Brunfelsia burchellii Plowman. Flowering branch. (From Burchell 8527.)
Additional specimens studied.— BRAZIL. GOlAS: Porto Real, Porto Real to Igar-
ape, 1828-1830, Burchell 8415 (L), about village, Burchell 8494 (K), at entrance to
village, Burchell 8653 (K); "about the Manga" (cf. Rio dos Mangues, where it
meets the Tocantins), Burchell 8752 (GH). MARANHAO: Island of Sao Luiz,
Estrada do Barreto, Feb.-March 1939, Frdes 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 Fr6es
from Sao Luiz Island, more than 1,000 km north of Porto Nacional, is tentatively
PLOWMAN: FIVE NEW SPECIES OF BRUNFELSIA 11
assigned to B. burchellii but lacks mature flowers or fruits and differs somewhat
in leaf venation.
Brunfelsia burchellii appears to be most closely related to B. guianensis Benth.
from which it differs by the ovate, acuminate leaves, longer tubular calyx, and
the broadly rounded corolla lobes. With B. guianensis, B. burchellii is provisionally
placed in Sect. Guianenses based on the gradually dilated and not apically con-
stricted corolla tube.
Two additional collections with ovate acuminate leaves appear to be related
to B. burchellii but cannot be placed with certainty. One of these, Ducke s.n. (RB
18141), was collected at Braganga in Para and bears small, white, terminal, sol-
itary 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 persistent, accrescent calyces.
5. Brunfelsia clandestina Plowman, sp. nov. Sect. Guianenses. Figure 5.
Frutex vel arbor parva. Ramuli glabri, subnitidi, cinerascentes vel obscure brunnei. Folia
breviter petiolata, elliptica vel oblongo-lanceolata, rarius lanceolata vel obovata, apice acuta
vel acuminata, apice ipso obtusa, basi cuneata vel obtusa, glabra, raro sparse in costis
abaxialibus puberulenta, nervis lateralibus 6-8, rectis, divaricatis. Inflorescentia in ramulis
hornotinis juvenilibus vel maturis terminalis vel subterminalis, plerumque pedunculo brevi
instructa, floribus uni- vel biflora. Pedicellus glaber, post anthesin ad apicem lenticellis
verrucosis instructus. Calyx tubulosus vel tubuloso-campanulatus, glaber, dentibus sub-
aequalibus, triangulari-ovatis, apice acutis vel acuminatis, apice ipso minute glanduloso-
papilloso, post anthesin persistens, demum nitidus, coriaceus, striato-nervatus. Corollae
tubus glaber, raro sparse glanduloso-puberulus, limbo inclinato patente, lobis subaequa-
libus aliquantum accrescentibus, late obovatis vel rotundahs. Capsula sicca, ovoidea vel
subglobosa, apice rotundata, breviter apiculata, laevis, nitida, pericarpio tenui crustaceo.
Semina oblongo-reniformia, teretia vel paulo applanata, reticulato-foveolata, obscure brun-
nea.
A Brunfelsia martiana foliis multo minoribus, nervis paucis et inflorescentiis uni- vel
bifloris differt.
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. Branchlets 1.5-2 mm in diameter, glabrous, grayish to dark reddish brown, more
or less shiny, cracked longitudinally. Leaves short petiolate, the blade elliptic to oblong-
lanceolate, rarely lanceolate or obovate, apically acute to acuminate, the apex itself obtuse,
basally acute or obtuse, 35-120 mm long, 15-50 mm wide, glabrous or rarely sparsely
puberulent on the costa beneath, chartaceous to subcoriaceous, rarely coriaceous, medium
green above, somewhat lighter green beneath, shiny on both surfaces, 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 2-6 mm long, glabrous or with
scattered glandular hairs. Inflorescence terminal or subterminal on mature or newly formed
twigs of current season, with or without a short peduncle, one- or two-flowered. Bracteoles
none to 3, linear or cymbiform, truncate at apex, 2-6 mm long, sparsely pubescent or
glandular-pubescent, caducous. Flowers white. Pedicel 3-9 mm long, 1 mm in diameter,
glabrous, becoming thicker, warty-lenticellate in fruit, to 3 mm in diameter. Calyx tubular
or tubular-campanulate, terete, 8-16 mm long, 3-7 mm in diameter, glabrous, light green,
membranaceous, teeth subequal, 2-5 mm long, triangular-ovate, apically acute or acu-
minate, the apex itself blunt and minutely glandular-papillose; calyx in fruit persistent,
11-16 mm long, shiny, coriaceous, striately nerved, tightly enclosing basal half of capsule.
Corolla tube 1.5 to 2 times as long as calyx, straight, 18-25 mm long, 1-3 mm in diameter,
glabrous, rarely with a few glandular hairs, orifice 3-5 mm across; limb spreading, inclined,
15-26 mm in diameter, lobes subequal, expanding somewhat with age, broadly obovate
to rounded, 6-10 mm long. Stamens inserted in upper part of corolla tube; filaments strap-
shaped, 0.6-1 mm wide, the longer anterior pair incurved at apex, included, 3-5 mm long,
the shorter posterior pair 1.5-3 mm long; anthers orbicular-reniform, 1 mm in diameter.
