Brittonia, 37(3), 1985, pp. 243-251.

9 1985, by the New York Botanical Garden, Bronx, NY 10458

NEW SPECIES OF HABRACANTHUS (ACANTHACEAE)

FROM ECUADOR

DIETER C. WASSHAUSEN

Wasshausen, Dieter C. (Department of Botany, Smithsonian Institution, Wash-

ington, DC 20560). New species of H a b r a c a n t h u s (Aeanthaceae) from Ecuador.
Brittonia 37:243-251.1985.--Four new Ecuadorian species o f H a b r a c a n t h u s , H .
asplundii from Pichincha, H. laxus and H. lugonis from Tungurahua, and H.
luteynii from Napo, are described, illustrated, and compared with their closest
relatives.

T h e r e are a p p r o x i m a t e l y 4 5 species o f H a b r a c a n t h u s in Mexico, Central Amer-

ica, a n d A n d e a n S o u t h A m e r i c a . M o s t are l i m i t e d to wet v i r g i n forests at e l e v a t i o n s
b e t w e e n 1700 a n d 2 8 0 0 m e t e r s . D u e to the superficial s i m i l a r i t y o f m a n y o f the
species a n d the fact t h a t they are p o o r l y k n o w n a n d o f e x t r e m e l y local d i s t r i b u t i o n ,
the t a x o n o m y o f t h e g e n u s is r a t h e r difficult. A n a t t e m p t to t r e a t the g e n u s for
the F l o r a o f E c u a d o r has r e s u l t e d i n t h e r e c o g n i t i o n o f f o u r p r e v i o u s l y u n d e s c r i b e d
species b a s e d o n r e l a t i v e l y r e c e n t c o l l e c t i o n s .

K e y to the E c u a d o r i a n species o f H a b r a c a n t h u s
1 Calyceswithout gland-tipped trichomes.
2 Corolla deep maroon, glabrous, 2.5 cm long, upper lip linear, 2 mm wide, tip recurved or
curled; stamens exserted 15 mm beyond mouth of corolla ........................................................ 1. H. l u t e y n i i
2 Corolla yellow, pilose, 1.8 em long, upper lip ovate, 2.75 mm wide, obtuse; stamens
exserted 6 mm beyond mouth of corolla ..................................................................................................... 2. H. a s p l u n d i i
l Calycesbearing at least a few gland-tipped trichomes.
3 Panicles rather lax and open, the branches of the inflorescence usually once-forked ...............
........................................................................................................................................................................................................................... 3. H. l a x u s
3 Panicles dense and conical, at least the terminal one, the branches of the inflorescence
more than once-forked.
4 Corolla densely pilose ................................................................................................................................................................ 4. H . lugonis
4 Corolla glabrous.
5 Leaf blades ovate, to 9 cm wide; corolla dark reddish-violet; calyx segments linear,
6.5 mm long .............................................................................................................................................................................. 5. H. aglaiis
5 Leaf blades oblong-ovate or oblong-elliptic, to 7 cm wide; corolla bright red; calyx
segments lanceolate, 4 mm long .......................................................................................................... 6. H. s a n g u i n e u s

Habracanthus luteynii W a s s h a u s e n , sp. n o v . (Fig. 1)

Herba suffruticosa subsucculenta, caulibus teretiusculis, glabris; foliorum lamina elliptica vel ovata,
breviter acuminata, basi cuneata, utrinque glabra vel subtus in costa et venis lateralibus parce puberula;
paniculae floriferae terminales, densae, ramis subhelicoideis, 1-furcatis; paniculae inferiores parvae,
forsan steriles, apetalae, calycibus reductis, parvis; bracteae ramos panicularum subtendentes cordatae,
rotundatae et apiculatae, rhachidibus et rhachillis parce pilosis, bracteis ultimis lanceolatis; calycis
segmenta elliptica-oblonga, glabra, subtruncata et apiculata, dilute 3- vel 5-nervata; corolla atrocas-
tanea, glabra, tubo anguste infundibuliformi,labio superiore lineari, apice recurvato vel crispato, labio
inferiore patulo, ovato, trilobato, lobis obtusis; stamina exserta, glabra.
Suffrutescent, s o m e w h a t s u c c u l e n t h e r b to 1.5 m tall; s t e m s subterete, g l a b r o u s ,
cystoliths n u m e r o u s a n d parallel; l e a f b l a d e s elliptic to o v a t e , 9 - 1 2 c m long a n d
3 - 5 . 5 c m wide, s h o r t - a c u m i n a t e , c u n e a t e at base or the s m a l l e r u p p e r m o s t b l a d e s
r o u n d e d to o b t u s e , r a t h e r t h i n , e n t i r e or u n d u l a t e , b o t h surfaces g l a b r o u s except
costa a n d l a t e r a l v e i n s , these s p a r i n g l y p u b e r u l o u s w i t h a p p r e s s e d or a s c e n d i n g
septate hairs, costa, lateral v e i n s ( 6 - 9 pairs) a n d v e i n l e t s p r o m i n e n t b e n e a t h , r a t h e r
244 BRITTONIA [VOL. 37

