Download with descriptions of four new species

A synopsis of Costa Rican Ruellia (Acanthaceae),
with descriptions of four new species
LUCINDA A. MCDADE 1 ° 2 AND ERIN A. T RIPP 3
Department of Botany, Academy of Natural Sciences, 1900 Ben Franklin Parkway, Philadelphia, PA
19103, U.S.A.; e-mail: [email protected]. 'Current address: Rancho Santa Ana Botanic Garden,
1500 N. College Avenue, Claremont, CA 91711, U.S.A.; e-mail: [email protected].
3 Department of Biology, Duke University, Durham, NC 27708, U.S.A.; e-mail: erin.tripp@
duke.edu
1
Abstract. We describe four new species of Ruellia from Costa Rica; three from the
Osa Peninsula, Puntarenas province (R. exilis, R. mira, R. odorata), and one from
the southwestern corner of San Jose province (R. norvegigratiosa). Ruellia matagal-
pae is reported as a new record for the country. New combinations are made in Ruellia for the two Costa Rican species of Blechum (R. blechum and R. costaricensis).
Three species are reduced to synonymy, and another species is considered doubtfully
worthy of recognition. Notable range extensions are reported for two species. Corollas and fruits of all 22 Costa Rican species of Ruellia are shown and compared as
easily used identification aids. An informal guide to species groups and a dichotomous key to the 22 species are presented.
Key words: Acanthaceae, Costa Rica, Osa Peninsula, Puntarenas, Ruellia.
Resumen. Se describen y se ilustran cuatro especies nuevas para la ciencia de Ruellia
para Costa Rica; tres de la Peninsula de Osa, provincia de Puntarenas (R. exilis, R.
mira, R. odorata) y una del extremo suroeste de la provincia de San Jose (R.
norvegigratiosa). Reportamos a Ruellia matagalpae como especie encontrada en el
pais por primera vez. Se proponen nuevas combinaciones en Ruellia para las dos especies de Blechum que se encuentran en Costa Rica (R. blechum y R. costaricensis). Tres especies son reducidas a sinonimia y una cuarta especie es presentada como
dudosa. Se reportan extensions de la distribucion geografica para dos especies. Presentamos ilustraciones de las corolas y los frutos de las 22 especies costaricenses de
Ruellia; estas estructuras presentan caracteristicas muy (stiles para la identificacion.
Las 22 especies de Ruellia se contrastan en un gufa informal para grupos de especies
y en una clave dicotomica.
In studying plants and specimens as part of
our work to prepare the treatment of Acanthaceae for the Manual de Plantas de Costa
Rica (Hammel et al., 2004 and subsequent
volumes), we have made a number of discoveries in Ruellia L. With approximately 300
species, Ruellia is the second largest genus
(after Justicia L.) in the family. Here we describe and illustrate four new species; three
of these have been discovered as a result of
intensive collecting on the Osa Peninsula in
the last couple of decades, and the fourth was
collected only in 2005 from the extreme
southwestern corner of San Jose province.
The junior author's doctoral research on the
entire genus Ruellia gives us a broad geographic perspective on the genus and convinces us of the merit of describing this last
new species although it is known only from a
single collection. We also report a new
species record for Costa Rica, propose two
new combinations and place three previously
Brittonia, 59(3), 2007, pp. 199-216.
© 2007, by The New York Botanical Garden Press, Bronx, NY 1 045 8-5 126 U.S.A.
ISSUED: 27 September 2007
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BRITTONIA
recognized species in synonymy. Finally, we
provide figures that illustrate the diversity of
corolla and fruit morphology in Costa Rican
Ruellia; these structures present a number of
traits that are extremely useful in distinguishing species. We also present an informal
guide that uses easily observed traits to restrict the identification process to groups of
species, and a dichotomous key to all 22
species of Ruellia that are known to occur in
Costa Rica.
New Species
Ruellia odorata E. Tripp & McDade, sp.
nov. Type: Costa Rica. Puntarenas: Canton
de Osa, Peninsula de Osa, Reserva Forestal
Golfo Dulce, Rancho Quemado, fila ca.
3 km al sur, 8°43'00"N, 83°34'50"W,
200m, Jan 1993, A guilar 1565 (holotype:
INB; isotype: MO). (Figs. 1, 2A, 3A)
Species nova floribus albis aromaticis et calycibus
pilosus praedita, ob corollarn albam R. golfodulcensi
Durkee, R. palustri Durkee, R. praeclarae Stand!. et R.
tubiflorae H. B. & K. simulans, sed a R. golfoculdcensi
corolla longiore et caule piloso differt, a R. palustri et R.
praeclara calyce breviore differt, a R. tubiflora calyce
longiore differt.
Herbs to 1 m, stems erect, younger stems
subquadrangular and densely pilose, basal
portions of plant terete, less densely pilose,
trichomes perpendicular to stem or slightly
ascending, ca. 2mm long. Leaves with petioles to 1.9 cm long, pubescent with dense,
appressed trichomes; laminae oblanceolate to
elliptic, 8.5-20 x 2.5-8.3 cm, 2.9-3.1 times
longer than wide, apices acuminate, bases
acute, margins entire to repand, abaxial surfaces pubescent, mostly along veins, adaxial
surfaces glabrous to very sparsely pubescent,
cystoliths visible on both surfaces but more
obvious above, both laminal surfaces (especially developing leaves) with inconspicuous,
sessile, patelliform glands, drying pale to
dark red. Inflorescences of few flowered
cymes in axils of distal nodes, sometimes appearing terminal. Bracts elliptic, acute or
blunt at apex, 10-23 mm long x 2-5 mm
wide, pubescent with appressed trichomes of
longer eglandular hairs and shorter often
glandular hairs, with patelliform glands as
on leaves, bracts sometimes falling postanthesis, bracteoles not seen. Flowers sub-
[VOL. 59
sessile, aromatic. Calyx 10-18 mm x 1-2.5
(-3) mm, pubescent with eglandular trichomes to 1.5 mm long and shorter, often
glandular trichomes to 0.5 mm long, lobes
linear, with patelliform glands as on leaves.
Corolla white, outer surface sparsely villous,
trichomes crumpled, of multiple lengths,
often glandular, with patelliform glands as on
leaves, unexpanded portion of tube
3.5-4.5 cm long, expanded portion ca.
2-2.5 x 1.3-1.5 cm (pressed), lobes ca.
1.0-1.5 x 1.8-2.0 cm. Stamens included,
slightly didynamous, apex of anther of
shorter pair 52-54 mm from base of corolla,
apex of anther of longer pair 53-55 mm from
base of corolla, free portions of filaments
8-9 mm (shorter), 9-12 mm (longer) long,
fused portion of filaments 41-45 mm long,
fused filament sheath ("curtain") enclosing
nearly all of unexpanded portion of tube, anthers 3.5-5 mm long, rounded at base; styles
4.9-5.8 mm long, not extending beyond
mouth of corolla and approximately equal in
height to anthers, stigma bifid, dorsal lobe reduc,ed to three quarters the length of ventral
lobe. Fruits clavate, to 17 mm long x 5-7 mm
wide, stipe 5-8 mm long, seed bearing portion 8-11 mm long, velutinous with short trichomes, sometimes glandular. Seeds to 8,
4-5 mm in diameter, margins with hygroscopic trichomes.
Distribution and ecology. Ruellia odorata
is known only from the Osa Peninsula of
Puntarenas Province, including central portions of Reserva Forestal Golfo Dulce and
areas around and west of Rincon. Plants occur
in the understory of forests at 50-200 m elevation. Flowering specimens of R. odorata
have been collected December—March, fruiting
specimens in January. Nothing is known about
the biology of the species but its white, aromatic flowers with long tubes suggest hawkmoth pollination.