12 FIELDIANA: BOTANY
Ovary conical-ovoid, 1-1.5 mm long, 0.8-1.2 mm in diameter, with about 10-12 ovules
per locule; style incurved at apex, 16-20 mm long; stigma briefly bifid, upper lobe slightly
larger, 1-1.5 mm long. Capsule dry at maturity, ovoid to subglobose, apiculate, 13-20 mm
long, 13-15 mm in diameter, smooth, shiny, dark green at maturity; pericarp thin, 0.5-1
mm thick, crustaceous, with about 9-15 seeds per capsule. Seeds oblong-reniform, terete,
or somewhat flattened on one side, 5-7 mm long, 2.5-3 mm in diameter, reticulate-pitted,
dark brown. Embryo straight, 3-6 mm long; cotyledons ovate to elliptic, 1-2 mm long,
radicle 2-4 mm long.
Type. — BRAZIL. State of Bahia: Municipio Itapebi, Fazenda Lombardia, BR
101 ao lado leste. Arbusto de 3 m de altura. Flor branca. Capoeira. 12 Aug. 1971.
I. S. dos Santos 1777 (holotype, CEPEC 7086; isotype, F 1849031).
Etymology. — From Latin dandestinus, "secret" or "hidden," referring to the fact
that the species remained unrecognized for over a century owing to its superficial
resemblance to Brunfelsia uniflora.
Distribution. — States of Bahia and Espiritu Santo, Brazil.
Additional specimens studied. — 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 Sincora, Brejao a Iracema, 17 Feb. 1943, Froes 20210
(IAN, NY, US); Municipio Jaguaquara, Jaguaquara a Apuerama, 4 Oct. 1972,
Pinheiro 1980 (CEPEC, F, NY); Municipio Itacare, Itacare-Ubaitaba, 14 April 1980,
dos Santos 689 (CEPEC, F); rodovia BA 654, km 6 ao oeste de Itacare, approx. 14°
18' S., 39° 02' W., altitude ca. 60 m, 12 April 1980, Plowman, Mattos Silva & dos
Santos 10066 (CEPEC, F, four duplicates to be distributed), Plowman, Mattos Silva
& dos Santos 10087 (CEPEC, F, 18 duplicates to be distributed); Municipio Itambe,
Itambe, 24 Nov. 1942, Froes 20067 (IAN, NY); Municipio Itaju do Colonia, 12 km
da estrada em diregao a Feirinha ao lado oeste, margem esquerda do Rio Corro,
23 Oct. 1969, dos Santos 433 (CEPEC, F); Municipio Belmonte, Estagao Experi-
mental Gregorio Bondar, km 58 da rodovia Belmonte/Itapebi, 16 May 1979, Mattos
Silva et al. 357 (CEPEC); Municipio Santa Cruz de Cabralia, Reserva Biologica
do Pau-Brasil, 18 Sept. 1971, dos Santos 1964 (CEPEC, F), cerca de 16 km a oeste
de Porto Seguro, 21 March 1978, Mori et al. 9775 (CEPEC, F), antiga rodovia que
liga a Estagao Ecologica de Pau-Brasil a Santa Cruz, 5-7 km ao N.E. da Estagao,
ca. 12 km ao N.W. de Porto Seguro, 16° 23' S., 39° 8' W., ca. 80-100 m altitude,
5 July 1979, Mori et al 12082 (CEPEC, F, US); Municipio Guaratinga, rodovia
Guaratinga/Sao Paulinho, km 25, 2 April 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). ESPIRITU SANTO: Linhares, Vale do Rio Doce, km
6 da rodovia BR 101, lado sul, 30 Sept. 1971, dos Santos 2015 (CEPEC, F, NY).
Brunfelsia clandestina was first collected by the Swiss collector Blanchet in the
state of Bahia over 100 years ago. Following earlier authors, I originally assigned
Blanchet's several collections of the species to Brunfelsia uniflora (Pohl) D. Don,
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. Recently, several excellent collections of B. clandestina
have been made by the staff at the CEPEC herbarium, Itabuna, Bahia. Accom-
panied by important field notes, these collections provided the basis for de-
scribing the plant as a distinct species of Brunfelsia Sect. Guianenses.
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-
BRUNFELSIA clandestina Plowman
FIG. 5. Brunfelsia clandestina Plowman. 1, flowering branch; 2, fruiting branch; 3, excised
apical portion of corolla tube showing anthers and stigma; 4, adaxial view of corolla limb;
5, seed. (1 drawn from dos Santos 1777, 2 from Pinheiro 1980, 3 and 4 from Pinheiro 2086
and 5 from dos Santos 1964.)
