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Full text of "Flora costaricensis"

tfuy. 



AC, 

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Botany 



FLORA COSTARICENSIS 



William 



Family #193 Scrophulariaceae 
Family #193a Schlegeliaceae 
Family #194 Bignoniaceae 
Family #195 Pedaliaceae 

^^^Hj 

Family #196 Martyniaceae 
Family #197 Orobanchaceae 



PUBLI 



FIELDIANA 



Botany 

NEW SERIES, NO. 41 



FLORA COSTARICENSIS 



William Burger, Editor 

Curator Emeritus, Department of Botany 
Field Museum of Natural History 
1400 South Lake Shore Drive 
Chicago, Illinois 60605-2496, U.S.A. 



Family #193 Scrophulariaceae Family #195 Pedaliaceae 

Kerry Barringer and William Burger William Burger 



Family #193a Schlegeliaceae 

William Burger and Kerry Barringer 

Family #194 Bignoniaceae 

William Burger and Alwyn Gentryt 



Accepted October 29, 1998 
Published April 28, 2000 
Publication 1508 



Family #196 Martyniaceae 

William Burger 

Family #197 Orobanchaceae 

Luis D. Gomez and William Burger 



PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY 



Introduction 



This is the eleventh issue in the Flora Costar- 
icensis series. The first dealt with the Piperaceae, 
family number 41 (Fieldiana, Bot. 35, 1971). The 
second included families numbered 42 through 
53, Chloranthaceae through Urticaceae (Fieldiana, 
Bot. 40, 1977). The third issue covered the Gra- 
mineae (Poaceae) and was authored by Richard 
Pohl (Fieldiana, Bot. n.s. No. 4, 1980). The fourth 
issue included families numbered 54 through 70, 
Podostemaceae through Caryophyllaceae (Fieldi- 
ana, Bot. n.s. 13, 1983). The fifth issue covered 
families 200 and 201, the Acanthaceae, authored 
by L. H. Durkee, and the Plantaginaceae (Fieldi- 
ana, Bot. n.s. No. 18, 1986). The sixth issue in- 
cluded families 80 and 81, the Lauraceae and the 
Hernandiaceae (Fieldiana, Bot. n.s. No. 23, 1990). 
The seventh issue included families numbered 97 
through 103, Krameriaceae through Zygophylla- 
ceae (Fieldiana, Bot. n.s. No. 28, 1991). The 
eighth issue included family 202, the Rubiaceae 
(Fieldiana, Bot. n.s. No. 33, 1993). The ninth is- 
sue included family 1 13, the Euphorbiaceae (Fiel- 
diana, Bot. n.s. No. 36, 1995). The tenth issue 
covered Tribe Maxillarieae of family 39, the Or- 
chidaceae (Fieldiana, Bot. n.s. No. 40, 1999). 

Illustrations of leafy or flowering stems are all 
drawn to the same scale with the exception of 
Figures 1, 2, and 28. Enlarged flowers, fruits, or 
seeds are drawn to the same scale on an individual 
plate unless otherwise noted. Bignoniaceous fruits 
are drawn to the same scale as the leafy stems. 
The closed scales represent centimeters and the 



open scales represent millimeters. The figures are 
somewhat diagramatic and represent a common or 
characteristic morphology for each species. Figure 
28 was done by Luis D. G6mez; the others are by 
William Burger. 

The circumscription of the Scrophulariaceae is 
difficult because the family is part of a closely 
related group of families that includes Oroban- 
chaceae, Bignoniaceae, Myoporaceae, Gesneri- 
aceae, Pedaliaceae, Martyniaceae, and Acantha- 
ceae. A few genera have been difficult to place. 
Gibsoniothamnus was described by Williams 
(1970) as a genus of Scrophulariaceae, and he 
noted its similarity to Schlegelia. These genera 
were placed in the Bignoniaceae by Gentry but 
are regarded as a separate family here. Reveal has 
recently (1995) opted for a finer division of the 
Scrophulariales and proposed segregating families 
from within Scrophulariaceae. We adhere to a 
more traditional circumscription of the family be- 
cause of its long history in providing a useful in- 
formation retrieval system. 

The treatment of the Bignoniaceae presented 
here is publication No. 2 in the Gentry Invitation 
Series. This series acknowledges Dr. Gentry's 
many contributions to our knowledge and under- 
standing of the Bignoniaceae. The late Alwyn 
Gentry's publications and his many authoritative 
determinations have served as the foundation for 
the treatment of the Bignoniaceae presented here. 
In addition, there are virtually no departures from 
his taxonomic concepts in this treatment, and he 
is listed as co-author for that reason. 



Acknowledgments 



We thank the staff of the Muse"o Nacional de 
Costa Rica for their assistance in collecting pro- 
grams over many years. The National Science 
Foundation and the National Geographic Society 
helped support many of these collecting activities. 
NSF grant DEB-8103184 helped support the work 
of Kerry Barringer while studying Costa Rica's 
Scrophulariaceae. We thank Warren Douglas Ste- 
vens for making available copies of treatments of 
Bignoniaceae and Scrophulariaceae prepared for 



the Flora of Nicaragua. We thank Michael Grayum 
for providing a copy of the treatment of the Big- 
noniaceae for the Manual Flora of Costa Rica. 
Noel Holmgren, William D'Arcy, and two anony- 
mous reviewers provided many helpful comments 
and corrections. The collections of the Field Mu- 
seum, the Missouri Botanical Garden, Duke Uni- 
versity, and the U.S. National Herbarium were con- 
sulted in preparing this treatment, and we thank 
those institutions for the use of their materials. 



FLORA COSTARICENSIS 
Family #193 Scrophulariaceae 
Family #193a Schlegeliaceae 
Family #194 Bignoniaceae 
Family #195 Pedaliaceae 
Family #196 Martyniaceae 
Family #197 Orobanchaceae 



SCROPHULARIACEAE 

By Kerry Barringer and William Burger 

REFERENCES W. D'Arcy, Scrophulariaceae, in 
R. Woodson et al., Flora of Panama. Ann. Mis- 
souri Bot. Card. 66: 173-272. 1979. F. Pennell, 
The Scrophulariaceae of eastern temperate North 
America. Acad. Nat. Sci. Philadelphia Monogr. 1 : 
1-650. 1935. P. Standley & L. O. Williams, Scro- 
phulariaceae, in Flora of Guatemala. Fieldiana, 
Bot. 24, 9(4): 319-416. 1973. D. Sutton, A Re- 
vision of the Tribe Antirrhineae, British Museum 
(Natural History) & Oxford Univ. Press, London, 
576 pp. 1989. D. Sutton & R. Hampshire, Scro- 
phulariaceae, in Flora of Nicaragua (unpublished 
manuscript). 1995. J. Thieret, The tribes and gen- 
era of Central American Scrophulariaceae. Ceiba 
4: 164-184. 1954. J. Thieret, Supraspecific clas- 
sification of Scrophulariaceae: A review. Sida 3: 
87-106. 1967. 

Herbs or subshrubs (rarely woody shrubs or 
trees), erect to decumbent or prostrate (rarely 
climbing), terrestrial to semiaquatic or aquatic, 
autotrophic or hemiparasitic, bisexual, stems 
without internal phloem, glabrous or with simple 
or branched, glandular or eglandular uni- or mul- 
ticellular hairs; stipules absent. Leaves alternate, 
opposite, or verticillate, simple (rarely deeply 
pinnately lobed), distal leaves often reduced and 
intergrading with the floral bracts, leaf blades 
serrate to entire or deeply pinnately lobed, ve- 
nation pinnate or less often palmate. Inflores- 
cences racemes, spikes, thyrses, or panicles of 
cymes or of solitary (several) flowers in leaf ax- 
ils, bracteoles present or absent on the pedicels, 
pedicels usually well developed. Flowers bisex- 
ual, small to large, often showy, calyx 4- or 5- 
lobed, or deeply divided to the base and sepals 
4 or 5 (2), imbricate or valvate in bud, persisting 



and often enlarging slightly in fruit; corolla unit- 
ed and tubular to campanulate, bilaterally sym- 
metric and usually 2-lipped (rarely radially sym- 
metric), lobes 4-5 (8), upper (adaxial) lip 2- 
lobed to emarginate, sometimes galeate, lower 
(abaxial) lip usually 3-lobed, tube saccate or 
spurred in some; stamens 2-4 (5), alternating 
with the lobes, usually of 2 unequal pairs, fila- 
ments borne on the tube, free, anthers with 2(1) 
equal or unequal thecae, distinct or confluent, 
rarely awned, a staminode present or absent 
(rarely 2 or 3), a disc present or absent around 
the ovary; pistil solitary, ovary superior (or half 
inferior), 2-locular (rarely 1-locular near the 
apex or 3-locular), ovules usually many on 2 ax- 
ile placentas, unitegmic, style 1, terminal, stig- 
mas simple or 2-lobed. Fruits usually dry cap- 
sules (rarely baccate), dehiscence loculicidal or 
septicidal or both (rarely indehiscent), placenta 
often persisting; seeds usually many and small, 
usually with an ornamented testa or exotesta 
(sometimes smooth or winged), endosperm usu- 
ally present, embryo small. 

The family Scrophulariaceae includes 285 gen- 
era with between 4,000 and 5,000 species. The 
family is primarily north temperate, but with trop- 
ical montane species and cosmopolitan weeds. 
One of the largest genera. Calceolaria (300-400 
spp.), is almost exclusively South American and 
montane. In this treatment we include 36 genera 
with 72 species, 1 8 of these being introduced or 
cultivated (see listing at end of family, p. 69). 
Costa Rica lies between two major areas of di- 
versity for the family: one in Andean South 
America and one in Mexico and Guatemala. 
Among the southern elements are Alonsoa, Cal- 
ceolaria, Scoparia, and Siemodia, while northern 
elements are Castilleja, Hemichaena, Lamouroux- 
ia, and Russelia. 