Etymology. Collectors indicate that the
flowers of R. odorata are "aromatica" and
"sweet smelling." Floral fragrance is unusual
among Acanthaceae and merits further study
in terms of floral phenology and pollinator
relationships.
Additional specimens examined. COSTA RICA.
Osa Peninsula, Rincon de Osa, 8°42"N,
83°31'W, 50-200m, near air field, in forest, Mar 1973,
PUNTARENAS:
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MCDADE & TRIPP: COSTA RICAN RUELLIA (ACANTHACEAE)
201
FIG. 1. Ruellia odorata. A . Habit. B. Close-up of stem showing trichomes. C. Calyx with close-up showing
trichomes. D. Androecium, ventral view with corolla dissected. E. Stigma and terminal portion of the style. F. Fruit.
(From Burger & Gentry 8915, F.). Illustration by Amanda Labadie.
Burger & Gentry 8915 (F, US); Osa Peninsula, Rincon
de Osa, 8°42N, 83°31'W, 50-200m, near airfield, Jan
1970, Burger & Liesner 7329 (F); Osa Peninsula,
Rincon de Osa, 8°42N, 83°31'W, 50-200m, moist
forests and areas of secondary vegetation, Jan 1970;
Burger & Liesner 7202 (NY); Canton de Osa, Rancho
Quemado, Rincon, 8°42'N, 83°34'W, 200 m, bosque primario, Dec 1990, Quesada 287 (INB, MO); Rincon,
Finca del Delfin, near Rancho Quemado, 8°42N,
83°34'W, 200m, Jan 1991, Quesada & ChavarrIa 414
(INB, MO).
Ruellia odorata most closely resembles R.
palustris Durkee but differs in having rela-
tively narrower leaves (ca. 3 x longer than
wide vs. usually < 2.5 x longer than wide in
R. palustris), a shorter and narrower calyx
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[VOL. 59
Three other white-flowered Ruellia species
occur on the Osa Peninsula. Plants of R.
praeclara are much larger (to 2 m), have
longer leaves (to 30 cm), and a much longer
calyx (30-55 mm). Plants of R. golfodulcensis
have nearly glabrous stems and shorter corollas, with the tube to only 3.0 cm long. Finally,
plants of R. tubiflora are shrubs or treelets
with spicate inflorescences, large leaf-like
bracts and very short calyces (8-10mm).
FIG. 2. Scanning electron micrographs of pollen of
four new species, showing the spheroidal shape and
distinctly reticulate surfaces typical of Ruellia species
(Daniel, 1998). A . Ruellia norvegigratiosa (Santamaria
932, INB). B. Ruellia mira (Herrera 4860, MO). C.
Ruellia odorata (Quesada 287, MO), one of three pores
visible. D. Ruellia exilis (Herrera 4548, MO), one of
three pores visible. Scale bar =20µm.
(10-18 mm long x l-2.5 [-3 mm] wide vs.
25-30 mm long x 4-5 mm wide in R. palustris), and a shorter fruit (to 17 mm long vs.
20-23 mm long in R. palustris); further, R.
palustris is not known from the Pacific slope.
FIG. 3. Scanning electron micrographs of glands on
abaxial leaf surfaces of plants of four new species; glands
are sessile and patelliform. A . Ruellia norvegigratiosa
(Santamaria 932, INB), showing droplets of an
unknown substance. B. Ruellia exilis (Herrera 4548,
MO). C. Ruellia mira (Herrera 4860, MO). D. Ruellia
odorata (Quesada 287, MO). Scale bar for A and
B= l O pm. Scale bar for C and D = 20µm.
Ruellia exilis McDade & E. Tripp, sp. nov.
Type: Costa Rica. Puntarenas: Peninsula de
Osa, Reserva Forestal Golfo Dulce, Cerro
de Oro, bosque secundario, en charral,
8°33'35"N, 83°29'55"W, 110m, Mar 1995,
A lfaro 138 (holotype: INB; isotype: PH).
(Figs. 2B, 3B, 4)
Species nova foliis lanceolatis 6.5-plo longioribus et
inflorescentia longipedunculata, oh inflorescentiam et
pedunculos longos R. pittieri Lindau, R stemonacanthoidi (Oerst.) Hemsl., R. norvegigratiosae McDade &
E. Tripp, R. malacospermae Lindau simulans, sed ab
omnibus earum foliis angustis, corolla brevi, fructibus
brevibus differt.
Herbs 0.2-0.4 m tall, stems erect, upper
portions quadrangular, glabrous to covered in
short (< 0.25 mm) erect or upwardly appressed trichomes. Leaves with petioles
5-20 mm long, pubescent as stems; laminae
narrowly lanceolate, 3.3-15 x0.4-2.5(-3) cm,
at least 6.5 times longer than wide, apices
and bases attenuate, margins entire to repand,
abaxial surfaces pubescent along veins but
otherwise glabrous, adaxial surfaces glabrous,
both surfaces with visible cystoliths and inconspicuous, patelliform glands, drying dark
red. Inflorescences numerous, from axils of
distal leaves, borne on mostly glabrous peduncles 2.9-8.0(-12.5) cm long, peduncles
diffusely branching, nodes with either paired
branchlets or one bud differentiating as a solitary flower, peduncles and inflorescence
branches with dense cystoliths. Bracts paired
at each node, narrowly elliptic to linear, most
proximal pair 0.9-2.5 x 1.5-2.0 mm, becoming
reduced distally, some flowers subtended by
paired bracteoles in addition to reduced inflorescence bracts, these narrowly subulate,
<1 mm long. Flowers sessile or on pedicels
to 4 mm long. Calyx 2.5-3 mm, fused portion
0.5-1.0 mm, free portions linear, 1.5-2.Ox
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MCDADE & TRIPP: COSTA RICAN RUELLIA (ACANTHACEAE)
203
FIG. 4. Ruellia exilis. A. Habit. B. Androecium, ventral view with corolla dissected. C. Stigma and terminal
portion of the style. D. Fruit. (Drawn from the holotype.) Illustration by Amanda Labadie.
<0.5 mm, puberulent. Corolla greenish-white
at base, distal portions of tube and lobes purple, 1.3-1.6 cm long excluding lobes, unexpanded portion of tube 7-9 mm long, expanded portion 7-9 mm long, lobes
4 x 3-4 mm. Stamens included, strongly didydamous, borne near the mouth of corolla,
apex of anther of shorter pair 13-14 mm from
base of corolla, apex of anther of longer pair
15-16 mm from base of corolla, free portions
of filaments ca. 3 mm long (shorter) and
5-6 mm long (longer), fused portion of filaments 7-8 mm long, fused filament sheath
("curtain") wholly enclosing the unexpanded
portion of corolla tube, anthers 1.5-2 mm long,
thecae basally mucronate, styles 1.2-1.4 cm
long, not extending beyond corolla mouth,
stigmas bifid but dorsal lobe nearly non-
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existent. Fruits clavate, 1.0-1.4 cm long,
glandular-puberulent to glabrous, stipe 7-8 mm
long, seed bearing portion 5-7 mm long. Seeds
to 8 per capsule, orbicular, 2-3 mm in diameter, margins with hygroscopic trichomes.