i l1 cm J

]
1 cm

F r JI 1 crn

]
1cm
~ $ ~ 1m~crn
cm
E ~

FIG. 1. A-D. Habracanthus luteynii (Luteyn et al. 8449). A. Flowering branch. B. Inflorescence
node with a three-flowered dichasium. C. Corolla and exserted stamens. D. Corolla expanded, showing
attachment o f filaments. E-I. Habracanthus laxus (Asplund 9781). E. Sterile branch. F. Inflorescence.
G. Calyx. H. Corolla and exserted stamens and style. I. Corolla expanded, showing attachment of
filaments.
1985] WASSHAUSEN" HABRACANTHUS 245

FIG. 2. SEM photomicrographs of Habracanthus pollen. A-B. Habracanthus luteynii (Luteyn et

al. 8449). A. Equatorial view, • 1500. B. Portion of surface, • C. H. laxus (Asplund 9781),
equatorial view, x 2500.

obscure above, cystoliths small and inconspicuous; petioles 1-3 c m long, glabrous
or sparingly puberulous, cystoliths minute, n u m e r o u s and parallel; flower-bearing
panicles terminal, 6-8 cm long and a b o u t 5 cm broad, dense; mostly sterile cymes
axillary to 2 cm long, branches o f terminal inflorescence subhelicoid, usually once-
forked; bracts subtending branches o f inflorescence cordate, 1-1.5 cm long, 1-1.5
cm wide, r o u n d e d and apiculate at apex; rachis and rachilla sparingly pilosulous;
pedicels 1-2 m m long, sparingly pilosulous; bracts subtending flowers lanceolate,
about 2 m m long and 1 m m wide near base; calyx 6 m m long, glabrous, segments
elliptic-oblong, 5 m m long, 1.5 m m wide, subtruncate and apiculate, faintly
3- to 5-nerved, ciliate, cystoliths obscure, calyces o f sterile lower axillary inflo-
rescence 1-2.5 m m long; corolla deep maroon, glabrous, 2.5 cm long from base
to tip o f upper lip, tube 5 m m broad near base, narrowed at 7 m m above base
to 2.5 mm, thence enlarged to 4.5 m m at mouth, upper lip linear, 13 m m long,
2 m m wide, the tip r e c u r r e d or curled, cucullate and entire, lower lip spreading,
246 BRITTONIA [VOL. 37

1 cm B .4 cm

~ c m

,4 ii
rj
D .15 cm

FIG. 3. A-E. Habracanthus asplundii (Asplund 17429). A. Flowering branch and leaf. B. Inflores-
cence node showing bracts, calyx, and capsule. C. Calyx. D. Corolla, stamens, and style. E. Corolla
showing attachment of filaments. F-I. Habracanthus lugonis (Lugo 1895). F. Flowering branch. G.
Calyx. H. Corolla, stamens, and style. I. Corolla expanded.
 1985] WASSHAUSEN: HABRACANTHUS 247