Most Scrophulariaceae are recognized by their 
sympetalous tubular (usually two-lipped) corollas, 



FIELDIANA: BOTANY, N.S., NO. 41, APRIL 28, 2000, PP. 1-174 



two-locular ovaries with many ovules on axile 
placentas, capsular fruits with many small seeds, 
and stems lacking internal phloem. Most species 
are herbs or small subshrubs, and the corollas are 
often showy. There is great diversity in the form 
of corolla and pollination vectors (see C. M. 
Kampny, Pollination and flower diversity in Scro- 
phulariaceae, Bot. Rev. 61: 35-366, 1995). Iridoid 
compounds are common in the family. The family 
includes many species cultivated as ornamentals 
and an important drug plant (Digitalis) but no im- 
portant food, spice, or fiber plants. Interestingly, 



a few of the most colorful genera (e.g., Castilleja 
and Lamourouxia) have not been utilized in or- 
namental horticulture because of difficulty in 
propagating their hemiparasitic species. Species 
likely to be mistakenly identified as Scrophulari- 
aceae are found in Lamiaceae, Acanthaceae, and 
Gesneriaceae, as well as the other families includ- 
ed in this volume. Current molecular studies in- 
dicate that the Scrophulariaceae, as traditionally 
defined, are polyphyletic (Wagstaff & Olmstead, 
1997). We retain the traditional circumscriptions 
of families in this series to facilitate information 
retrieval. 



Key to Genera and Unusual Species of Scrophulariaceae 

la. Plants fully aquatic, only the inflorescences held above the water level; leaves submerged or 

floating, whorled, with pinnatisect filiform divisions [rarely collected] Benjaminia reflexa 

Ib. Plants aquatic, partly aquatic, or terrestrial, if aquatic the leaves floating or emergent and not with 

submerged filiform pinnatisect divisions 2 

2a. Plants slender-stemmed twining or climbing, usually found in gardens and near habitations; leaves 

mostly alternate 3 

2b. Plants erect or creeping on the ground or aquatic, not twining or climbing, found in a great many 

habitats; leaves alternate or opposite 5 

3a. Leaf blades reniform to orbicular, cordate at base, 10-35 mm long, alternate or rarely op- 
posite; corollas 7-9 mm long, with a basal spur Cymbalaria muralis 

3a. Leaf blades often triangular-subcordate, to 60 mm long, always alternate; corollas 30-80 mm 

long, without a spur 4 

4a. Corollas 60-70 mm long; calyx 15-24 mm long; seeds with lateral wings around much of 

the seed Lophospermum erubescens 

4b. Corollas 30-40 mm long; calyx 9-15 mm long; seeds without thin wings .... Maurandya 
5a. (from 2b) Leaves deeply dissected or compound, the lobes or leaflets > 30% the width of the 

blade 6 

5b. Leaves entire to crenate, dentate or lobed, the lobes or teeth < 30% the width of the blade . . 8 
6a. Fruits linear; plants small (40 cm), erect, in open weedy lowland (0-600 m) sites; leaves 

opposite, to 20 mm long Schistophragma mexicana 

6b. Fruits broader; plants usually taller, in montane (1 100-3800 m) habitats; leaves opposite or 

alternate, 10-100 mm long 7 

7a. Leaves opposite; corolla bright yellow with the lower lip sac-like; calyx 4-lobed, green . . . 

Calceolaria 

7b. Leaves alternate; corolla and bracts red, orange, yellow, or green, the lower lip usually 
narrowed and with acute lobes; calyx 4-parted but usually united to form 2 large lateral lobes, 

often colored Castilleja 

8a. (from 5b) Lower leaves alternate, plants usually with leaves alternate all along the stem 9 

8b. Lower leaves opposite or whorled, distal leaves opposite or alternate along the stem or the leaves 

rosulate from the base 13 

9a. Corolla expanded at the base and spurred [yellow or blue, 5-30 mm long; lower leaves linear 

to lanceolate; cultivated for ornament] Linaria 

9b. Corolla tubular, not expanded at the base to become saccate or spurred; wild, naturalized, or 

cultivated species 10 

lOa. Plants repent with creeping stems rooting at most nodes; leaf blades rounded-orbicular [to 
25 mm wide]; corolla 4-8-lobed, rotate, 3-5 mm wide Sibthorpia repens 



FIELDIANA: BOTANY 



lOb. Plants erect, not rooting at distal nodes; leaf blades not rounded-suborbicular; corolla not 

rotate 11 

11 a. Corolla with the upper lip galeate, elongate, narrowly 1-lobed and entire, lower lip shorter 
than the upper, small and often recurved; distal bracts often becoming colorful [not found 

below 700 m elevation] Castilleja 

lib. Corolla not as above, lobes rounded with the lower lobes usually longer than the upper; 

bracts not becoming colorful 12 

12a. Corolla 7-10 mm long, white, campanulate; stamens 5; native plants at 0-500 m elevation. . . . 

Capraria biflora 

12b. Corolla 20-50 mm long, usually pink marked with purple dots within, tubular; stamens 4; 
in gardens or naturalized at 1800-3300 m elevation (if at lower elevations and with viscous 

hairs or slime glands, see Martyniaceae and Pedaliaceae) Digitalis purpurea 

13a. (from 8b) Plants lacking erect stems, < 7 cm tall; found only in wet sites above 3300 m elevation 
in Central America; leaves linear to oblanceolate, in dense fascicles or rosulate at the nodes; corolla 

2-3 mm long, rotate, with 3-5 lobes Limosella acaulis 

13b. Plants with erect stems or without the above combination of characteristics 14 

14a. Leaf blades linear to linear-lanceolate, often scabrous, larger blades not exceeding 7 mm in width 

[plants with stiff erect stems but rarely exceeding 0.7 m height; corollas 8-15 mm long] ... 15 

14b. Leaf blades narrowly lanceolate to broadly ovate or rounded, scabrous or smooth; larger blades 

usually > 7 mm in width 18 

15a. Corolla 20-45 mm long or with spreading lobes to 2 cm wide, magenta, red, blue-violet, or 
purple to yellowish, glabrous or densely puberulent externally; flowers pedicellate in leaf 
axils or in distal racemes; plants not wiry or scabrous herbs or subshrubs, 0.4-1.5 m tall . . 

... 16 

15b. Corolla 7-15 mm long, lavender to pale purple, glabrous externally; flowers sessile or ped- 
icellate; plants wiry and scabrous, herbs to 0.6 m tall 17 

16a. Corolla densely minutely puberulent externally, narrowly tubular with small distal lobes 

Lamourouxia spp. 

16b. Corolla glabrous externally, with short tube and broadly spreading large lobes 

Angelonia angustifolia 

17a. Flowers borne on pedicels 10-35 mm long; fruits 4-7 mm diam.; erect stems usually with 

several prominent lateral branches Anisantherina hispidula 

17b. Flowers sessile or subsessile on pedicels < 3 mm long; fruits 2-3 mm diam.; erect stems 

usually with few or no prominent lateral branches Buchnera weberbaueri 

18a. (from 14b) Corollas with the lower lip forming a distal sac and slipper-like, usually bright yellow 

[10-25 mm long; plants both wild and cultivated for ornament] Calceolaria 

18b. Corollas not slipper-like, the lower lip not forming a distal sac, variously colored, usually with a 

short or long tube and spreading distal lobes 19 

19a. Corollas becoming > 15 mm long at anthesis 20 

19b. Corollas usually < 15 mm long at anthesis 29 

20a. Corollas 70-120 mm long, white and salverform (with a narrow tube and broad subequal 

rotate lobes) Escobedia grandiflora 

20b. Corollas 14-45 mm long, variously colored, usually tubular and somewhat 2-lipped with the 

lobes unequal 21 

2 la. Corolla yellow or yellow and white [flowers in axillary groupings; leaves sessile and auric- 

ulate at the base, blades narrowly lanceolate; native wild plants] 22 

21b. Corolla pink to red or purple, marked with dark spots if white or yellowish (note that Mar- 
tynia and Sesamum may key out in this dichotomy; see Martyniaceae and Pedaliaceae, Fig. 

27) 24 

22a. Plants weak-stemmed herbs to 0.4 m tall; leaves petiolate with blades rounded or trun- 
cate at the base; flowers solitary in leaf axils [corolla 14-20 mm long] 

Mimulus glabratus 

22b. Plants erect subshrubs, often to 1 (2) m tall; leaves sessile with blades lanceolate and 
gradually narrowed to the base; flowers usually several in leaf axils 23 

BURGER: FLORA COSTARICENSIS 3 



23a. Corolla 25-45 mm long, calyx 11-16 mm; fruits dry, brownish, oblong, 13-17 mm 

long Hemichaena fruitcosa 

23b. Corolla 14-23 mm long, calyx 5-9 mm long; fruits fleshy, white, globose, 7-10 mm 

long Leucocarpus perfoliatus 

24a. Corolla densely minutely puberulent or glandular puberulent on the exterior 25 

24b. Corolla glabrous on the exterior 26 

25a. Corollas densely minutely puberulent externally, the throat not closed by a palate; native 

wild species Lamourouxia 

25b. Corollas sparsely puberulent, throat usually closed by a palate; cultivated ornamentals 

Antirrhinum 

26a. Corolla tube ca. 2-4 mm diam., with small distal lobes, deep red; plants shrub-like orna- 
mentals, often without leaves and with many slender green stems Russelia equisetiformis 
26b. Corolla tube 7-10 mm diam., with prominent distal lobes, purple to rose- white or marked 

with dark coloring; plants not shrub-like, rarely without leaves 27 

27a. Plants of wet forests; stems usually unbranched and < 30 cm tall; leaf blades oblanceolate, 

to 3 1 cm long Tetranema 

27b. Plants of gardens and ornamental plantings; stems usually with a few branches, 0.4-1 m tall; 

leaf blades not oblanceolate, to 10 cm long 28 

28a. Plants becoming ca. 1 m tall; calyx united for only a short distance, lobes > half the length 

of the calyx; staminode prominent; grown as ornamentals in Costa Rica 

Penstemon gentianoides 

28b. Plants to 0.4 m tall; calyx united for half its length or more, lobes < half the length of the 

calyx; wild and ornamental species in Costa Rica Torenia 

29a. (from 20b) Calyx united for > half its length at anthesis, a calyx tube clearly present (note that 
flowers may appear to have separate sepals after anthesis as the fruit develops and the calyx tube 

splits) 30 

29b. Calyx with the sepals united only at the base, a calyx tube not evident 35 

30a. Stems and leaves scabrid or hispid, stems often becoming 1 m tall; flowers sessile or sub- 
sessile in distal spike-like inflorescences 31. 