Distribution and ecology. Ruellia exilis is
known only from the Osa Peninsula of
Puntarenas Province, including central (Quebrada Vaquedano) and western (Rio Agujitas)
portions of the border between Parque Nacional Corcovado and Reserva Forestal Golfo
Dulce, and the region just south of Sierpe
(Rio San Juan). Plants occur in primary
forests at 100-600 m elevation, frequently in
riparian habitats. Flowering specimens of R.
exilis have been collected from early November to late January and fruiting specimens
from early November to mid-February. This
species should be sought in other areas of
the Osa from riverbanks and forest interior
habitats.
Etymology. The specific epithet means
`thin, slender, meager' in Latin, and refers
to the narrow leaves, delicate inflorescence
branches, and diminutive size of flowers and
fruits.
Additional specimens examined. COSTA RICA.
PUNTARENAS: Osa Peninsula, Parque Nacional Corcov-
ado, Rio Tigre, Rio Agujas, Estacion Agujas. 8°31'30"N,
83°25'40"W, 200-300m, riparian, May 1997, A zofeifa
308 (INB, MO); Parque Nacional Corcovado, Jimenez,
Estacion Los Patos, 8°3359N, 83°30'59"W, loom, Jul
1998, Rodriguez 3555; Parque Nacional Corcovado,
Cerro Brujo, 8°38'N, 83°35'W, 600m, Jan 1991, Castro
261 (INB, MO); Parque Nacional Corcovado, Estacion
Cerro de Oro, Rio Termo, 8°3250"N, 83°3045"W, Apr
1996, A ngulo 592 (INB); Sierpe, San Juan, cuenca
media del Rio San Juan, 8°4350"N, 83°32'10"W, 200m,
Nov 1990, Herrera 4548 (INB, MO); Parque Nacional
Corcovado, Cerro Brujo, Orilla de la Quebrada Vaquedano, 8°38N, 83°35'W, 400m, Jan 1991, Cordero 138
(INB, MO); Parque Nacional Corcovado, Los Planes (La
Gloria), orillas del Rio Agujitas, 8°37'30"N, 83°40'50"W,
loom, bosque primario, Feb 1991, R. Gonzalez 43
(INB, MO).
Ruellia exilis resembles a number of other
species that we refer to informally as the
"long-pedunculate" group. These are plants
with diffusely branching cymose inflorescences arising from the leaf axils. Unbranched basal peduncles are longer than
distal branches and generally raise the inflorescence above the level of the foliage. As we
treat the species belonging to this informal
group, five others occur in Costa Rica:
[VOL. 59
R. pittieri (generally below 300m, Pacific
slope, southern portion of the country), R.
stemonacanthoides (almost always above
500 m, Pacific slope, northern portion of the
country, one Atlantic slope record), R.
norvegigratiosa (600-700 m, single collection from Pacific slope, central portion of
country), R. biolleyi Lindau (Atlantic slope
only) and R. malacosperma (nativity uncertain, we have seen collections only from near
Puerto Limon, Atlantic slope). It is, however,
easily distinguished from all these by its
much narrower leaves (3 cm wide at most,
leaves >6 x longer than wide), inflorescences
with slender peduncles and branches, diminutive corollas (tubular portion < 1.6 cm) and
fruits (< 1.5 cm long), and anthers with thecae basally mucronate (except R. norvegigratiosa which also has basally mucronate anthers).
Picado & Gamboa 145 (INB), from near
Sirena station, Corcovado National Park, is
typical of R. exilis except in having leaves
considerably shorter (to only 6 cm long) than
other specimens of this species. As multiple
leaf morphs are known in a number of other
species of Acanthaceae (e.g., A phelandra aurantiaca Lindl., A. scabra (Vahl) Sm.) and
are thought to represent minor genetic variants, we provisionally refer this specimen to
R. exilis, while acknowledging the need for
additional collections to understand fully the
morphological variation encompassed by this
new species.
Ruellia mira McDade & E. Tripp, sp. nov.
Type: Costa Rica. Puntarenas: Canton de
Osa, Sierpe, San Juan, cuenca superior del
Rio San Juan, 8°43'50"N, 83°3310"W,
600m, Jan 1991, Herrera 4860 (holotype:
INB; isotypes: MO, NY, PH).
(Figs. 2C, 3C, 5)
Species nova habitu arbusculo notabilis, a congeneribus bracteis grandibus densis viridi-flavis et corollis
grandibus viridi-flavis differt.
Shrub or treelet to 4m, upper stems subquadrangular, lower stems not seen, glabrous
or bearing few, scattered trichomes. Leaves
with petioles 1.5-3.8 cm long, glabrous, cystoliths dense; laminae elliptic to oblanceolate, 10-22 x 3-7.2 cm, 3.1-3.3 times longer
than wide, apices acuminate, bases acute,
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MCDADE & TRIPP: COSTA RICAN RUELLIA (ACANTHACEAE)
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Fin. 5. Ruellia mira. A . Habit. B. Androecium, ventral view with corolla dissected. C. Stigma and terminal
portion of the style. D. Fruit. (Drawn from the holotype.) Illustration by Amanda Labadie.
margins entire to repand, both surfaces
glabrous or abaxial surface with few scattered trichomes, adaxial veins raised slightly
above surface, cystoliths visible on both surfaces, more notable on adaxial surface, very
dense along veins, both laminal surfaces with
inconspicuous, patelliform glands, drying
dark red or black. Inflorescences terminal,
of 1-2 pedunculate racemes, peduncles 1.54 cm long, densely covered in short, erect,
glandular trichomes, racemes to 14 cm long.
Bracts paired at each node, elliptic to
oblanceolate, greenish-yellow, most proximal
pair usually leaf-like, 50-60 x 5-10 mm de-
creasing to ca. half this size toward distal
nodes, apices of proximal bracts acuminate,
more distal bracts oblanceolate, apically
rounded, pubescent as peduncles, one bract
per node usually sterile (i.e., only one flower
per node), paired bracteoles subtending each
flower resembling bracts in color and shape
but smaller, 22-27 x (2—)4-7 mm, with scattered often glandular trichomes. Flowers
pedicellate, pedicels to 5 mm long, pubescent
as peduncles. Calyx pale yellowish-green,
(1 1—)15-18 x 1.5-2.0 mm, fused portions
ca. 2 mm long, free portion 9-16 mm long,
pubescent as bracteoles. Corolla yellow or
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yellowish-green basally and green distally,
narrow unexpanded portion of the corolla
tube 30-34 x 2-3 mm (pressed), expanded
portion 38-40 mm x 12-13 mm (pressed) at
mouth, lobes ca. 1 cm wide, length uncertain
but very short compared to tube, orientation
uncertain, with scattered glandular and eglandular trichomes, denser on tube. Stamens
strongly didydamous, apex of anther of
shorter pair 66-69 mm from base of corolla,
apex of anther of longer pair 72-74 mm from
base of corolla, free portions of filaments
ca. 13 mm long (shorter) and 18 mm long
(longer), fused portion of filaments ca.
50 mm long, fused filament sheath ("curtain") enclosing ca. 3/a of unexpanded portion
of tube, anthers 5-6 mm long, dorsal connective with two parallel rows of glandular
trichomes, styles ca. 7 cm long, stigmas
2.5-3 mm long, bifid, the dorsal lobe reduced
to ca. '/4 the length of longer lobe. Fruits
clavate, 16-18 mm long x 6 mm wide, stipe
5-6 mm long, seed bearing portion 11-12 mm
long, puberulent, trichomes often glandular
Seeds not seen.
Distribution and ecology. Ruellia mira is
known only from Puntarenas Province, Osa
Peninsula, central portions of Reserva Forestal Golfo Dulce, areas southwest of Sierpe
and southeast of Drake. Plants occur in riparian areas around the upper basin of the Rio
San Juan and in high forests along logging
roads, from sea level to 600 m elevation.