ovate, 12 m m long, 8 mm wide, rounded and 3-lobed at tip, lobes 0.5 mm long,
l mm wide, rounded; stamens exserted 15 m m beyond mouth of corolla, glabrous;
anthers linear, 5 m m long, 1 mm broad, muticous at both ends; pollen 2-porate,
bilateral, 47 x 53 tzm, pores more or less circular, nexine densely granular, sexine
tectate (Fig. 2); ovary glabrous; capsules not seen.
TYPE: ECUADOR. NAPO; Santa Barbara-La Bonita road, trail towards La
Bonita, ca 25-28 km S of Santa Barbara, along Rio Chingual, 2134-2348 m a l t ,
17 May 1982, J. L. Luteyn, H. Balslev & B. M. Boom 8449 (HOLOTYPE:US;
ISOTYPES[ A A U , C A S , C O L , GB, MO, NY, QCA).
Known only from the type locality.
Habracanthus luteynii superficially resembles H. floriferus Leonard, a Colom-
bian species, but it is easily distinguished from that species by its shorter (5 mm
long), elliptic-oblong, subrotundate, and apiculate calyx segments, smaller (2 mm
long) floral bracts, and shorter stamens which are exserted 15 m m beyond the
mouth of the corolla tube. In H. floriferus, the calyx segments are lanceolate, 7
mm long and acute, the floral bracts about 4 m m long, and the stamens are
exserted 25 m m beyond the mouth of the corolla tube.

Habracanthus laxus Wasshausen, sp. nov. (Fig. 1)

Suffrutex, caulibus teretiusculis, glabris; foliorum lamina elliptica vel ovata, breviter acuminata,
basi cuneata, utrinque parce pilosula, pilis praecipue in costa et venis lateralibus positis; paniculae
terminales et axillares, aliquanto densae, virides, ramis subhelicoideis, 1-furcatis; bracteae ramos
inflorescentiae subtendentes ovatae, acuminatae, cordatae et sessiles, rhachidi et rhachillis puberulis,
bracteolae subulatae, mucronulatae, glabrae vel parce pilosulae; calycis segmenta viridia, lanceolata,
infra glabra, pilosula ad apicem, pills glanduliferis intermixtis; corolla atrosanguinea, glabra, tubo
anguste infundibuliformi, labio superiore lineari, apice recurvato vel cuspidato, parce pilosulo, labio
inferiore patulo, ovato, trilobato, lobis obtusis; stamina exserta, glabra, antheris linearibus.
Subshrub about 1 m tall; stems subterete, glabrous, the cystoliths inconspicuous,
parallel and subpunctiform; leaf blades elliptic to ovate, I 1.5-13 cm long and 4-
6 cm wide, slenderly acuminate, more or less curved at tip, cuneate at base,
moderately thin, entire or lightly undulate, sparingly pilose above with flaccid,
septate hairs, obscurely pilosulous beneath, especially on costa, lateral veins (8-
10 pairs), and veinlets, these prominent both above and beneath, cystoliths slender
but prominent under a lens; petioles 3-5 cm long, sparingly pilose, cystoliths
minute, numerous and parallel; panicles green, rather lax, the terminal 15 cm
long and 11 cm broad, the axillary panicles to 9 cm long and 6 cm wide, the
branches of the inflorescence subhelicoid, usually once-forked; bracts subtending
branches of the inflorescence ovate, to 2.5 cm long and 1.4 cm wide, acuminate,
cordate and sessile at base, glabrous and ciliate, cystoliths purplish-punctate; rachis
and rachilla sparingly puberulous; pedicels to 3 mm long, sparingly pilose; bracts
subtending flowers subulate, 3 mm long and 0.75 m m wide, mucronulate at tip,
glabrous or rather sparingly pilosulous; calyx green, 5 mm long, glabrous below,
sparingly pilosulous toward tip, simple hairs intermixed with gland-tipped tri-
chomes, segments narrowly lanceolate, 4 m m long and 1 m m wide at base, acute;
corolla dark-red, glabrous except tips of lips (these sparingly pilosulous), 1.8 cm
long from base to tip of upper lip, tube 4.5 m m broad near base, narrowed to 2.5
mm at 6 m m above base, 4 mm broad at mouth, upper lip linear, 4 mm long,
1.5 m m wide, tip recurved or curled, cucullate and acute, lower lip spreading,
ovate, 9 mm long, 7-8 mm wide, rounded, 3-lobed at tip, lobes 1 mm long, 1.5
mm wide, rounded; stamens exserted about 12 mm beyond mouth of corolla,
glabrous; anthers linear, 3.5 mm long, 0.5 mm broad, muticous at both ends;
pollen 2-porate, subspheroidal, 50 • 46 #m, exine spinulose, spinulae widely
spaced (Fig. 2); style exserted about 1.5 cm beyond mouth of corolla; stigma
248 ~RITTONIA [VOL. 37