30b. Stems and leaves neither scabrid nor hispid, stems rarely > 40 cm tall; flowers not sessile 

in spike-like terminal inflorescences (except in Bacopa sessiliflora) 32 

3 la. Corolla yellow, campanulate; flowers subtended by large bracts differing little from the 

leaves; distal leaves narrowly triangular Alectra aspera 

31b. Corolla white or marked with purple or blue, salverform; flowers subtended by bracts 

much smaller than the leaves; distal leaves narrowly lanceolate . . . Buchnera pusilla 

32a. Lower leaves forming a rosette, blades narrowly obovate with long-attenuate base; flowers 

in lax terminal racemes; rarely encountered introduced weeds Mazus pumila 

32b. Lower leaves not forming dense rosettes, blades narrowly obovate to suborbicular; flowers 

mostly axillary to distal leaves 33 

33a. Calyx 4-lobed; stems terete; leaf blades varying from obovate to suborbicular (rarely ovate 

with truncated base) Bacopa egensis 

33b. Calyx 5-lobed; stems 4-angled or with 2-4 longitudinal ridges; leaf blades mostly ovate- 
triangular or ovate-ellipsoid with truncated base 34 

34a. Calyx tube < 4 mm long, usually campanulate; fruits 3-4 mm long, rounded ovoid 

Lindernia 

34b. Calyx tube usually > 7 mm long, tubular; fruits 8-10 mm long, narrowly ellipsoid-oblong 

Torenia thouarsii 

35a. (From 29b) Fruits usually lenticular in cross-section, truncated or rounded and obcordate at the 
apex; stamens 2; corolla tube very short, with corolla lobes usually in a single plane (rotate); 

introduced weeds, 1000-3300 m elevation Veronica 

35b. Fruits usually round in cross-section, never truncated or obcordate at the apex, and without the 

other combination of characteristics 36 

36a. Sepals very unequal, the outer usually much broader than the inner and imbricate in bud (note 
that small linear bracteoles may be present at the base of the calyx) 37 

4 FIELDIANA: BOTANY 



36b. Sepals equal or subequal, valvate or less often imbricate in bud 38 

37a. Flowers yellow; bracteoles present at the base of the pedicel; leaf blades broad and abruptly 

narrowed to a short petiole Mecardonia procumbens 

37b. Flowers purple or blue to white (sometimes yellowish in age); bracteoles present in the middle 
or apex of the pedicel or absent; leaf blades usually gradually narrowed to the base or sessile 

if rounded at the base Bacopa 

38a. Corollas bright red or orange; plants erect herbs or subshrubs to 2 m tall, stems often stiff and 

longitudinally ridged 39 

38b. Corollas white to blue, purple, or yellowish; plants mostly weak-stemmed herbs 41 

39a. Leaf blades 15-31 cm long; corollas 26-55 mm long; peduncles 9-24 cm long, axillary, 

slender and flexuous; rarely collected endemic species Tetranema 

39b. Leaf blades 1-11 cm long; corollas 5-16 mm long; peduncles less than 2 cm long or the 

flowers on terminal erect racemes; common and widespread species 40 

40a. Flowers in axillary fascicles of 2-20, corolla deep red, tube 9-12 mm long with small acute 

lobes; seeds surrounded by hairs within the fruit Russelia sarmentosa 

40b. Flowers solitary in bract or leaf axils on a terminal raceme, corolla red to orange, tube < 2 
mm long with large rounded spreading-rotate lobes; seeds not associated with hair-like struc- 
tures Alonsoa meridionalis 

4 la. Corolla < 4 mm long, rotate or campanulate 42 

41b. Corolla 4-14 mm long, usually tubular and bilabiate 43 

42a. Plants terrestrial, erect, to 1 m tall; leaf blades to 35 mm long, oblanceolate and dentate; 

fertile stamens 4 Scoparia 

42b. Plants floating or prostrate, small; leaves to 8 mm long, rounded and entire; fertile stamens 

2 Micranthemum umbrosum 

43a. Fertile stamens 2; leaves sessile; stems glabrous Lindernia dubia 

43b. Fertile stamens 4; leaves petiolate (sessile and amplexicaul in Stetnodia durantifolia); stems gla- 
brous or puberulent 44 

44a. Anthers glabrous; stems and leaves puberulent; sepals acute at the apex Stemodia 

44b. Anthers with hairs; stems and leaves glabrous; sepals bluntly acute and often thickened at the apex 
Darcya 



Agalinis Rafinesque 
Nomen conservandum 

Herbs or shrubs, annual or perennial, usually 
erect and branched, glabrous or pubescent, hem- 
iparasitic on roots, turning dark when dried. 
Leaves opposite or alternate, becoming smaller or 
bract-like on distal stems, subsessile or sessile, 
blades usually narrow, entire or lobed, glabrous 
or puberulent. Inflorescences of solitary flowers 
in axils of reduced distal leaves (often resembling 
open racemes, spikes, or panicles), peduncles 
slender, subtended by small bracts, pedicels slen- 
der, bibracteolate or ebracteolate. Flowers showy, 
calyx tubular to campanulate or hemispheric, with 
5 prominent lobes or teeth, slightly imbricate or 
open in bud; corolla campanulate and often some- 
what bilabiate, glabrous or puberulent, pink to 
purple (yellow or white), tube straight or curved, 
expanded distally into a broad throat, lobes 5, 
rounded, subequal or the posterior (upper) small- 
er; stamens 4, of 2 unequal pairs, borne from the 



middle of the corolla tube, shorter than the corol- 
la, filaments usually pilose, anthers glabrous or 
puberulent, 2-thecous with parallel thecae, un- 
equal in some species; ovary glabrous, 2-locular, 
style straight and slender, deciduous, stigma soli- 
tary and linear. Fruits rounded capsules, woody 
to chartaceous or leathery, dehiscence loculicidal 
(and sometimes septicidal); seeds many, oblong to 
angular, testa reticulate. 

A distinctive New World genus with ca. 40, 
mostly North American, species. Species of this 
genus were formerly placed in Gerardia, but the 
Linnaean type proved to be a species of Acantha- 
ceae. This genus has not been collected in Costa 
Rica, but ranges southward as far as northern Nic- 
aragua. The following key includes Anisantherina 
hispidula, which occurs in Costa Rica (q.v.), and 
two species of Agalinis known from Nicaragua: 
Agalinis albida Britton & Pennell (? = Agalinis 
harperi Pennell) and Agalinis peduncularis 
(Benth.) Pennell. All three species have linear 
leaves. 



BURGER: FLORA COSTARICENSIS 



Bacopa 
monnieri 



Micranthemum umbrosum 



.imosella aquatica 
jar. americana 




FIG. 1. Scrophulariaccac: aquatic and semiaquatic species. 



FIELDIANA: BOTANY 



Stemodia 
angulata 



Stemodi a 
verticillata r 



veronica 
serpyl li folia 



Mecardonia 
procumbens " 




Lindernia diffusa' 

FIG. 2. Scrophulariaccac: small herbs with opposite leaves and broad blades. 



V. polita 



BURGER: FLORA COSTARICENSIS 



// Darcya 
costaricensis 



Stemodi a 
peduncularis 



Calceolaria 
irazuensis 




FIG. 3. Scrophulariaceae: herbs with opposite leaves and lanceolate to ovate or dissected blades. 



FIELDIANA: BOTANY 



Lindernia 
dubi a 



Schistophragma mexicana 



Anisanthenna 
hispidula 



Stemodi a 
duranti folia 




Scoparia dulcis 



FIG. 4. Scrophulariaceae: herbs with opposite leaves and narrow leaf blades or narrow pinnatifid lobes. 



BURGER: FLORA COSTARICENSIS 



Alectra aspera 



Russell a 
sarmentosa 



Lamourouxia 
guttierrezii 




L. lanceolata 

FIG. 5. Scrophulariaccac: herbs with stiff opposite leaves and erect or clambering stems. 



10 



FIELDIANA: BOTANY 



Calceolaria 
perfol i ata 



Leucocarpus 
perfol iatus 



Escobedia 
grandi flora 



Stemodia 

duranti folia 
v 




FIG. 6. Scrophulariaccac: erect herbs with larger sessile or pcrfoliatc leaves. 



BURGER: FLORA COSTARICENSIS 



11 



Castilleja 
tayloriorum 



Castilleja 
irazuensis 




FIG. 7. Scrophulariaccac: erect herbs with consistently alternate leaves and unusual curved flowers (Castilleja) or 
simple, almost regular flowers (Capraria). 



12 



FIELDIANA: BOTANY 



Maurandya barclaiana 



Torenia 
fournieri 



Lophospermum 
erubescens 




Antirrhinum 
majus 



Penstemon /gent i anoi des 
Angel onia'angusti folium " ^ 



FIG. 8. Scrophulariaceae: wild and cultivated species with large colorful corollas and alternate or opposite leaves. 



BURGER: FLORA COSTARICENSIS 



13 



Key to Species of Agalinis and Anisantherina 

la. Flowers usually I/node, pedicels shorter than the calyx; distal leaves mostly alternate; stems and 

leaves smooth to the touch Agalinis albida 

Ib. Flowers usually 2/node, pedicels much longer than the calyx; distal leaves mostly opposite or 

suhopposite; stems and leaves slightly scabrous 2 

2a. Corolla 20-30 mm long, edge of corolla lobes with multicellular hairs; anther thecae equal in size 

and shape; calyx lobes narrow; pedicels without nodes or bracts Agalinis peduncularis 

2b. Corolla 10-16 mm long, edge of corolla lobes glabrous; anther thecae slightly unequal in size and 

shape; calyx lobes triangular; pedicels usually with a node or bracts (reduced leaves) near the middle 
Anisantherina hispidula 



Alectra Thunberg 
nomen conservandum 

REFERENCE H. Melchior, Die Gattung Alectra 
Thunb. Notizbl. Hot. Gart. Berlin 15: 423-447. 
1941. 

Erect annual herbs, hemiparasitic, turning dark 
when dried, unbranched to few-branched, with 
hispid or scabrous hairs often enlarged at the base. 
Leaves opposite or subopposite (or alternate 
above), reduced in size distally (rarely absent), 
simple and sessile or subsessile, margins serrate 
to dentate (entire), often with 3 prominent veins 
from the base. Inflorescences racemose or spi- 
cate, elongate, of solitary flowers in axils of distal 
leaves, pedicels (peduncles) short, with 2 (1) dis- 
tal bracts or bracts absent. Flowers with campan- 
ulate calyx, 10- veined, persisting and enclosing 
the fruit, with 5 acute to obtuse lobes, valvate in 
bud, equaling the tube in length; corolla usually 
yellow or orange, personate to subglobose or cam- 
panulate, as long or slightly longer than the calyx, 
5-lobed and slightly bilabiate, lobes rounded, 
shorter or equaling the tube; stamens of 2 shorter 
and 2 longer pairs, attached near the base of the 
corolla, included, filaments glabrous or puberu- 
lent, anthers 2-thecous, often barbate abaxially, 
disc annular and fleshy; ovary ovate or com- 
pressed, 2-locular with thick fleshy placenta and 
many seeds, style linear, elongate, and inflexed, 
stigma entire or bifid, thickened. Fruits loculici- 
dal capsules enclosed within the dry calyx, round- 
ed or compressed; seeds very numerous, small, 
with transparent exotesta often truncated and open 
at the 2 ends. 