Flowering specimens have been collected in
January and February, and fruiting specimens
in February.
Etymology. The specific epithet means
`wonderful, astonishing, extraordinary' in
Latin and reflects the distinctiveness of this
new species in terms of stature and elongate
spikes with large and remarkably colored
bracts and corollas. Mira also means `look!'
in Spanish and this is likely to be what
botanists say to each other upon encountering
this extraordinary plant in the field.
Additional specimens examined. COSTA RICA.
PUNTARENAS: Peninsula de Osa, Reserve Forestal Golfo
Dulce, Rancho Quemado, camino a Drake, 8°43'00"N,
83°3450"W, 200-350m, Feb 1991, Chavarrfa 428
(INB, MO); Puntarenas, Cant6n de Sierpe, high forest W
of Rancho Quemado on road to Drake and new logging
road, 8°4200"N, 83°36'00"W, 1-300m, Feb 1991, Maas
et al. 7872 (INB, MO).
[VOL. 59
Ruellia mira is a very distinctive species;
with spikes of large, yellowish-green flowers
and similarly colored large bracts, it resembles no other Ruellia species in Costa Rica.
Only a few other Costa Rican species of Ruellia are woody shrubs or treelets. Among
these, R. tubiflora has more congested spikes,
greenish-white bracts and white flowers. Ruellia praeclara has axillary white flowers that
are longer than those of R. mira, and also has
much longer calyces. Finally, plants of R.
jussieuoides Schltdl. & Cham. can also be
shrubs or treelets, but flowers are axillary and
purple.
Ruellia norvegigratiosa McDade & E. Tripp,
sp. nov. Type: Costa Rica. San Jose: Dota,
San Isidro, 2 km antes de San Isidro, a orillas del camino, 84°01'20"N, 9°29'00"W,
600-700 m, Mar 2005, D. Santamaria 932
(holotype: INB). (Figs. 2D, 3D, 6)
Species nova inflorescentis longi-pedunculatis, et
corollis viridi-flavis praedita, ob inflorescentiam longipedunculatum R. exili, R. pittieri, R stemonacanthoidi,
R. malacospermae simulans, sed ab omnibus earum
foliis grandibus, corolla viridi-flava differt.
Shrub to 1 m, young stem weakly quadrangular, densely pilose with minute trichomes < 0.25 mm long, erect to upwardly
ascending, older stems not seen. Leaves
with petioles to 10.5 cm long, pubescent as
young stem, laminae broadly ovate, 14-24 x
8-16 cm, ca. 1.6 times longer than wide,
apices acute to acuminate, bases rounded,
margins crenate to entire, abaxial surfaces
pubescent as stem, mostly along veins, adaxial surfaces essentially. glabrous, cystoliths
visible on both surfaces but more obvious
above, both laminal surfaces with sessile,
patelliform glands, drying pale to dark red.
Inflorescence axillary, of much branched cymose panicles, peduncles 14.5-17 cm long,
pubescent as young stems, proximal nodes
mostly bearing branches, more distal nodes
with flowers either terminally or in one of leaf
axils. Bracts subtending first major branches
elliptic, 4-5 x 0.7-1.3 cm, bracts at more distal nodes similar but smaller and appearing
bracteole-like, these narrowly lanceolate,
rounded apically, 51 mm long, pubescent as
leaves, with patelliform glands as on leaves,
bracteoles not seen. Flowers with pedicels
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FIG. 6. Ruellia norvegigratiosa. A. Habit. B. Androecium, ventral view with corolla dissected, with close-up
showing basally mucronate anthers. C. Stigma and terminal portion of the style. D. Fruit. (Drawn from the holotype.)
Illustration by Amanda Labadie.
1-3 mm long, trichomes pubescent, <0.25 mm crescent in fruit to 10-12 mm long, pubescent
long, mostly eglandular, few glandular, mixed as pedicels, often more densely pubescent
orientation. Calyx 7-10 x ca. 1 mm, fused within. Corolla pale green, outer surface
portion 1-1.5 mm long, free portions 6-9 mm sparsely pubescent with trichomes as on petilong, narrowly lanceolate, calyx slightly ac- oles, unexpanded portion of tube 2.1-2.5 cm
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long, expanded portion 1.8-2.0 cm, lobes
4-5 x 4-5 mm, rounded at apex. Stamens extending to mouth of corolla, not strongly didynamous, apex of anthers ca. 45 mm from
base of corolla, free portions of filaments
16-18 mm long, fused portions of filaments
25-28 mm long, fused filament sheath ("curtain") not seen, anthers 3-4 mm long, basally
mucronate, styles nearly 5 cm long, stigma
extending just beyond anthers at anthesis,
bifid, ventral lobe ca. 2 mm long, dorsal lobe
reduced ca. 0.5 mm. Fruits clavate, to 25 mm
long x 6 mm wide, 4 mm thick, stipe 15 mm
long, seed bearing portion 10 mm long, pubescent with minute erect eglandular and
glandular trichomes. Seeds to 8, at least 3 mm
in diam, but mature seeds and hygroscopic
trichomes not seen.
Distribution and ecology. Known only
from the type collection on the Pacific slope,
extreme southwest corner of San Jose
province. The plant was collected in flower
and fruit in early March.
Etymology. The name of this new species
is a compound adjective, combining the Latin
terms for Norway (Norwegi-) and full of favor
(-gratiosa). It gives us great pleasure to name
this attractive new species in recognition of
the government and people of Norway whose
commitment to biodiversity is manifested in
their generous support of the Instituto Nacional de Biodiversidad and the herbaria of
Central America.
Ruellia norvegigratiosa is a very distinctive species. Based on inflorescence structure, plants of this species might be confused
with others in the "long-pedunculate" group
discussed above (i.e., R. biolleyi, R. exilis, R.
malacosperma, R. pittieri, R. stemonacanthoides). However, it is easily distinguished
from all of these by its very large, broadly
ovate leaves that are >— 8 cm in width (all others
are < 8 cm wide). Also, R. norvegigratiosa
has pale green flowers whereas corollas of
the other species are purple or pink (portions
of the corolla in R. biolleyi can be pale green,
but this species is known only from the Caribbean slope and has flowers about half the
size of those of R. norvegigratiosa). Among
Costa Rican species with yellowish or greenish flowers, R. standleyi Leonard has much
smaller leaves and corollas, is densely glan-
[VOL. 59
dular pubescent, and has elliptical fruits. Ruellia mira has spicate inflorescences with
much larger bracts and flowers.
New Record for Costa Rica
Ruellia matagalpae Lindau, Bull. Herb.
Boissier 3: 364. 1895. Type. Nicaragua:
Matagalpa, Canada, Yerica, Rothschuh 395
(B, destroyed-n.v.).
Recent collections from the volcanoes of
the Guanacaste chain have included the first
known specimens of R. matagalpae from
Costa Rica. This is a wide-ranging species
that is known from southern Mexico to
Nicaragua such that its presence in northern
Costa Rica is not especially surprising.
Plants of this species can be distinguished
from other Costa Rican Ruellia as set forth in
the key. Ruellia matagalpae is probably related to R. tubiflora in that it possesses similar ovate-elliptic bracts that are leaf-like and
pale green to whitish in color. Also,
yellowish-orange glands that cover leaves,
bracts, and calyces are readily visible as in R.
tubiflora. To date, glands in Ruellia have
been poorly studied (but see Daniel, 1990,
Ezcurra, 1993). Research by the junior author suggests that foliar glands may be
nearly ubiquitous in the genus, although
there is a great deal of variation in apparency
among species. As foliar glands are unusual
among Acanthaceae, we illustrate these in
the four new species described here (Fig. 3)
and note that this character merits further
study.