FIG. 4. SEM photomicrographs of Habracanthus asplundii pollen. A. Asplund 17429, equatorial

view, • 1900. B. Asplund 7374, equatorial view, x 1600.

minute; o v a r y glabrous; capsule clavate, glabrous, 18 m m long, 5 m m wide; seeds

wanting.
TYPE: E C U A D O R . TUNGURAHUA: Cordillera de Llanganates, valley o f Rio
Sangarinas (Desaguadero), Rio San Josr, 3 1 0 0 - 3 2 0 0 m a l t , 19 Jan 1939, E.
Asplund 9781 (HOLOTYPE: S).
Distribution. K n o w n only f r o m the type locality.
 1985] WASSHAUSEN" HABRACANTHUS 249

FIG. 5. SEM photomicrograph of Habracanthus lugonis (Lugo 1895) pollen. A. Equatorial view,
• 1650. B. Portion of surface, x 2500.

Habracanthus laxus is perhaps nearest in relationship to the rather widespread

H. sanguineus Nees but differs m a r k e d l y in that the latter species has a glabrous
corolla, 2-2.5 c m long, b o t h the u p p e r a n d lower corolla lips 12 m m long, a n d
the calyx segments 9 - 1 0 m m long. In contrast, H. laxus has a sparingly pilosulous
corolla, 1.8 c m long, an u p p e r corolla lip 4 m m long, a lower corolla lip 9 m m
long, and calyx segments only 5 m m long.

H a b r a e a n t h u s asplundii Wasshausen, sp. nov. (Fig. 3)

Herba vel suffrutex, caulibus subquadrangularibus, glabris vel parce pilosis; foliorum lamina late
elliptica vel ovata, graciliter acuminata, basi rotundata in petiolum decurrens, supra modice tomentosa
subtus parce puberula; paniculae terminales et axillares modice densae, bracteae ramos inflorescentiae
subtendentes conspicuae, foliaceae, orbiculares, sessiles vel subsessiles, rhachidi, rhachillis et pedicellis
pilosis, bracteis ultimis anguste lanceolatis pilosis; calycis segmenta anguste triangularia, infra glabra
parce pilosa ad apices; corolla lutea, pilosa, tubo basi angusto, supra abrupte dilatato, in fauce leviter
angustato, labio superiore ovato, erecto, apice obtuso, ciliato, labio inferiore apice trilobato, lobis
obtusis; stamina exserta, glabra, antheris utrinque muticis; ovarium glabrum.
Herbaceous or suffrutescent plant a b o u t 1 m tall; stems ascending, subquadran-
gular, glabrous or sparingly pilose n e a r tip; leaf blades broadly elliptical or ovate,
14 c m long, 8-8.3 c m wide, slenderly short-acuminate, short-cuneate (7 m m long)
f r o m a r o u n d e d base, thin, entire, m o d e r a t e l y t o m e n t o s e above, sparingly puber-
ulous beneath, the hairs confined chiefly to costa and lateral veins (about 10 pairs),
spreading or ascending, veinlets coarsely reticulate, these and the costa and lateral
veins rather p r o m i n e n t especially beneath, cystoliths p r o m i n e n t u n d e r a lens,
brownish; petioles 2 - 5 cm long and 1.75 m m thick, sparingly pilose, cystoliths
n u m e r o u s , conspicuous and parallel; flowers borne in m o d e r a t e l y dense, terminal
a n d axillary panicles, these to 30 c m long a n d 7 c m broad, axillary panicles 4 - 8
c m long; bracts subtending branches o f panicle conspicuous, leaf-like, orbicular,
sessile or subsessile, l o w e r m o s t pair 2.5 c m long and 3 c m wide, b e c o m i n g grad-
ually smaller t o w a r d tip o f panicle, all obtuse to acuminate, the pubescence similar
to that o f stem leaves; rachis pilose, the hairs spreading; rachilla a n d pedicels (to
250 BRITTONIA [VOL. 37