A genus of 41 species, according to Melchior 
(1941, above). Nearly all the species are from Af- 
rica and India. The one American species is native 
to South America and the West Indies; it differs 
from the Old World species in having divergent 



rather than parallel anther thecae. Alectra is close- 
ly related to Melasma in the tribe Buchnereae and 
was once united with that genus. Few characters 
reliably distinguish the two. In Melasma the co- 
rolla is twice as long as the calyx and is more 
campanulate than that of Alectra. 

Alectra aspera (Cham. & Schldl.) L. O. Wil- 
liams, Fieldiana, Bot. 34: 118. 1972. Pedicu- 
laris melampyroides L. C. Rich., Actes Soc. 
Hist. Nat. Paris 1: 111. 1792. Glossostylis as- 
pera Cham. & Schldl., Linnaea 3: 22. 1828. 
Scrophularia fluminensis Veil., Fl. flumin., 263. 
1829. A. brasiliensis Benth. in DC., Prodr. 10: 
339. 1846. A. melampyroides (L. C. Rich.) 
Kuntze, Rev. Gen. PI. 2: 458. 1891, non A. me- 
lampyroides Benth. 1846. Melasma melampy- 
roides (L. C. Rich.) Pennell in Britton & Wil- 
son, Bot. Porto Rico 6: 188. 1925. A. fluminen- 
sis (Veil.) Steam, J. Arnold Arbor. 52: 636. 
1971. Figure 5. 

Herbs 0.3-1.5 m tall, stems erect and simple 
to many-branched, main stems 1.7-6 mm diam., 
scabrid with straight stiff whitish hairs 0.3-1.5 
mm long, short (0.1-0.2 mm) thin hairs also pre- 
sent. Leaves becoming gradually smaller distally, 
opposite, subopposite (rarely alternate distally), 
subsessile with petioles 0.5-4 mm long; leaf 
blades l-5(-7) cm long, 4-16(-25) mm wide, 
narrowly ovate-triangular to ovate-lanceolate or 
lanceolate, gradually narrowed to the acute or 
acuminate apex, margin coarsely serrate with 
teeth 0.3-2.5 mm high, base truncated or obtuse, 
surfaces scabrous with stiff whitish hairs 0.1-0.9 
mm long, venation subpalmate with midvein and 
2 prominent ascending basal 2 veins (triplivei- 
ned). Inflorescences of spike-like or raceme-like 
distal stems or with solitary flowers in the axils 
of smaller leaves (1-2 flowers/node), pedicles 1- 
2(-7) mm long, ca. 0.5 mm diam., bracts linear 



14 



FIELDIANA: BOTANY 



or absent, at the base of the calyx, ca. 3 mm long. 
Flowers with campanulate calyx 6-9 mm long (to 
12 mm in fruit), 5-8 mm wide at the mouth, calyx 
lobes -2-6 mm long, triangular and acute, with 
scabrous hairs along the margins and veins; co- 
rolla 10-13 mm long, included within the calyx 
or slightly exserted, campanulate, yellow, gla- 
brous, tube 5-8 mm long, lobes subequal; stamens 
with pubescent filaments, anthers 1.5 mm long, 
thecae divergent, without awns; ovary ca. 3 mm 
long, ovoid, style 6-8 mm long, curved, stigma 
lanceolate. Fruits 5-8 mm long, 4-6 mm diam., 
oblate-globose, included within the persisting ca- 
lyx; seeds 1-1.2 mm long, oblong-triangular with 
truncated ends, translucent yellowish, exotesta re- 
ticulate. 

Weeds of open sunny sites in lowland Central 
America. It is a recent introduction and is spread- 
ing. It has been collected near Upala and Villa 
Neilly, at Tarrazu, and on the lower slopes of the 
Cerros de Puriscal. The species ranges from Gua- 
temala and the West Indies to Brazil, Parguay, and 
Bolivia. 

Alectra aspera is recognized by its short erect 
stems, stiff pustulate-hispid leaves, tissues drying 
dark, solitary flowers in distal leaf axils, corolla 
tube only as long as the calyx, persisting campan- 
ulate-globose calyx, and minute seeds with trans- 
lucent testa. The corolla of this species is open for 
a very short time. It soon withers, closing over 
the anthers and the stigma. The anthers are borne 
close to the stigma, suggesting that the species is 
largely self-pollinating. 



Alonsoa Ruiz Lopez & Pavdn 

REFERENCES J. L6pez Guilldn, El Genero 
Alonsoa en el Peru: 1. Revision de las especies 
endemicas. Raymondiana 3: 155-246. 1970 
(1971). Brian Wrigley, A taxonomic revision of 
Alonsoa. Ph.D. diss., Univ. Connecticut, 1968. 

Herbs or subshrubs, annual or perennial, often 
with distal branching, usually woody at the base, 
stems quadrangular in cross-section, glabrous or 
sparsely minutely puberulent. Leaves opposite or 
ternate (leaf-like floral bracts alternate), sessile or 
petiolate, leaf blades ovate to linear, serrate or 
rarely entire, venation pinnate. Inflorescences ra- 
cemes, terminal or axillary to distal leaves (rarely 
flowers axillary to distal leaf pairs), floral bracts 
alternate along the rachis, proximal bracts leaf- 
like, pedicels solitary, well developed, and be- 
coming twisted, bracteoles absent. Flowers small. 



resupinate because of pedicel twisting, calyx 
deeply 5-parted. lobes narrow and slightly un- 
equal, glabrous or sparsely and minutely puberu- 
lent, valvate in bud; corolla rotate with very short 
tube, bilaterally symmetric with the 2 lower lobes 
small and divided nearly to the base, lateral lobes 
short and broad, upper lobe much larger and usu- 
ally held erect and convex, reddish to orange or 
purple; stamens 4, subequal, borne on the base of 
the corolla, filaments thick or slender, anthers 
closely positioned around the style, thecae parallel 
or divergent; ovary 2-locular, style curved up- 
ward, stigma capitate. Fruits capsules, ovate to 
oblong, septicidal, 2-valved, valves chartaceous, 
entire or bifid at the apex; seeds many, punctate- 
rugose or longitudinally ridged. 

Alonsoa is a Neotropical genus of six to fifteen 
species. The flowers are upside-down (resupinate) 
because of the twisted pedicel, with the result that 
the morphologically lower lobe is the upper lobe. 
The reddish to yellow resupinate flowers with en- 
larged median lobe are distinctive. The genus is 
placed in the tribe Hemimeridae, with Angelonia, 
but is probably more closely related to Scrophu- 
laria and Verbascum, with which it shares char- 
acters of seed and anther morphology. Alonsoa 
warscewiczii Regel is often used as a potted or- 
namental plant; only one species is found in the 
higher mountains of Central America. The genus 
is currently being studied by Fanny Astholm 
(GB). 



Alonsoa meridionalis (L.f.) Kuntze, Rev. Gen. 
PI. 2: 457. 1891. 

Scrophularia meridionalis L.f., Suppl. 280. 
1781. Figure 3. 

REFERENCE F Astholm & Y. Nyman, Morpho- 
metric variation in the Alonsoa meridionalis com- 
plex (Scrophulariaceae). Plant Syst. Evol. 193: 
53-68. 1994. 

Herbs or subshrubs 0.2-1.5 m tall, often 
woody at the base, usually with lateral branches, 
leafy stems 0.7-8 mm diam., glabrous or sparsely 
minutely puberulent (rarely densely puberulent at 
the node), with 4 prominent longitudinal ridges or 
wings (from decurrent petiole margins). Leaves 
opposite (sometimes alternate below the flowering 
nodes), petioles 3-15(-22) mm long, 0.4-1.7 mm 
diam., glabrous or with few hairs less than 0.3 
mm long, with lateral margins continuous with the 
blade margins; leaf blades 1.5-8(-ll) cm long, 



BURGER: FLORA COSTARICENSIS 



15 



1-4 cm wide, ovate- lanceolate to oblong-lanceo- 
late or narrowly ovate, apex acute, margin with 
4-7 acute teeth/cm, base obtuse to slightly cune- 
ate and decurrent, drying greenish or dark, sparse- 
ly puberulent on both surfaces, 2 veins 5-10/side, 
strongly ascending. Inflorescences terminal ra- 
cemes 3-30 cm long, flowers opposite or distally 
alternate, subtended by progressively smaller, 
leaf-like to subulate bracts 4-14 mm long, pedi- 
cels 6-18(-25) mm long, glabrous or with few 
minute (0.2 mm) gland-tipped hairs, upcurved in 
fruit. Flowers resupinate, calyx 3-6 mm long, di- 
vided to near the base, lobes 1-1 .7 mm wide, sub- 
equal or unequal, glabrous or with few minute 
hairs near the base; corolla 5-1 1 mm long, rotate, 
yellow to orange, tube ca. 2 mm long, with prom- 
inent lobes, median lobe to 15 mm long, larger 
than the lateral lobes; stamens 3-4 mm long, fil- 
aments 0.4-0.6 mm diam., anthers ca. 2 mm long; 
ovary 1.5-4 mm long, narrowly ovoid, style 1.5- 
2 mm long, stigma 0.8 mm wide. Fruits 7-15 mm 
long, 3.7-6 mm wide, narrowly ovoid with acute 
to acuminate apex, smooth, glabrous, pale brown, 
sulcate along the plane of dehiscence, septicidal; 
seeds many, 1.1-1.4 mm long, 1-1.2 mm diam., 
oblong, dark, with ca. 6 deep longitudinal sulci. 

Plants of open sites in high montane forest for- 
mations, 1600-3200 m elevation. Flowering in 
October-December. In Costa Rica they are found 
in the Cordillera Central and the Cordillera de Tal- 
amanca. The species ranges from southern Mex- 
ico to Bolivia. 

Alonsoa meridionalis is recognized by its ter- 
minal racemes with solitary, usually alternate 
flowers subtended by bracts gradually diminishing 
in size, unusual little flowers with red-orange co- 
rolla with enlarged median uppermost lobe, short 
thick filaments, large anthers, and capsules grad- 
ually narrowed to the apex. Because the pedicels 
are twisted 1 80, the flowers are upside-down (re- 
supinate). The leaves often have new shoots with 
small leaves in their axils. 