13
I
I
03
D
-S
I
CO
O
H
I
PLOWMAN: FIVE NEW SPECIES OF BRUNFELSIA 15
pears to be most closely related to B. martiana Plowman, a species that also
grows in the moist forests of Bahia. Brunfelsia clandestine! differs in having much
smaller leaves with fewer lateral nerves and one- or two-flowered inflorescences
that are borne terminally. In dried specimens, B. dandestina may also be confused
with B. uniflora, a species known from Bahia but belonging to a different section
of the genus (Sect. Frandscea). Brunfelsia dandestina 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 flow-
ers of B. dandestina are white; those of B. uniflora are violet fading to white with
age.
Brunfelsia dandestina grows in the moist coastal forests of southern Bahia and
Espiritu 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, interior part of Bahia was made in 1943.
ACKNOWLEDGMENTS
Research reported in this paper was supported in part by a National Institutes
of Health Training Grant (T T01 GM 00036-13, Harvard University) and by a
National Science Foundation Evolutionary Biology Training Grant (GB 7346,
Reed Rollins, Principal Investigator, Harvard University). Publication was sup-
ported in part by a grant from the National Institutes of Drug Abuse (1-R01 DA
02210-01, R. E. Schultes, principal investigator). This support is gratefully ac-
knowledged. I am also grateful to Field Museum of Natural History for financial
assistance for field work in Brazil in 1980.
I would like to thank the following persons who assisted in the collection,
preparation, or forwarding of herbarium materials for this study: Pedrito Silva
(Salvador, Brazil), Paulo Alvim, Luis A. Mattos Silva, Sergio da Vinha and Talmon
S. dos Santos (CEPLAC, Itabuna, Brazil), Scott Mori (New York Botanical Gar-
den), Craig Greene (Harvard University), and Julian A. Steyermark and Carlos
Blanco (Caracas, Venezuela). I would also like to thank the curators of the
following herbaria who kindly lent specimens cited in this work: BM, BR, C,
CEPEC, ECON, F, G, GH, IAN, K, L, LE, MG, MY, NY, P, RB, S, UB, UC, UEC,
US, YEN.
I am grateful to John Leary and Jan-Erik Lindgren, who performed the chem-
ical analyses, and to Bo Holmstedt for authorizing phytochemical work on Brun-
felsia in his laboratories at the Karolinska Institutet, Stockholm. I also wish to
Jhank Peter Paterson and Joan Ulrich, volunteers at Field Museum, who prepared
the line drawings for Figures 1 and 5 and Figures 3 and 4, respectively; to Rolf
Singer who corrected the Latin diagnoses; and to Michael Nee and Julian Stey-
ermark who offered comments on the manuscript.
LITERATURE CITED
BROWN, K. S., JR. 1979. Ecologia Geografica e Evolucao nas Florestas Neotropicais. Thesis.
Institute de Biologia, Universidade Estadual de Campinas, Campinas, Sao Paulo, Brazil.
LEARY, J. 1973. Letter to the author. March 22, 1973.
LINDGREN, J.-E. 1973. Letter to the author. March 6, 1973.
MORI, S. A., AND L. A. MATTOS SILVA. 1979. The Herbarium of the "Centro de Pesquisas
do Cacau" at Itabuna, Brazil. Brittonia, 31: 177-196.
16 FIELDIANA: BOTANY
PLOWMAN, T. 1973. The South American Species of Brunfelsia (Solanaceae). Doctoral Dis-
sertation, Harvard University, Cambridge, Mass.
1977. Brunfelsia in Ethnomedicine. Bot. Mus. Leafl. Harvard Univ., 25:
289-320.
_. 1979. The genus Brunfelsia: a conspectus of the taxonomy and biogeography,
pp. 475-491. In Hawkes, J. G., R. N. Lester, and A. D. Skelding (eds.), The Biology and
Taxonomy of the Solanaceae. Linnean Society Symposium Series 7, London, Academic
Press.
PRANCE, G. T. 1973. Phytogeographic support for the theory of Pleistocene forest refuges
in the Amazon basin, based on evidence from distribution patterns in Caryocaraceae,
Chrysobalanaceae, Dichapetalaceae and Lecythidaceae. Acta Amazonica, 3: 5-27.
SMITH, L. B. 1962. Origins of the Flora of Southern Brazil. Contr. U.S. Natl. Herb., 35:
215-249.
STEYERMARK, J. A. 1968. Contribuciones a la flora de la Sierra de Imataca, Altiplanicie de
Nuria y region adyacente del Territorio Federal Delta Amacuro al sur del Rio Orinoco.
Acta Bot. Venez., 3: 49-175.
1979. Plant refuge and dispersal centers in Venezuela — Their relict and
endemic element, pp. 185-221. In Larsen, K., and L. Holm-Nielsen (eds.), Tropical
Botany. London, Academic Press.
URBAN, I. 1906. Vitae, itineraque collectorum botanicorum. In von Martius, C. P. F., (ed.),
Flora Brasiliensis, 1(1): 138.
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