7 mm long and 0.5 mm thick) pilose with erect and ascending hairs; bracts
subtending flowers narrowly lanceolate, 2 m m long and 0.75 m m wide, blunt,
ciliate at tip, pilose; calyx 5 m m long, glabrous below, sparingly pilosulous toward
tip, segments narrowly triangular, 4 mm long and 1.5 m m wide at base, acute;
corolla yellow, 18 m m long, pilose with long spreading septate hairs, tube 2 m m
broad at base, at 4 m m above base abruptly enlarged to 7 mm, 4.5 mm broad at
throat, lips erect, subequal, 3 m m long, upper lip ovate, 2.75 m m wide, obtuse
and ciliate, lower lip 3-lobed, lobes 1 mm long and 1.5 m m broad, obtuse; stamens
exserted 6 m m beyond mouth of corolla, glabrous; anthers 3 mm long, 0.5 m m
thick, muticous at both ends; pollen 2-porate, subspheroidal, 40 x 38/~m, exine
spinulose, spinulae numerous and densely spaced (Fig. 4); style exserted 5 mm
beyond mouth of corolla, stigma minute; ovary glabrous; capsule clavate, glabrous,
18 m m long, 5 mm wide; seeds immature.
TYPE: ECUADOR. PICHINCHA: E1 Volante on road from Chiriboga to Santo
Domingo de los Colorados, 1900 m a l t , 22 Aug 1955, E. Asplund 17429
(HOLOTYPE: S).
Additional specimen examined: ECUADOR. PICHINCHA:Las Palmeras on road from Quito to Santo
Domingo de los Colorados, 1900 malt, 28 Jun 1939, E. Asplund 7374 (S).
Occasional, in shade along banks of rivulet, in Pichincha Province, Ecuador.
This species is readily distinguished by its long, narrow, moderately dense
terminal panicle, the conspicuous, leaf-like bracts subtending the branches of the
panicle, and the small (18 mm), yellow, pilose corolla. It is not very closely allied
to any of the known species of Habracanthus.

Habracanthus lugonis Wasshausen, sp. nov. (Fig. 3)

Herba vel suffrutex, caulibus ascendenfibus, subquadrangularibus, pilosis; foliorum lamina elliptica
vel anguste ovata, breviter aeuminata, basi euneata, utrinque modice pilosa; paniculae ovoidae, modice
densae, bracteis ramos panicularum subtendentibus eonspicuis, foliaeeis, cordatis; rhachidi dense
pilosa, rhachillis et pedicellis pilosis, braeteis ultimis anguste lanceolatis, pilosis; calyeis segmenta
viridia infra glabra dense pilosa ad apicem, pilis glanduliferis intermixtis; corolla rubra, dense pilosa,
tubo anguste infundibuliformi, labio superiore lineari, labio inferiore patulo, obovato, trilobato, lobis
parvis, obtusis; stamina exserta, rubra, glabra, antheris linearibus.
Herbaceous or suffrutescent plant about 1 m tall; stems ascending, subquad-
rangular, pilose, the hairs spreading, septate; leaf blades elliptic to narrowly ovate,
15-18 cm long, 6.5-8.5 cm wide, short-acuminate, cuneate at base, thin, entire,
both surfaces moderately pilose, especially costa and lateral veins (9-11 pairs),
these and reticulate veinlets moderately conspicuous, the hairs erect, septate,
cystoliths minute, sparse, inconspicuous; petioles 3.5-5 cm long and 1.75 m m
thick, pilose, the hairs spreading, septate interspersed with brownish glands, cys-
toliths inconspicuous; panicles ovoid, the terminal about 14 cm long and 5 cm
wide, moderately dense, the axillary panicles shorter, 3-7 cm long; bracts sub-
tending branches of panicle conspicuous, leaf-like, cordate, sessile or subsessile,
lowermost pair 1.5 cm long and 1 cm wide, becoming gradually smaller toward
tip of panicle, all short-acuminate, moderately puberulous, rachis densely pilose,
the hairs spreading, rachilla and pedicels (to 5 mm long and 0.5 m m thick) densely
pilose with spreading, septate hairs; bracts subtending flowers narrowly lanceolate,
4 mm long and 0.75 mm wide, blunt at tip, densely pilose; calyx green, 6.5 m m
long, glabrous below, densely pilose toward tip, hairs intermixed with gland-tipped
trichomes, segments narrowly lanceolate, 6 m m long and 1.25 m m wide at base,
acute; corolla red, densely pilose, 2.4 cm long, tube 4 m m broad near base,
narrowed at 7 mm above base to 2.5 mm, throat about 2 m m broad, upper lip
erect linear, 11 mm long, 1.25 m m wide, obtuse, lower lip spreading, obovate, 8
 1985] WASSHAUSEN: HABRACANTHUS 251