Angelonia Humboldt & Bonpland 

REFERENCE K. Barringer, A Revision of An- 
gelonia (Scrophulariaceae). Ph.D. diss. Univ. 
Connecticut, 1981. 

Erect herbs or subshrubs, annual or perennial, 
stems terete to 4-anguIate, simple or branching 
from the base, puberulent with multicellular hairs 
or glabrous. Leaves opposite or rarely alternate 
distally, petioles short or absent; blades ovate to 



linear, serrate or rarely entire, apex acute, vena- 
tion pinnate. Inflorescences racemes, terminal or 
axillary, bracts leaf-like to rounded, flowers 1-3/ 
axil, pedicels bibracteolate or ebracteolate. Flow- 
ers showy, calyx of 5 free or partly united sepals, 
sepals equal, lanceolate, entire, acute to acumi- 
nate; corolla strongly bilaterally symmetric, cu- 
pular-campanulate with short tube and 2 broadly 
flaring lips, bisaccate at the base of the median 
lobe, the sacs with a dense mat of glandular hairs 
within, lobes 5, upper lobes 2, the lower median 
(abaxial) lip 3-lobed, variously ornamented with 
a ridge or a crateriform palate and a bifid tooth; 
stamens 4, of 2 unequal pairs, held against the 
upper part of the corolla tube, filaments short, the- 
cae divaricate, without spurs; ovary ovoid, 2-loc- 
ular, ovules many, style longer than the ovary, 
stigma entire and minute. Fruits dry capsules, 
ovoid to broadly ellipsoid, chartaceous or leath- 
ery, septicidal, often secondarily loculicidal; seeds 
many, obconical, with a loose reticulate exotesta, 
endosperm absent. 

A genus of 26 species whose major concentra- 
tion is in the dry caatinga and cerrado formations 
of Brazil. The Central American and Caribbean 
species form a distinctive group within the genus. 
The genus is easily distinguished by the bisaccate 
corolla and the ornamented median corolla lobe. 
One species is found in Costa Rica; a second, An- 
gelonia ciliaris B. L. Robinson (with puberulent 
leaves auriculate at the base), ranges from south- 
ern Mexico and the Antilles to Nicaragua. 

Angelonia angustifolia Bentham in DC., Prodr. 
10: 254. 1846. Figure 8. 

Herbs 20-120 cm tall, erect, few-branched or 
unbranched, leafy stems 1.5-5 mm diam., terete 
or slightly 4-angled, sparsely puberulent with thin 
multicellular hairs 0.5-2 mm long at the nodes or 
along longitudinal lines. Leaves becoming small- 
er distally (intergrading with the floral bracts), op- 
posite or subopposite, sessile or subsessile, often 
clasping the stem; leaf blades 2-11 cm long, 4- 
20 mm wide, linear to narrowly elliptic-oblong or 
lanceolate, apex acute, margin obscurely serrate 
with short (0.2-0.6 mm) teeth 1-4/cm, base cu- 
neate to acute, drying chartaceous and brown or 
dark grayish green, surfaces subglabrous with few 
thin hairs, 2 veins 3-6/side and strongly ascend- 
ing. Inflorescences 5-40 cm long, flowers soli- 
tary in distal leaf/bract axils (usually 2/node), 
pedicels 4-18 mm long, 0.2-0.4 mm diam., gla- 
brous, ascending, subtended by 2 linear bracteoles 
0.5-2 mm long. Flowers glabrous, calyx 2-4 mm 



16 



FIELDIANA: BOTANY 



long, 1-2 mm wide at the base, narrowly ovoid 
to campanulate, calyx lobes 2-3.5 mm long, lan- 
ceolate to triangular; corolla 8-25 mm wide at the 
mouth, purple to lavender or bluish, minutely 
punctate, tube white to yellowish or green, often 
marked with white on the palate (basal entry of 
the throat), corolla lobes to 9 mm long, rounded; 
stamens included, filaments glandular-pubescent, 
anthers 2 mm wide, thecae divergent, equal, gla- 
brous; ovary glabrous. Fruits 4-6 mm long, 4-7 
mm diam., globose to ovoid-rounded with trun- 
cated base, glabrous; seeds 1.3-1.5 mm long, con- 
ic to oblong, exotesta strongly reticulated with 
prominent thin walls. 

Native to southern Mexico, this species has 
spread into much of Central America, where it is 
a favorite garden ornamental. In Costa Rica it has 
been collected in both the deciduous and ever- 
green lowlands and from gardens in the Meseta 
Central; it flowers throughout the year. The spe- 
cies is now cultivated throughout the world. 

Angelonia angustifolia is recognized by its 
small stature, narrow subglabrous opposite leaves, 
showy corolla with prominent spreading rounded 
lobes, and unusual seed surface. This species is 
probably pollinated by Centris bees, which collect 
a thick oil from the corolla sacs with specialized 
combs on their front legs. Common names used 
for this species are boca de la vieja (Guatemala), 
porto hello (Nicaragua), and angeldn (Colombia). 



Anisantherina Pennell 

Herbs, annual, erect with ascending branches, 
hispidulous with multicellular hairs with dark 
cross-walls, hemiparasitic on roots, turning dark 
when dried. Leaves opposite or subopposite dis- 
tally, becoming smaller or bract-like on distal 
stems, sessile, blades linear and entire, scabrous. 
Inflorescences of solitary flowers in the axils of 
reduced distal leaves (raceme-like with well-sep- 
arated flowers), pedicels often longer than the ca- 
lyx, bibracteolate. Flowers showy, calyx campan- 
ulate with 5 prominent equal lobes; corolla tubu- 
lar-campanulate, slightly bilabiate, glabrous, pink 
to purple, tube straight or curved, expanded dis- 
tally into a broad throat, 2-lipped, upper lip 2- 
lobed, lower lip 3-lobed, lobes rounded, subequal, 
spreading; stamens 4, of 2 unequal pairs, inserted 
near the mouth of the corolla tube, shorter than 
the corolla, anterior filaments longer than the pos- 
terior, pilose above, anthers 2-thecous with un- 
equal divergent thecae, glabrous; ovary ovoid, 
glabrous, 2-locular, style longer than the ovary. 



straight and slender, stigmas linear and lateral on 
the liguliform style apex. Fruits globose capsules, 
chartaceous, loculicidal, style base persistent, pla- 
centa persistent; seeds many, linear, exotesta re- 
ticulate. 

Anisantherina is a monotypic Neotropical genus 
related to African genera in the tribe Buchnereae. 
Pennell distinguished Anisantherina from Agalinis 
by its bibracteolate pedicels, unequal anther thecae, 
and narrow oblong-linear seeds. Canne (1980) 
showed that the structure and ornamentation of An- 
isantherina seeds are also distinctive. This species 
was once placed in Gerardia, but that name is no 
longer valid; see the discussion under Agalinis. 

Anisantherina hispidula (Man.) Pennell, Mem. 
Torrey Bot. Club 16: 106. 1920. Gerardia his- 
pidula Mart., Nov. Gen. Sp. PI. 3: 13. 1829. 
Agalinis hispidula (Mart.) D'Arcy, Ann. Mis- 
souri Bot. Card. 65: 4. 1978 (1979). Figure 4. 

Annual herbs, 30-50 cm tall, unbranched or 
with few branches arising in the lower half, leafy 
internodes 0.5-1.5 mm diam., with few short 
(0.2-0.4 mm) stiff hispidulous hairs; usually dry- 
ing dark. Leaves opposite or subopposite (rarely 
alternate distally), sessile or subsessile, sometimes 
clasping the stem; leaf blades 8-80 mm long, 
0.5-4 mm wide, linear, entire, scabrous with short 
(ca. 0.2 mm) whitish, hairs above and along the 
margin, 2 veins obscure. Inflorescences of soli- 
tary flowers in axils of reduced distal leaves, ra- 
ceme-like with 8-14 flowers, pedicels 10-35 mm 
long, 0.3-0.4 mm diam., glabrous, usually with a 
node or a pair of small (0-2 mm) bracts near the 
middle. Flowers glabrous, drying dark, calyx tube 
4-6 mm long, 3-5 mm diam., campanulate-tu- 
bular, abruptly rounded and truncated at the base, 
calyx lobes 1.2-2.5 mm long, triangular and 
acute; corolla 10-15 mm long, campanulate, pink 
to light purple with darker spots within, tube 8- 
10 mm long, lobes 2-3 mm long, glabrous, round- 
ed; filaments 2-3 mm long, anther thecae unequal, 
divergent. Fruits 5-9 mm long, 4-7 mm diam., 
abruptly rounded at apex and base (short-cupu- 
late), glabrous, drying black, slightly exserted be- 
yond the thin persisting calyx; seeds 0.6-0.8 mm 
long, 0.2-0.3 mm diam., linear, dark brown. 

Rarely collected plants of open sunny seasonal 
pools and moist savannas in deciduous or ever- 
green forest areas, 0-600 m elevation. We have 
seen only two collections from Costa Rica, both 
from near La Cruz in northern Guanacaste (L. D. 
Gomez 18965 & J. Gomez-Laurito 9097); flow- 



BURGER: FLORA COSTARICENSIS 



17 



ering and fruiting in November. The species rang- 
es from southern Mexico and Cuba to Brazil. 

Anisaniherina hispidula is recognized by its 
short wiry habit, parts drying dark, linear sca- 
brous leaves, glabrous distant upright pink flow- 
ers, and rounded fruits. It is confined to wet hab- 
itats in open sunny savannas. See the key and dis- 
cussion under Agalinis. 



Antirrhinum Linnaeus 

REFERENCE D. Sutton, A Revision of the Tribe 
Antirrhineae. British Museum (Natural History) 
& Oxford Univ. Press, 1988. 

Herbs, annual or perennial, erect or procum- 
bent, usually few-branched, stems terete, often 
glandular pubescent. Leaves opposite to subop- 
posite, sometimes alternate on distal stems, sessile 
or short-petiolate, leaf blades lanceolate to ovate, 
usually narrow, entire to denticulate. Inflores- 
cences usually showy terminal racemes or with 
solitary flowers in axils of distal bracts or reduced 
leaves, pedicels ebracteolate. Flowers with calyx 
united at the base, sepals 5, imbricate in bud; co- 
rolla bilaterally symmetric, 2-lipped and very ir- 
regular, corolla tube gibbous or saccate at the base 
(not spurred), broader than high, upper lip erect 
and 2-lobed, lower lip spreading and 3-lobed, 
base of the lower lobe forming a palate and press- 
ing against the front of the throat (closing off easy 
entrance to the tube); stamens 4, of 2 unequal 
pairs, included in the corolla tube, filaments slen- 
der and slightly dilated at the apex, anther cells 
divergent, staminode absent; ovary ovoid, 2-loc- 
ular, many-ovulate, style filiform, longer than the 
ovary, stigma bilobed. Fruits ovoid or globose 
capsules, opening below the apex by pores or slits 
(septicidal); seeds many, oblong, truncated, exo- 
testa smooth or rugose (without wings). 