Specimens examined. COSTA RICA.
GUANACASTE:
Parque Nacional Guanacaste. Estacion Cacao, 900 m, Feb
1995, V illalobos 78 (INB); Parque Nacional Guanacaste,
Cerro El Hacha, 300-600m, Sep 1991, Espinoza 151
(INB); Parque Nacional Rincon de la Vieja, Estacidn Las
Pailas, Sendero Santa Maria, 1200m, Jan 1993, D. Garcia
54 (INB).
Revised Taxonomic Concepts
Ruellia blechum L., Syst. nat. ed. 10, 2: 1120.
1759 (as "blechnu"). Blechum brownei Juss.
Ann. Mus. Natl. Hist. Nat. 9:270. 1807.
"Blechum blechum" (L.) Millsp. Publ. Field
Columbian Mus., Bot. Ser. 2: 100. 1900.
Lectotype: Illustration of Sloane (Voy. Jamaica 1: 5. 109, fig. 1. 1707).
20071
MCDADE & TRIPP: COSTA RICAN RUELLIA (ACANTHACEAE)
Blechum pyramidatum (Lam.) Urb., Repert. Spec.
Nov. Regni Veg. 15: 323. 1918. Barleria pyramidata Lam. Encycl. 1: 380. 1785. Type: Illustration
of Plumier (Pl. amer. 2: t. 42, fig. 3. 1756).
Ruellia costaricensis (Oerst.) E. Tripp &
McDade, comb. nov. Blechum costaricense Oerst., Vidensk. Meddel. Dansk
Naturhist. oren. Kjobenhavn 1854: 168.
1855. TYPE: Costa Rica: Paa Bjerget Aguacate med Blomst i November, Oersted s.n.
(holotype: C-n.v.).
Blechum dariense Lindau, Repert. Spec. Nov. Regni
Veg. 12: 423. 1913. Type: Panama: ad Sambu River
in prov. Darien meriodionali supra eastuum limites,
fl. et fruct. Feb, Pittier 5549 (Holotype: US [photo
seen]; isotype: NY [photo seen]).
The genus Blechum P.Br. has been recognized as distinct from Ruellia on the basis of
pollen characters; given the remarkable homogeneity of pollen morphology among Ruellia
species (Daniel, 1998), this indeed seems a
notable distinction. However, to our knowledge, macromorphological characters that distinguish species of the two genera are lacking.
Further, results from phylogenetic analysis of
molecular data indicate that the two Costa
Rican species of Blechum are sister species
and that they are embedded among lineages of
Ruellia (Tripp, unpubl. data). These results indicate that the pollen type of Blechum is autapomorphic, and macromorphological and
molecular data reflect the relationship of these
species to Ruellia. We prefer to transfer these
two species of Blechum (including the type
species of Blechum, i.e., Blechum pyramidatum) to Ruellia rather than to break the genus
Ruellia into numerous new genera as would be
required to recognize monophyletic taxa.
Tripp's work has not yet included other
species of Blechum and it would be premature
to combine the entire genus with Ruellia.
Ruellia biolleyi Lindau, Anales Inst. Fis.Geogr. Nac. Costa Rica 9: 188. 1898. Syntypes: Costa Rica: Confluent du R. Puerto
Viejo et du Sarapiqui Biolley 7396 (B?n.v., US [image seen]); Costa Rica: Talamanca, dans la foret a Xirores, 100 m, Tonduz 9297 (location unknown).
Ruellia cooperi Leonard, Pub. Field Mus. Nat. Hist.,
Bot. Ser. 18: 1251. 1938. Type: Panama, Bocas del
209
Toro: Daytonia Farm Region of Almirante,
Jan—Mar 1928, Cooper 185 (holotype: F [photo
seen]; ISOTYTES: US, YU)
In Leonard's (1938) treatment of the Acanthaceae of Costa Rica, he stated that R.
cooperi is "distinguishable from R. pittieri by
its pubescent capsules
and nearly
glabrous corollas." He made no comparison
of R. cooperi to R. biolleyi but his descriptions of both include "sparingly puberulent
corollas," puberulent capsules, and oblong
leaves. Further, R. cooperi has corollas
2.5 cm long, within the range of those of R.
biolleyi. The type of R. cooperi (Cooper 185)
was collected in Bocas del Toro, Panama, a
region that is within the range of R. biolleyi
but not R. pittieri. We have studied isotypes
of R. cooperi and find that it is not distinguishable from plants of R. biolleyi. Thus,
both plant specimens and descriptions indicate that R. cooperi should be considered a
synonym of R. biolleyi.
The status and locations of type material of
R. biolleyi require additional research. The
idea that a specimen of Biolley 7396 is or was
at B comes from floristic treatments of Central American plants and has not been verified
by us; further, there is no photo of such a
specimen among the collection of images of
European types at F. We have seen the sheet
of Biolley 7396 at US but the collection is
mixed: the majority of the plant material on
the sheet is of R. metallica Leonard and there
is only a portion of an inflorescence of R. biolleyi. It seems unwise to designate this sheet
as the lectotype until more authentic material
is sought. The location of the one or more
sheets of Tonduz 9297 is unknown. Unlike
some of Lindau's publications, the Anales
Inst. Fis.-Geogr. Nac. Costa Rica has no protologue that would permit an inference about
the location of the specimens cited.
...
Ruellia pittieri Lindau, Bull. Herb. Boissier 5:
655. 1897. Type: Costa Rica: in silvis Cerro
del Volcan prope Boruca, 1200m. Pittier
6743 (holotype: B?-n.v.; isotype: US).
Ruellia tonduzii Lindau, Anales Inst. Fis.-Geogr. Nac.
Costa Rica 8: 184. 1898, and in Pittier, Prim. Fl.
Costaric. 2: 302. 1900. Type: Costa Rica: Punta
Mala, zone littorale du Pacifique, Tonduz 6778
(holotype: B?-n.v.; isotype: US).
210
BRITTONIA
Ruellia pittieri and R. tonduzii are names
that have been applied to a taxonomically
challenging group of plants. We have examined numerous specimens identified as belonging to both species. Some morphological
distinctions can be made that hold in many
cases, and there is a degree of geographic
structure to the morphological variation. Ruellia pittieri is often the name applied to
specimens with longer (and sometimes
broader) leaves, longer peduncles, and wider,
oblong bracts. Plants with these characteristics have been most commonly collected at
low elevations on the mainland, especially on
the southwestern slopes of the Cordillera
Central and at the "base" of the Osa Peninsula where it connects to the mainland (e.g.,
Rio Sierpe region). Ruellia tonduzii is frequently applied to specimens with shorter
and usually narrower leaves, shorter pedunIles, and shorter bracts from the Osa Peninsula proper (e.g., Parque Nacional Corcovado).
Nonetheless, for all characters and combinations of characters that distinguish most specimens, other specimens are intermediate and
bridge the putative gaps. The geographic pattern is also far from absolute, as numerous
"pittieri-like" specimens are from the Osa
proper and plants with "tonduzii-like" traits
have been collected from the mainland (i.e.,
excluding Osa). Study of the type specimens
of both species indicates that they are not
distinguishable, and that they represent neither the morphological distinctions nor geographic patterns (i.e., both are from the mainland and are more robust than most
collections from the Osa) that often seem
to distinguish two entities. Thus, we synonymize R. tonduzii with R. pittieri, while
acknowledging that further taxonomic,
phylogenetic, or biosystematic data might reveal more than one entity worthy of species
recognition.