mm long, 6 m m wide, the tip rounded, 3-lobed, lobes small, 0.5 m m long, 1 mm
wide, rounded; stamens red, exserted about 15 mm beyond mouth of corolla,
glabrous; anthers linear, 4.5 mm long, 0.75 mm thick; pollen 2-porate, subspheroi-
dal, 42 ~tm in diam, exine spinulose, spinulae widely spaced (Fig. 5); style exserted
about 1 cm beyond mouth of corolla, stigma minute; ovary glabrous; capsule not
seen.
TYPE: ECUADOR. TUNGURAHUA: Colonia Regina, 5-6 km NW of Rio Verde,
15 Apr 1972, H. Lugo S. 1895 (HOLOTYPE: GB).
Additionalspecimenexamined:ECUADOR. NAPO:road from Baezato Tena, vicinityof Cosanga,
El Paso de Guacamayo,2500 malt, 17 Dec 1976, E. IV. Davis 528 (S).
Primary cloud forest in Northern Ecuador.
Habracanthus lugonis superficially resembles H. lamprus Leonard. However,
in H. larnprus, the leaf blades are ovate and to 11 cm long and the dark red
corollas are 1.5 cm long and glabrous. In H. lugonis, the leaf blades are elliptic
to narrowly ovate, 15-18 cm long, and 6.5-8.5 cm wide and the red corollas are
2.4 cm long and densely pilose.

Acknowledgments
I wish to thank Cathy Pasquale for preparing the excellent line drawings and
Dr. Stephen F. Smith and the staffofthe Smithsonian Institution SEM laboratory
for their continued help in preparing the high-quality pollen photomicrographs.

BOOK REVIEWS
Plant Variation and Evolution. Second edition. By D. Briggs and S. M. Walters.
Cambridge University Press, 32 East 57th Street, New York, NY 10022. ISBN
0-521-257069 (cloth), 0-521-276659 (paper). 1984. 412 pp. $59.50 (cloth),
$17.95 (paper).
One is often disappointed when a book appears in a second edition, and this
is the ease with the Briggs and Walters text. The 1969 edition (published by
Weidenfield and Nicolson) was a delightful little book, extremely reasonable in
price, a nice size to handle, and well-illustrated, as well as being an interesting
and useful text. In contrast, the second edition is larger in format, relatively
expensive, and rather poorly illustrated in some cases (e.g., figs. 4.6, 7.10, and
7.14). Of course the text has been updated, with a full 34 percent of the bibli-
ography being post-1969 publications. However, the new material is not well
integrated into the text, and it is difficult to recommend the book with any sin-
cerity.--P. MIcK RICHARDSON, New York Botanical Garden.
The Chemistry of Allelopathy: Biochemical Interactions among Plants. Edited by
Alonzo C. Thompson. American Chemical Society, 1155 Sixteenth Street, N.W.,
Washington, DC 20036. ISBN 0-8412-0886-7. 1984. 470 pp. $79.95 (cloth).
This book is based on a symposium sponsored by the American Chemical
Society in April 1984. It contains 31 papers, ranging from host selection by
parasitic angiosperms through soil-borne allelopathic chemicals to the synthesis
of strigol, a plant exudate stimulating the germination of Striga seeds. Two in-
teresting chapters cover alleloehemicals from fresh- and saltwater plants. Although
this book may be of little direct interest to taxonomists, every botanist will find
something of interest in many of the chapters.--P. MICK RICHARDSON,New York
Botanical Garden.