A genus of about 42 species from the western 
United States, northwestern Mexico, Europe, and 
the Mediterranean region of Eurasia. The genus is 
a member of tribe Antirrhineae, with Cymbalaha 
and Linaria. A number of species and many va- 
rieties have been developed as garden ornamen- 
tals with bright red, purple, yellow, or white 
forms. The mouth of the corolla is closed by the 
curved palate of the lower lip, which is held 
against the base of the upper lip. Thus, bees must 
force their way into the interior of the corolla to 
gain access to nectar. The following species is 
commonly cultivated in cool highland gardens 
throughout Central America. 



Antirrhinum ma jus L., Sp. PL 617. 1753. Figure 8. 

Herbs, erect, 0.4-1 m tall, leafy stems 2-8 mm 
diam., glabrous and terete. Leaves opposite or ter- 
nate below, alternate or rarely ternate distally, pet- 
ioles 1-14 mm long but poorly differentiated from 
the blade; leaf blades 2-9 cm long, 3-16 mm 
wide, lanceolate to ovate-lanceolate or linear, 
apex acute, margin entire, base gradually nar- 
rowed, drying stiffly chartaceous and greenish. 
Inflorescences to 30 cm long, racemes of alter- 
nate flowers, rachis usually densely puberulent 
with gland-tipped hairs ca. 0.3 mm long, bracts 
ovate, 2-10 mm long, acute, pedicels 2-10 mm 
long. Flowers showy, calyx glandular puberulent, 
sepals 5, 6-8 mm long, ovate to rounded; corolla 
2.5-4 cm long, pink to red or purple, externally 
glandular hairy, the palate closing the throat, yel- 
low within. Fruits 1-1.5 cm long, oblong cap- 
sules with unequal valves, opening near the apex, 
glandular pubescent to glabrous; seeds light 
brown, reticulate-tuberculate. 

Antirrhinum majus is a native of the Pyrenees 
of northern Spain and southern France. It is wide- 
ly cultivated in temperate and subtropical regions. 
It can be recognized by its showy racemes and 
slightly saccate corolla with an enlarged palate 
that "closes" the throat. In some horticultural 
forms the throat is open or the flowers are highly 
modified. Common names are "snapdragon," 
boca de leon, and boca de dragon. 



Bacopa Aublet 

REFERENCE F. W. Pennell, Reconsideration of 
the Bacopa-Herpestis problem of the Scrophular- 
iaceae. Proc. Acad. Nat. Sci. Philadelphia 98: 83- 
98. 1946. 

Herbs, erect to decumbent or procumbent, usu- 
ally growing in moist soil or in standing water, 
stems simple or profusely branched, glabrous or 
pubescent, often glandular punctate. Leaves op- 
posite, sessile or petiolate, leaf blades often slight- 
ly succulent, entire to dentate (or dissected), ve- 
nation pinnate or palmate, the minor venation usu- 
ally obscure. Inflorescences of 1-6 flowers in leaf 
axils, less often the distal flowering stems race- 
mose or spicate (cymose, paniculate), pedicels 
short or absent, bracteoles present or absent at the 
base of the calyx. Flowers usually with 5-parted 
calyx, sepals subequal to strongly unequal with 
the 3 adaxial sepals usually much wider than the 



18 



FIELDIANA: BOTANY 



abaxial, imbricate in bud (if 4-parted, the calyx 
tube equaling the lobes); corolla tubular, strongly 
or weakly bilabiate, blue-violet to white, tube cy- 
lindriq, the lips spreading, with 3, 4, or 5 lobes, 
the upper lip exterior in bud and 2-lobed or emar- 
ginate (1-lobed), the lower lip 2- or 3-lobed; sta- 
mens (5) 4 (2, 3), usually in 2 unequal pairs, in- 
serted on the upper half of the corolla tube, in- 
cluded, anthers approximate or distant, thecae 
contiguous, parallel or divergent, staminode ab- 
sent; ovary 2-locular, style usually straight, stigma 
terminal, bilobed or entire. Fruits dry capsules, 
globose to ovoid, bisulcate, loculicidal, often sec- 
ondarily septicidal into 4 valves; seeds many, 
small, oblong, longitudinally reticulate. 

Bacopa is a genus of about 50 to 60 species, 
widespread in warm temperate and tropical areas 



throughout the world. It is most diverse in South 
America. The species are varied but they can usu- 
ally be recognized by their small opposite leaves, 
strongly unequal sepals, and preference for wet 
habitats. They are often found in open sunny sites 
in shallow standing water of seasonal pools, along 
the edges of watercourses, and in moist savannas. 
Most of the species described below are rarely 
collected in Costa Rica, perhaps because of their 
seasonally inundated habitats. Because the sepals 
differ so much in size, they may appear to be 
bracts enclosing the flower. The inner sepals are 
usually much narrower than the outer. The brae - 
teoles are borne at the apex of the pedicel when 
present. The synonymy of this genus is very large; 
D'Arcy (1979, p. 183) provides a long list of ge- 
neric synonyms. 



Key to the Species of Bacopa 

la. Flowers sessile or subsessile, pedicels up to 2 mm long; plants erect; rarely collected in Costa Rica 

2 

Ib. Flowers borne on conspicuous pedicels > 2 mm long; plants erect, prostrate or floating; rare or 

common 4 

2a. Stems densely puberulent with hairs to 1.3 mm long; [flowers often in dense verticels of 3-12 

flowers/node in the axils of leaves; outer calyx lobes 3-4 mm long] B. axillaris 

2b. Stems glabrous or minutely puberulent, the hairs < 0.5 mm long 3 

3a. Stems puberulent; fruits with smooth surfaces; outer sepals 1-2 mm long; flowers in distal 

fascicles separated by conspicuous slender internodes; leaves subentire ... B. monnierioides 

3b. Stems glabrous; fruits with pitted-reticulated surfaces; outer sepals 2-3.5 mm long; flowers 

often in distal crowded fascicles and spike-like with obscure internodes; leaves serrate 

B. sessiliflora 

4a. Plants mostly erect with few to many distal branches, rooting only at the base, stems glabrous, 
leaves serrate, sessile and auriculate at the base (petiolate in B. lacertosa), opposing leaves not 

united and usually without an interpetiolar line 5 

4b. Plants usually prostrate, floating or creeping, with few or no distal branches, rooting at base and 
lower nodes, stems glabrous or puberulent; leaves serrate or entire, sessile or petiolate, opposing 
leaves slightly united across the stem to form an interpetiolar line or ridge or the node pubescent 

7 

5a. Outer sepals to 5 mm long, sepals usually narrowed at the base, rarely covering the fruits, the 
venation not conspicuously raised; distal flowers subtended by greatly reduced leaves (ca. 8 

mm long) and easily seen [distal internodes usually longer than the adjacent leaves] 

B. laxiflora 

5b. Outer sepals to 8 mm long, rounded at the base, usually covering the fruits, venation often 
conspicuous on the outer surface; distal flowers subtended by normal-size or reduced leaves, 

usually easily visible 6 

6a. Outer sepals not developing a lustrous surface, venation only slightly elevated; plants to 40 

(-50) cm tall; distal stems glabrous or glandular puberulent B. bacopoides 

6b. Outer sepals with smooth lustrous surface and conspicuous elevated venation; plants to 70 cm 

tall; distal stems glabrous [Belize to Nicaragua] B. lacertosa 

7a. Internodes densely villous with hairs to 1 .5 mm long; leaves often suborbicular [outer sepals 4-6 

mm long, ciliolate; commonly collected in Costa Rica] B. salzmanii 

7b. Internodes glabrous or with thin hairs < 0.5 mm long; leaves broadly to narrowly obovate (rarely 
suborbicular) 8 



BURGER: FLORA COSTARICENSIS 



19 



8a. Braeteoles 2, slender, below the calyx; outer sepals 5-7 mm long; fruits 4-7 mm long; flowers 

usual 1\ I/node; pedicels to 30 mm long; internodes glabrous B. monnieri 

8b. Braeteoles none; outer sepals 2.5-4 mm long; fruits 2.3-4 mm long; flowers 1-4/node; pedicels to 

1 8 mm long; internodes glabrous or puberulent 9 

9a. Sepals (4) 5, unequal and united only at the base; leaves usually cuneate or rounded at the base, 

sessile; widespread B. repens 

9b. Sepals 4, subequal and united in the lower half; leaves cuneate with a narrowed petiole-like base; 

rarely collected B. egensis 



Bacopa axillaris (Benth.) Standl., J. Wash. Acad. 
Sci. 15: 460. 1925. Herpestis axillaris Benth. in 
DC, Prodr. 10: 396. 1846. Monniera axillaris 
(Benth.) O. Kuntze, Rev. Gen. PI. 463. 1891. 
Caconapea axillaris (Benth.) Pennell, Proc. 
Acad. Nat. Sci. Philadelphia 72: 152. 1920. Fig- 
ure 1. 

Erect herbs 10-30 cm tall, aquatic or paludal, 
rooting mostly at the base, leafy stems 1.3-4 mm 
diam., spongy, villous with thin multicellular hairs 
to 1 .3 mm long. Leaves opposite, sessile with op- 
posing leaves slightly united at the base and form- 
ing a line or ridge across the stem (clasping the 
stem); leaf blades 1.2-4.8 cm long, 2-12 mm 
wide, oblanceolate to narrowly obovate or nar- 
rowly elliptic-oblong, apex bluntly obtuse to 
acute, distal % of the margin serrate, teeth 0.2-0.5 
mm high, 1-2.3 mm wide, base gradually nar- 
rowed and cuneate, drying yellowish brown or ol- 
ive green, glabrous, conspicuously pellucid punc- 
tate beneath, venation pinnate. Inflorescences 
verticellate, of dense axillary fascicles with 3-12 
flowers/node, pedicels 0.2-1.5 mm long, expand- 
ed at the apex and with 2 small (0.4-0.7 mm) 
slender bracteoles. Flowers glabrous externally, 
outer sepals 3-4 mm long, 1 .5-3 mm wide, ovate 
with rounded to obtuse apex, palmately veined, 
punctate, glabrous or ciliate; corolla 3-4 mm 
long, white, slightly exserted, the upper lip 1- 
lobed; stamens 4, inserted in the upper half of the 
tube; ovary ca. 1 mm long, style 1.5-2 mm long. 
Fruits 2-3 mm long, ca. 1.4 mm diam., very nar- 
rowly ovoid or conical, 4-valved; seeds 0.5-0.6 
mm long, 0.2-0.3 mm wide, oblong or somewhat 
curved, brown, reticulate with parallel longitudi- 
nal ridges. 