We have not been able to determine with
confidence the locations of the holotypes of
these two taxa. In the introduction to the
paper in which Ruellia pittieri is published,
Lindau (1897) indicates that he discovered a
number of new Acanthaceae among specimens at the Botanical Museum in Berlin.
This permits a strong inference that the holotype was at B; however, that specimen is not
among the collection of images of European
,
[VOL. 59
types at F. Lindau described R. tonduzii in
two publications, as presented above, in both
cases without a general introduction or protologue for the new species. There is also no
image of the specimen in F's collection of
images of European types. Our suggestion
that the holotype was at B is more speculative and additional type material of this
taxon should be sought. We are fortunate
that US holds isotypes of these collections.
There has also been confusion regarding the
dates of publication of the two descriptions
of R. tonduzii, apparently related to the fact
that both are in multi-fascicle volumes that
were published serially over a number of
years. The dates used above reflect the date
of publication of the fascicle containing the
treatment of Acanthaceae, to the best of our
ability to determine those dates; we also correct the page numbers which have been incorrect in some regional floristic treatments.
Finally, as the Anales Inst. Fis.-Geogr. Nac.
Costa Rica is not widely available, it merits
noting that the description of R. tonduzii
published there is identical to that in the
somewhat more widely available Prim. Fl.
Costaric.
Ruellia geminiflora H. B. & K., Nov. Gen.
Sp. (quarto ed.) 2: 240. 1817 [1818]. Type:
Colombia: von Humboldt & Bonpland s. n.
(holotype: P [photo seen: F-039431]).
Ruellia campestris (Oerst.) Hems!., Biol. Cent.-Amer.,
Bot. 2(12): 504. 1882. Gymnacanthus campestris
Oerst., Vidensk. Meddel. Dansk Naturhist. Foren.
Kjobenhavn 1854 (8-12): 126-127. 1855. Type:
Costa Rica: Oersted s.n„ no date (holotype: C).
Oersted described R. campestris from a
single specimen of a plant he collected at Hacienda Santa Rosa in the then department of
Guanacaste. Interestingly, in the same paper,
he reports a new specimen of R. geminiflora
from Mexico but without describing that
plant or indicating how it differed from his
new species. To our knowledge, R.
campestris has not been applied to other
specimens from Guanacaste or elsewhere in
the intervening 150+ years since Oersted collected in the area. Durkee (1986) did not include this species in his treatment of the
Acanthaceae of Costa Rica. Leonard (1938),
in his treatment of Acanthaceae for the Flora
2007]
211
MCDADE & TRIPP: COSTA RICAN RUELLIA (ACANTHACEAE)
of Costa Rica, mentioned R. campestris only
briefly, saying "(p)robably a small-leaved
form of R. geminiflora." He provided no description. The type specimen (C) has floral
characteristics that are consistent with R.
geminiflora. The specimen seems to have
been nearly leafless or to have lost its leaves;
still, so far as can be determined, leaves,
stems,, and enlarged, woody rootstock are
likewise consistent with those of plants of R.
geminiflora. Plants of R. geminiflora have
been collected near the type locality for R.
campestris. Thus, there appears to be no morphological or geographic basis for recognizing R. campestris as distinct from R. geminiflora.
Doubtful Species
Ruellia barbillana Cufod., Archiv. Bot. 10:
47. 1934 versus R. metallica Leonard,
Publ. Field Mus. Nat. Hist., Bot. Ser. 18:
1253. 1938.
Ruellia barbillana is known only from the
type collection (Cufodontis 657, W ) made
along the Rio Barbilla in Lim6n. It is described as having glabrous capsules and
corollas. Leonard later described R. metallica
to accommodate a very similar and widespread plant, but one with slightly pubescent
capsules and corolla. We have seen a relatively high quality image of the type specimen, kindly posted to the local server by curatorial staff at W but have not been able to
study the specimen. The plant appears to be
consistent with R. metallica although inflorescence characters are obscure and we cannot determine whether it has the distally
gradually reduced floral bracts that are distinctive for that species. Certainly, the plant
was collected from within the range of R.
metallica. However, synonymizing these two
species would be disruptive as R. metallica is
the junior name and is in widespread use.
Thus, we defer making changes until we can
study the actual type specimen.
Range Extensions
Ruellia stemonacanthoides. Plants of this
species have been collected from above
500 m on the Pacific slope of San Jose, Ala-
juela, and Guanacaste provinces. A single
collection from the Caribbean slope of this
species (Limon, Guapiles, Bosques de Toro
Amarillo, 300m, Apr 1941, J. Leon 669 [ F])
has been found, and this area should be explored further.
Ruellia golfodulcensis. As the specific epithet indicates, this species is known primarily
from the Golfo Dulce region, Pacific slope,
southwestern Costa Rica. Two collections
from the Caribbean slope of northeastern
Costa Rica are apparently of this species
(Heredia, Canton de Sarapiqui, Llanura de
San Carlos, Lomas Sardinal; ca. 15 km Linea
recta N de Puerto Viejo, 10°3410"N
84°02'50W, 250-350 m, Mar 1994, Hammel
& Garita 19478 [INB]; Alajuela, Cuenca del
San Carlos, Boca Tapada, Finca San Jorge,
100m, Feb 1996, A. Rodriguez 1068 [INB,
F]). These collections suggest that R. golfodulcensis may be more widespread than previously known. This plant should be sought
in other areas of lowland wet forest.
,
Identification Aids
Figure 7. Corolla morphology. All
species of Costa Rican Ruellia have sympetalous tubular corollas with a narrow basal
portion, expanded distal portion, and five
free lobes. As shown in Fig. 7, the corollas
of Costa Rican Ruellia species differ considerably in total length and also in the relative
dimensions of the different portions. The
two distinct regions of the tube may be similar (e.g., Fig. 7A) or markedly different in
relative lengths (e.g., Fig. 7H and 7S with
narrow tube shorter than expanded portion,
Fig. 7B and 7C with narrow tube much
longer than expanded portion; see also Fig. 3
in Ezcurra, 1993). The corolla lobes also
vary among species in terms of absolute size
and size relative to the tube length (e.g., contrast Fig. 7D and 7Q). In combination with
each other and with color, these traits offer
considerable power to distinguish species although they have not always been clearly
characterized in species descriptions. Orientation of the lobes is also potentially informative (e.g., many neotropical hummingbirdpollinated species have retrorse lobes),
although this can be difficult to assess from
herbarium specimens. Because orientation of
212
BRITTONIA
[VOL. 59
FIG. 7. Corolla variation among Costa Rican species of Ruellia. Corollas are white throughout (A—E), purple or
pinkish (F—P, T—V); yellow or greenish (Q—S). A. R. tubiflora (Hammel 8030, DUKE), with bars to right identifying
the "narrow unexpanded portion" (below) and "expanded portion" (above) of corolla tubes. B. R. odorata (Burger &
Gentry 8915, US). C. R. palustris (Grayum 1467, DUKE). D. R. praeclara (Rodriguez 1928, DUKE). E. R.
golfodulcensis (Herrera 4600, MO). F. R. jussieuoides (V alerio 83, DUKE). G. R. matagalpae (Davidse & Brant
32422, MO). H. R. malacosperma (W ilbur 25729, DUKE). I. R. nudiflora (McDade 192, DUKE). J. R. metallica
(W ilbur 39311, DUKE). K. R. geminiflora (Liesner & Lockwood 2567, MO). L. R. paniculata (Breedlove 23703,
MO). M. R. biolleyi (W ilbur 30115, DUKE). N. R. pittieri (McDade 390, DUKE). O. R. exilis (A lfaro 138, PH). P. R.
inundata (Burger et al. 11335A , US). Q. R. mira (Herrera 4860, INB). R. R. norvegigratiosa (Santamaria 932, INB).
S. R. standleyi (Tripp 147, DUKE). T. R. stemonacanthoides (Espinoza 1206, MO). U. R. costaricensis (Grayum
9310, MO). V. R. blechum (Kernan 49, MO).
the lobes varies and can be altered by pressing, we emphasize dimensions of the corolla
tube in characterizing and distinguishing
species. These dimensions are readily taken
on specimens with reasonably well-pressed
anthetic corollas.
Figure 8. Capsule Morphology. All
species of Ruellia have woody capsules
within which the seeds are borne on modified
funiculi termed retinacula; fruits are explosively dehiscent and seeds are ballistically
dispersed. In R. costaricensis and R. blechum,
placentas and retinacula fracture from capsule walls during fruit dehiscence, possibly
dispersing the entire structure (including all
seeds) as a single unit. This trait has been referred to as `elastic dehiscence of the placenta' and is known from some species of
Ruellia that do not occur in Costa Rica (e.g.,
R. erythropus (Nees) Lindau and R. hypericoides (Nees) Lindau, Ezcurra 1993; R.
beyrichiana (Nees) S. Moore and R. leucantha Brandegee, Tripp 2007), as well as other
acanths (e.g., species of Dicliptera Juss., Diclipterinae, Justicieae sensu McDade et al.
[2000]). There is considerable variation in
fruit size and shape as well as seed number
and seed position within fruits among Costa
Rican Ruellia species (Fig. 8; see also Fig. 5
in Ezcurra, 1993). Among our species of Ruellia, fruit length varies from R. geminiflora
(6.5-10 mm, Fig. 8T) to R. tubiflora
(25-30 mm. Fig. 8A). Fruit characters can
often distinguish sympatric species when
flowering material is unavailable (e.g., R.
paniculata L. and R. inundata H. B. & K.,
Fig. 8J vs. 8S). Most notably, some species
of Ruellia have clavate fruits with a narrow
sterile stipe and a wider seed-bearing portion
(Fig. 8A-8G, R. tubiflora through R. golfodulcensis, and Fig. 8L-8R, R. norvegigratiosa through R. exilis). Others have elliptical
fruits that lack a distinct stipe and have a
seed-bearing portion not substantially wider
than the stipe (Fig. 8H-8K, R. malacosperma
through R. standleyi). Two of our species, R.
inundata (Fig. 8S) and R. geminflora (Fig. 8T),
are clavate in lateral view but this is not readily visible in frontal view as drawn. Seeds
can be evenly distributed along the elongated
seed-bearing portion of the capsule (e.g., Fig.
8L-8R, R. norvegigratiosa through R. exilis)
20071
MCDADE & TRIPP: COSTA RICAN RUELLIA (ACANTHACEAE)
213
FIG. 8. Fruit variation among species of Costa Rican Ruellia. Shown are both locules, but only one-half of each.
A. R. tubiflora (W ilbur 15088, DUKE), with bars to right identifying the "stipe" or "sterile portion" (below) and
"seed-bearing portion" (above) of capsules; B. R. praeclara (Herrera 2604, DUKE). C. R. palustris (W ilbur 38646,
DUKE). D. R. matagalpae (Rees 182, MO). E. R. odorata (Quesada 414, INB). F. R. mira (Herrera 4860, MO). G.
R. golfodulcensis (Hammel 19478, INB). H. R. malacosperma (Rodriguez 193, MO). I. R. nudiflora (Gdmez 230117,
MO). J. R. paniculata (Contreras 8521, DUKE). K. R. standleyi (Tripp 147, DUKE). L. R. norvegigratiosa
(Santamaria 932, INB). M. R. jussieuoides (Tripp 156, DUKE). N. R. stemonacanthoides (Utley 3909, DUKE). O. R.
metallica (W ilbur 39311, DUKE). P. R. pittieri (McDade 390, DUKE). Q. R. biolleyi (Tripp 142, DUKE). R. R. exilis
(A lfaro 138, MO). S. R. inundata (Burger 11335A , DUKE). T. R. geminiflora (Liesner 2567, MO). U. R.
costaricensis (Aguilar 853, INB). V. R. blechum (Kernan 49, MO). Illustrations by K. Deregibus.
or, alternatively, may be clustered on a reduced, compact seed-bearing portion (e.g.,
Fig. 8A-8G, R. tubiflora through R. golfodulcensis). Number of ovules per capsule varies
among species (4-20), with minor intraspecific variation. Fruits also vary in the thick-
ness of the walls. Though difficult to measure
and not generally mentioned in species descriptions, this character seems to be entirely
consistent within species and can sometimes
be used to characterize groups of related
species within Ruellia (Ezcurra, 1993; Tripp,
214
BRITTONIA
unpubl. data). For example, R. stemonacanthoides (Fig. 8N) and R. metallica (Fig. 80)
can readily be distinguished by their thickversus thin-walled fruits, respectively. This
trait merits further study.
QUICK GUIDE TO GROUPS OF SPECIES WITH
DISTINCTIVE TRAITS
1. Plants> 1 m high (treelets): R. mira, R. norvegigratiosa, R. praclaera, R. tubiflora.
2. Plants with corollas white throughout: R. golfodulcensis (sometimes pale purple), R. odorata, R.
palustris, R. praeclara, R. tubiflora.
3. Plants with yellow, yellowish-green or greenish
corollas: R. norvegigratiosa, R. mira, R. standleyi.
4. Plants with purple or pinkish corollas, at least in
part (i.e., tube often paler than lobes),
these <3.5 cm long (excluding lobes): R. biolleyi,
R. costaricensis, R. exilis, R. geminiflora, R. gol-
[VOL. 59
fodulcensis, R. inundata, R. metallica, R. nudiflora (Engelm. & A. Gray) Urb., R. paniculata, R.
pitteri, R. blechum, R. stemonacanthoides.
5. Plants with purple or pinkish corollas, at least in
part (i.e., tube often paler than lobes),
these > 3.5 cm long (excluding lobes): R.
jussieuoides, R. malacosperma, R. matagalpae.
6. Plants with clavate fruits (Fig. 8A-8G, 8L-8T):
R. biolleyi, R. blechum, R. costaricensis, R. exilis, R. geminiflora, R. golfodulcensis, R. inundata, R. jussieuoides, R. matagalpae, R. metallica, R. mira, R. norvegigratiosa, R. odorata, R.
palustris, R. pittieri, R. praeclara, R. stemonacanthoides, R. tubiflora.
7. Plants with elliptical fruits (Fig. 8H-8K): R.
malacosperma, R. nudifora, R. paniculata, R.
standleyi.
8. Plants with fruits with placentae that fracture
from the wall at dehiscence (Fig. 8U-8V): R.
costaricensis, R. blechum.