Plants of swamps and the muddy edges of 
standing water in seasonally deciduous and ever- 
green forest areas, 0-500 m elevation (to 1 500 m 
in Guatemala). Rarely found north of Panama, the 
species has been collected near Bagaces and along 
the Rio Grande de Tarcoles in Costa Rica; it flow- 
ers in September. The species ranges from Gua- 
temala to Colombia. 



Bacopa axillaris is recognized by the pubescent 
stems, sessile oblanceolate serrate leaves, verticil- 
late flower clusters with small subsessile flowers, 
and wet habitat. The outer sepals are often flat, 
translucent, and conspicuously punctate. 

Bacopa bacopoides (Benth.) Pulle, Enum. PI. Su- 
rinam. 415. 1906. Herpestis bacopoides Benth. 
in DC., Prodr. 10: 399. 1846. B. bracteolata 
Pennell ex Standl., Contrib. U.S. Natl. Herb. 27: 
336. 1927. 

Erect herbs to 50 cm tall, aquatic or paludal, 
leafy stems 1.4-4.5 mm diam., glabrous, terete or 
slightly 4-angled, with minute sessile glands, node 
lacking interpetiolar lines. Leaves sessile, clasp- 
ing the stem, opposing leaves not united at the 
base; leaf blades 1.5-3.8 cm long, 2.5-7 mm 
wide, linear-oblanceolate to linear-oblong or 
ovate-elliptic, gradually narrowed in the lower 
half but slightly expanded and subauriculate at the 
base, apex acute, margin serrate with short (0.2- 
0.3 mm) broad (1-3 mm) teeth, drying dark 
brown, glabrous, minutely punctate, venation pin- 
nate, obscure. Inflorescences of 1-2 axillary 
flowers (2-4 flowers/node), pedicels 2-7 mm 
long, 0.2-0.3 mm diam., minutely papillate-pu- 
berulent with whitish hairs, slightly expanded at 
the apex, paired bracteoles 1-2 mm long or ab- 
sent, linear. Flowers glabrous externally or the 
outer sepals sometimes puberulent, outer 3 sepals 
4-8 mm long, 3-5 mm wide, broadly ovate, 
rounded but obtuse at the base, with raised ve- 
nation; corolla 4-7 mm long, white, stamens 4, 
inserted near the middle of the corolla tube, an- 
thers ca. 0.6 mm long; ovary glabrous, style ca. 
2 mm long. Fruits ca. 3 mm long, globose, en- 
closed within the enlarged (to 15 mm) wing-like 
sepals. 

Partly aquatic plants of marshes and wet areas 
at low elevations. The species has not been col- 
lected in Costa Rica but is common in central 
Panama and flowers in December-January in Nic- 
aragua. This species ranges from Guatemala to 
Brazil. 



20 



FIELDIANA: BOTANY 



Bacopa bacopoides is recognized by its erect 
habit, sessile narrow serrate leaves, one or two 
pedicellate flowers in leaf axils, large rounded 
outer sepals enclosing the fruits, and wet habitat. 

Bacopa egensis (Poepp.) Pennell, Proc. Acad. 
Nat. Sci. Philadelphia 98: 96. 1946. Hydran- 
thelium egense Poepp. in Poepp. & Endl., Nov. 
Gen. Sp. PI. 3: 75, tab. 287. 1845. 

Aquatic or terrestrial herbs, floating or pros- 
trate on wet soil, with short erect flowering stems, 
rooting at proximal nodes, leafy stems terete, pu- 
berulent on the upper surfaces. Leaves opposite, 
larger in aquatic plants, petioles not clearly distin- 
guished from the cuneate base; leaf blades 7-23 
mm long, 3-14 mm wide, obovate to spatulate, 
rhombic or suborbicular, apex obtuse to rounded, 
margin serrate distal ly, base cuneate from the 
middle of the blade, puberulent beneath, venation 
palmate with 5-7 1 veins. Inflorescences of sol- 
itary axillary flowers (usually 2/node), pedicels 3- 
6 mm long, slender, glabrous or puberulent, brac- 
teoles absent. Flowers with 4-parted calyx 2-4 
mm long, lobes united below the middle, sube- 
qual, obtuse, subglabrous or the outer surface mi- 
nutely puberulent; corolla 3-5 mm long, funnel- 
form, white, with 3 unequal lobes rounded distal- 
ly; fertile stamens 3; ovary with many ovules, 
stigma bilobed. Fruits 3-4 mm long, ovoid, bi- 
valved with membranaceous walls; seeds many, 
cylindric-curved, rugulose. 

Rarely collected plants of standing water and 
wet depressions in southern Nicaragua and north- 
eastern Costa Rica, 0-200 m elevation. Flowering 
and fruiting in September. The species is also 
found in Colombia and Brazil. 

Bacopa egensis is recognized by its unusual 
palmately veined leaves with cuneate or attenuate 
base, flowers with four equal sepals, three-lobed 
white corollas, and curved cylindric seeds. The 
terrestrial plants of this species have smaller 
leaves with more puberulence than their aquatic 
conspecifics. We have not seen Costa Rican ma- 
terial and follow the report in Flora of Nicaragua. 

Bacopa lacertosa Standley is found along the 
Caribbean coast, from Belize to Nicaragua. It has 
erect stems to 70 cm tall, lanceolate leaves to 70 
X 15 mm, axillary flowers on short pedicels, and 
broadly ovate outer sepals (to 8 mm long) with a 
parchment-like texture and lustrous surface on 
which the venation is conspicuously elevated. 
Sutton and Hampshire (Flora of Nicaragua, 1995) 
suggested that this species may be conspecific 



with the African B. decumbens (Fernald) F. N. 
Williams. 

Bacopa laxiflora (Benth.) Wettst. ex Edwall. Bol. 
Commiss. Geogr. Estado Sao Paulo 13: 180. 
1897. Herpestis laxiflora Benth. in DC, Prodr. 
10: 396. 1846. H. auriculata Robinson, Proc. 
Am. Acad. Arts 26: 172. 1891. B. auriculata 
(Robinson) Greenman, Publ. Field Mus. Nat. 
Hist. Bot. Sen 2: 262. 1907. Caconapea auri- 
culata (Robinson) Pennell, Proc. Acad. Nat. 
Sci. Philadelphia 72: 150. 1920. Mella laxiflora 
(Benth.) Pennell, Notul. Nat. Acad. Nat. Sci. 
Philadelphia 46: 1. 1940. 

Erect herbs 10-40 cm high, aquatic or paludal, 
rooting only at the base, main stem simple or with 
few lateral branches, leafy stems 0.5-3 mm diam.. 
glabrous, 4-angled with longitudinal ridges. 
Leaves sessile, smaller and bract-like at distal 
flowering nodes, often appearing perfoliate but 
the opposing leaves not united across the stem; 
leaf blades 6-30 mm long, 2-10 mm wide, lan- 
ceolate to lanceolate-oblong or ovate-lanceolate, 
apex acute, margin serrate, teeth ca. 0.3 mm high 
and 1-2 mm wide, base rounded to auriculate. 
drying grayish green, glabrous and punctate, ve- 
nation pinnate. Inflorescences of solitary axillary 
flowers (1 -2/node) or racemose and terminal (re- 
sembling panicles in some collections), pedicels 
3-8 mm long, minutely papillate puberulent, 
ebracteolate at base of calyx or with 2 slender 
bracteoles 1-1.5 mm long. Flowers glabrous, out- 
er sepals 3-5 mm long, 1-3.3 mm wide, ovate, 
usually narrowed at the base, acute, palmately 
veined, inner sepals linear; corolla 7-10 mm 
long, lilac to purple, bilabiate, tube ca. 5 X 1.7 
mm, lobes 2-3 mm long; stamens 4, inserted on 
the upper half of the tube, anthers 0.5 mm long; 
style 2.5 mm long. Fruits 2.5-4 mm long, subgl- 
obose, surface slightly reticulate with minute pits; 
seeds ca. 0.4 X 0.2 mm, oblong with 1 or 2 trun- 
cated ends, longitudinally ridged/reticulate, 
brown. 

Plants of wet sites and shallow standing water, 
in lowland deciduous and evergreen formations, 
0-300 m. The species has been collected near La 
Cruz, Guanacaste Province; it flowers in Decem- 
ber-January. The species ranges from Mexico to 
Brazil but is rarely collected in Central America. 

Bacopa laxiflora is recognized by its short erect 
habit, wet habitat, sessile serrate leaves appearing 
to clasp the stem, solitary axillary flowers form- 
ing distal racemes, and globose or ovoid fruits. 



BURGER: FLORA COSTARICENSIS 



21 



Stems and leaves are usually glabrous, but the 
pedicels arc minutely papillate puberulent. 

Bacopa monnieri (L.) Pennell, Proc. Acad. Nat. 
Sci. Philadelphia 98: 94. 1946. Lysimachia 
monnieri L., Sp. PI. Cent. 2: tab. 9. 1756. Fig- 
ure 1. 

Prostrate to procumbent herbs to 30 cm tall, 
rooting from the lower nodes, leafy stems 0.7-2.5 
mm diam., glabrous, nodes marked by interpetio- 
lar lines. Leaves slightly succulent, subsessile or 
with poorly defined petioles 0.5-2 mm long, 
clasping the stem at the base; leaf blades 3-18 
mm long, 1 .5-7 mm wide, obovate to oblong-ob- 
ovate, oblanceolate, or spatulate, apex rounded, 
margin entire, base gradually narrowed and cu- 
neate, drying yellowish green, glabrous, often 
punctate, venation pinnate but obscure (some- 
times appearing tripliveined). Inflorescences of 
solitary axillary flowers, usually with only 1 flow- 
er/node, pedicels 7-30 mm long, 0.3-0.6 mm 
diam., glabrous, bracteoles 2, opposite, 1.5-3 mm 
long, 0.4-0.7 mm wide, near the apex of the ped- 
icel and resembling the larger sepals in texture. 
Flowers glabrous, the 2 outer sepals 5-7 mm 
long, 2-3 mm wide, 3 inner sepals narrower, 
acute; corolla 6-10 mm long, white to lavender, 
blue or pale purple, lobes rotate and essentially 
regular, ca. 2 cm wide, tube 3-5 mm long; sta- 
mens 4, filaments arising near the apex of the 
tube, glabrous, anthers ca. 1 .5 mm long; ovary ca. 
4 mm long, ovate-oblong, style ca. 4 mm long, 
stigma flattened, ca. 0.8 mm wide. Fruits 4-7 mm 
long, ovoid, enclosed within the persisting sepals; 
seeds ca. 0.5 mm long, reddish brown, longitu- 
dinally reticulate. 