Key for plants in reproductive condition
Inflorescences mostly terminal, densely bracteate spikes, bracts < 16 mm long, ovate; fruits with placentae
that fracture from the capsule wall at dehiscence
2. Plants lax, sprawling, rooting at nodes; leaves ca. 2x longer than wide; corolla 9-16 mm long......... R. blechum
2. Plants erect, leaves>2.5x longer than wide; corolla 20-25 mm long
R. costaricensis
Inflorescences variable, if densely bracteate then bracts>30mm long, elliptic or oblanceolate; fruits with
placentae remaining attached to capsule wall at dehiscence
3. Plants with corollas white throughout
4. Corollas (excluding lobes) ca. 3 cm long
R. golfodulcensis
4. Corollas (excluding lobes) > 5 cm long
5. Inflorescences of dense, short terminal spikes, old inflorescences often appearing as a scarred,
cone-like stump; bracts leaf-like, pale green to whitish, ovate to elliptic; calyx lobes < 1 cm long
...................................................................................................................................................... R. tubiftora
5. Inflorescences of axillary or terminal few-flowered clusters, not as above; bracts not leaf-like;
calyx lobes > 1.5 cm long
6. Woody plants to 2 m tall; calyx lobes (28-)37-55 mm long, narrow; basal portion of corolla
tube approximately equal to expanded portion; thecae > 5 mm long; fruits > 25 mm long... R. praeclara
6. Herbaceous plants mostly 51 m tall; calyx lobes <30 mm, narrow; basal portion of the corolla
tube ca. 2x longer than expanded portion; thecae <_ 5 mm long; fruits <25 mm long
7. Calyx (in flower) 10-18 x 1-2 mm, villous; bracts 10-23 x 2-5 mm; fruits to 17 mm long;
forest understory, Osa Peninsula ......................................................................................... R. odorata
7. Calyx (in flower) 25-30x4-5 mm, puberulous; bracts 22-40 x 0.6-2.3 mm; fruits 20-23 mm
long; swamps and water-logged soils, northeast Costa Rica ............................................. R. palustris
3. Plants with corollas not white throughout
8. Corollas yellow, yellowish-green, or green
9. Plants herbaceous; leaves to 12 cm long; flowers with tube < 20 mm long; fruits elliptical, central
highlands above 1000 m; Cartago province ................................................................................ R. standleyi
9. Plants woody shrubs or treelets; at least some mature cauline leaves >20 cm long; flowers with
tube> 35 mm long; fruits clavate, below 1000 m; Puntarenas and San Jose Provinces
10. Inflorescences of elongate spikes, with imbricate bracts 30-60mm long; calyx (11-)
15-18 mm long; corolla (excluding lobes) ca. 5.5 cm long, yellowish green; fruits ca. 16 mm
long; Osa Peninsula, Puntarenas province .................................................................................. R. mira
10. Inflorescences of long pedunculate, spreading dichasia, bracts much shorter and less dense
than above; calyx mostly < 10 mm long; corolla (excluding lobes) <4 cm long, green; fruits ca.
25 mm long; San Jose province ................................................................................. R. norvegigratiosa
8. Corollas wholly or in part (i.e., with at least the lobes) purple or pinkish (tube may be white)
11. Inflorescence of terminal or axillary, essentially sessile flowers, sometimes appearing as an elongate, terminal spike due to short internodes and distally progressively reduced leaves
..........................................
...............................................................................
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MCDADE & TRIPP: COSTA RICAN RUELLIA (ACANTHACEAE)
215
12. Plants<_0.5m; mature cauline leaves<_8cm long; fruits<_llmm long, clavate but with a very
shortstipe
R. geminifiora
12. Plants > 0.6 m; mature cauline leaves > 9 cm long; fruits > 11 mm long, clavate with a marked
stipe or elliptical in shape
13. Calyx lobes (in flower) < 11 mm long
14. Corolla (excluding lobes) <_ 23 mm long; inflorescences many flowered, flowers
borne in the axils of distal, progressively reduced leaves so that inflorescence appears as an elongate, terminal spike; plants of Atlantic slope ............................. R. metallica
14. Corolla (excluding lobes)? 24 mm long; inflorescence of reduced cymes, I to fewflowered in axils of upper leaves; plants thus far known only from the Cordillera de
Guanacaste, Pacific slope ................................................................................. R. matagalpae
13. Calyx lobes (in flower) >11 mm long
15. Corolla to 30 mm long; bracts >5 mm long, inflorescence of terminal clusters of flowers, if axillary, then flowers in axils of distalmost reduced leaves .................. R. golfodulcensis
15. Corolla >35 mm long (usually >40 mm long); bracts <5 mm long; flowers always
axillary............................................................................................................. R. jussieuoides
11. Inflorescences distinctly pedunculate and usually freely branching, branches terminating in 1 to
many flowered cymes
16. Calyx> 7 mm long
17. Plants <_ 0.5 m tall; leaves frequently oblanceolate or obovate .................................... R. nudiflora
17. Plants >0.6 m tall; leaves elliptic to ovate to lanceolate (not as above)
18. Corollas fuchsia or pinkish colored; capsules <_ 10mm long, with maroon-colored
splotches................................................................................................................ R. inundata
18. Corollas purple at least in part (i.e., corolla tubes may be white); capsules>_ 11 mm
long, without maroon splotches
19. Capsules > 24 mm long; leaves mostly narrowly elliptic ...................... R. malacosperma
19. Capsules <23 mm long; leaves elliptic to ovate
20. Capsules elliptical; plants often densely glandular-pubescent and odorous;
plants of low elevations, xeric habitats, Pacific slope .......................... R. paniculata
20. Capsules clavate; plants not densely glandular-pubescent and not odorous;
plants of mid-elevation, mesic habitats, Pacific slope ............ R. stemonacanthoides
16. Calyx <7 mm long
21. Corolla (excluding lobes) <_ 16 mm long, narrow, unexpanded portion of tube <_ 9 mm
long; leaves narrowly elliptic, plants <_ 0.5 m tall .............................................................. R. exilis
21. Corolla (excluding lobes) >_ 17 mm long, narrow, unexpanded portion of tube ? 10mm
long; leaves elliptic to ovate to oblanceolate, plants >0.6 m tall
22. Corolla (excluding lobes) > 25 mm long, narrow, unexpanded portion of
tube > 13 mm long; calyx mostly >_ 6 mm long ..................................... R. stemonacanthoides
22. Corolla (excluding lobes) <25 mm long, narrow, unexpanded portion of
tube < 13 mm long; calyx mostly <_ 5 mm long
23. Plants of Atlantic slope; capsules > 18 mm long; leaves oblanceolate to elliptic,
often bi-colored; narrow unexpanded portion of corolla tube>_ 10mm long;
calyx mostly >4mm long ................................................................................. R. biolleyi
23. Plants of Pacific slope; capsules < 18 mm long, leaves ovate to elliptic, never bicolored; narrow unexpanded portion of corolla tube 5 10mm long; calyx
mostly<_ 4 mm long ............................................................................................ R. pirtieri
...........................................................................................................................
Acknowledgments
This research was supported in part by a
grant from the National Science Foundation
(DEB 0108589 to LAM) and by a Grant-InAid from the Biology Department of Duke
University and a McHenry Fellowship from
the Academy of Natural Sciences in
Philadelphia (EAT). We are grateful to
Amanda Labadie for the new species illustrations (Figs. 1, 4-6) and to K. Deregibus
for the fruit illustrations (Fig. 8). M. Grayum
helped us by locating rare literature in the li-
brary at MO, T. F. Daniel helped us to access
information regarding specimens at CAS and
on loan there, and curators at a number of
herbaria permitted study of specimens or facilitated loans (ARIZ, CR, F, INB, MO, NY,
US, W). F. Barrie and D. Nicolson provided
advice on nomenclature, and P. Eckel corrected the Latin diagnoses. D. Hearn, L.
Kelly, and two anonymous reviewers provided useful comments. The fieldwork and
active collecting of botanists based at INBio
have yielded the discoveries that we report
BRITTONIA
216
here and substantially improved our knowledge of Costa Rican Acanthaceae. Finally,
we are deeply indebted to the government
and citizens of Costa Rica for their support
of conservation and biodiversity research.
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