Plants of open sunny sites in marshes, sandy 
stream edges, and standing water, often creeping 
on wet mud and tolerant of some salinity, 0-1000 
m elevation. The species has rarely been collected 
in Costa Rica, where it is usually found near the 
coasts. It probably flowers throughout the year. 
We have seen a single sterile collection (Crow 
9465 INBIO) from 820 m in Alajuela. The species 
ranges from the southeastern United States to Ar- 
gentina. 

Bacopa monnieri is recognized by its small en- 
tire obovate or spatulate leaves, glabrous parts, 
narrow bracteoles beneath the sepals, somewhat 
larger flowers, and ability to survive in slightly 
salty water. The species has been called verdolaga 
in Nicaragua. 



Bacopa monnierioides (Cham.) Robinson, Proc. 
Am. Acad. Arts 44: 614. 1909. Ranaria mon- 
nierioides Cham., Linnaea 8: 31. 1833. Herpes- 
tis ranaria Benth. in J. D. Hook., Companion 
Bot. Mag. 2: 57. 1836, based on R. monnierioi- 
des Cham. B. ranaria (Benth.) Chod. & Hassl., 
Bull. Herb. Boissier ser. 2, 4: 288. 1904. Ca- 
conapea parviflora Pennell, Proc. Acad. Nat. 
Sci. Philadelphia 72: 152. 1920. B. parviflora 
(Pennell) Pennell ex Standl., Contrib. U.S. Natl. 
Herb. 27: 336. 1928. B. parviflora (Pennell) 
Standl. ex L. O. Williams, Fieldiana Bot. 34: 
118. 1972. 

Erect herbs to 40 cm tall, aquatic or in wet soil, 
with much distal branching, rooting mostly at the 
base, leafy stems 0.5-2.5 mm diam., minutely pu- 
berulent with slender crooked hairs 0.1-0.4 mm 
long on upper surfaces. Leaves sessile, opposing 
leaves not united across the stem but sometimes 
producing a line across the stem, distal leaves 
conspicuously smaller than leaves of the main 
stem; leaf blades 8-30 mm long, 2-7 mm wide, 
oblong-lanceolate to narrowly oblong, apex ob- 
tuse, margin subentire or obscurely dentate with 
teeth 1-2 mm wide, drying yellowish brown or 
dark grayish, glabrous, glandular punctate on both 
surfaces, venation subpalmate or tripliveined. In- 
florescences mostly fasciculate, of (1)2-4 axillary 
flowers, 2-8 flowers/node, pedicels ca. 0.5 mm 
long, bracteoles minute (0.3 mm) at apex of ped- 
icel. Flowers glabrous, outer sepals 1-2 mm long, 
0.5-1 mm wide, narrowly ovate-oblong, with an 
unusual glandular-pitted surface; corolla 1.5-2.5 
mm long, tubular, white or bluish white, upper 
lobe entire, lower lip 3-lobed; stamens 4, inserted 
near the apex of the tube; ovary glabrous. Fruits 
ca. 1 .5 mm long, smooth, enclosed within the stiff 
persisting sepals (to 2.5 mm long); seeds 0.6-0.7 
mm long, 0.2-0.3 mm thick, oblong-ellipsoid, 
longitudinally ridged, reddish brown. 

Plants of inundated areas, the margins of water 
bodies, and wet savannas, 0-1500 m in Central 
America. In Costa Rica, this species has been col- 
lected only in lowland Guanacaste. The species 
ranges from Guatemala to Paraguay. 

Bacopa monnierioides is recognized by its erect 
branching habit, sessile lanceolate-oblong leaves 
with subserrate margins, small fasciculate flowers, 
and smooth fruit. The glandular pit at the base of 
the outer sepal is unusual but difficult to see. 
Compare B. axillaris and B. sessiliflora. 



22 



FIELDIANA: BOTANY 



Bacopa repens (Sw.) Wettst. in Engl. & Prantl, 
Nat. Pflanzenfam. 4(3b): 76. 1895. Gratiola re- 
pens Sw., Prodr. 14. 1788. Herpestis repens 
(Sw.) Schldl. & Cham., Linnaea 5: 107. 1830. 
Macuillamia limosa Pennell, Proc. Acad. Nat. 
Sci. Philadelphia 72: 158. 1920. Bacopa limosa 
(Pennell) Standl., Contrib. U.S. Natl. Herb. 27: 
336. 1928. M. repens (Sw.) Pennell, Acad. Nat. 
Sci. Philadelphia Monogr. 1: 60. 1935. B. cur- 
tipes Standl. & L. O. Williams. Ceiba 3: 129. 
1952. Figure 1. 

Herbs, aquatic or terrestrial, floating to pros- 
trate or repent, to 40 cm long, rooting from lower 
nodes, stems 0.4-3 mm diam., glabrous or with 
short (0.1-0.3 mm) thin hairs distally. Leaves ses- 
sile, bases of opposing leaves partly united and 
obscuring the node; leaf blades 6-20(-30) mm 
long, 3-15(-20) mm wide, obovate to obovate- 
oblong or suborbicular, apex rounded or obtuse, 
margin entire, base cuneate, drying yellowish 
green to dark green or blackish, glabrous except 
near the base, venation palmate with 3-11 major 
veins. Inflorescences of axillary flowers, 2-4 
flowers/node, pedicels 6-18 mm long, 0.2-0.3 
mm diam., glabrous or minutely puberulent, brac- 
teoles absent or minute. Flowers with 4 or 5 se- 
pals, outer sepals 2.5-4 mm long, 0.7-1.5 mm 
wide, narrowly oblong, rounded to obtuse at the 
apex, margin often ciliolate, inner sepals narrow- 
er; corolla 3-4 mm long, white or pale violet, 
campanulate and regular or nearly so, throat 
sometimes marked with yellow, 2-lipped, the up- 
per lip 2-lobed; stamens 4, filaments inserted near 
the apex of the tube, anthers ca. 0.7 mm long; 
ovary ca. 1.3 mm long, style 1-3 mm long, slen- 
der, stigma capitate, slightly 2-lobed. Fruits 2.3- 
4 mm long, ca. 2 mm diam., glabrous, enclosed 
within the persisting (3-4 mm long) sepals; seeds 
ca. 0.4 mm long, oblong, dull whitish to brown, 
surface reticulate. 

Aquatic plants of lake edges and marshes, in 
both evergreen and deciduous forest areas, 0- 
1000 m elevation. Rarely collected in Costa Rica, 
this species is found in the Caribbean lowlands 
and on the Pacific coast in seasonal ponds at Ba- 
gaces and Palo Verde National Park. Flowering in 
August-October. The species ranges from south- 
ern Mexico and the West Indies to Argentina. 

Bacopa repens is recognized by its aquatic hab- 
itat, glabrous or sparsely puberulent stems, ob- 
ovate sessile leaves slightly united (across the 
stem) at the base, and small flowers. The distal 
leaves and stems are often floating and can be- 
come a floating mat. This species may be vege- 



tatively very similar to B. monnieri, but that spe- 
cies has larger flowers and fruits and lacks brac- 
teoles. Compare B. salzmannii, which occurs in 
similar habitats but has villous stems, larger flow- 
ers, and separate leaf bases. 

Two collections from water 1 .5 m deep at Palo 
Verde National Park (G. Crow 5977 & 60603) are 
provisionally placed here. These collections have 
larger (20-40 mm) leaves, longer fruits (3-5 
mm), and linear-oblong seeds to 0.7 mm long. 
They conform to the description of B. valerii 
Standl. & L. O. Williams (Ceiba 1: 163, 1950) 
based on material from 20 m elevation in we'stern 
Honduras. Although quite distinctive, it seems 
likely that all these plants represent no more than 
an unusually robust form of B. repens. 

Bacopa salzmannii (Benth.) Wettst. ex Edwall, 
Bol. Commiss. Geogr. Estado Sao Paulo 13: 
176, 181. \S91.ScrophulariaprocumbensVe\\., 
Fl. tl nun n. Ic. 6, tab. 85. 1827, non B. procum- 
bens (Miller) Greenman. Herpestis salzmannii 
Benth. in J. D. Hook., Companion Bot. Mag. 2: 
58. 1836. B. salzmannii (Benth.) Chod. & 
Hassl. Bull. Herb. Boissier Ser 2,4: 290. 1904. 
Monocardia humilis Pennell, Proc. Acad. Nat. 
Sci. Philadelphia 72: 157. 1920. M. violacea 
Pennell, loc. cit. 156. 1920. B. humilis (Pennell) 
Standl., J. Wash. Acad. Sci. 15: 460. 1925. B. 
violacea (Pennell) Standl., loc. cit. 460. 1925. 
Herpestis ciliata Pennell, Notul. Nat. Acad. Sci. 
Philadelphia 46: 2. 1940. Figure 1. 

Herbs, aquatic or terrestrial, prostrate or de- 
cumbent, sometimes floating or forming mats, 
rooting from lower nodes, stems succulent, 1-2 
mm diam., densely hirsute with thin multicellular 
hairs 0.7-1.5 mm long. Leaves sessile and decus- 
sate, opposing leaves not united at the base, often 
subtended by a tuft of hairs; leaf blades 6-20 mm 
long, 5-18 mm wide, ovate to broadly ovate-el- 
liptic or ovate-orbicular, apex rounded or bluntly 
obtuse, sometimes slightly notched, margin entire, 
base rounded and truncate to subcordate or auric- 
ulate, drying greenish to yellowish, glabrous or 
hirsute near the base beneath, venation palmate 
with 3-7 poorly defined primary veins. Inflores- 
cences of solitary axillary flowers, 1-2/node, ped- 
icels 8-23 mm long, 0.2-0.4 mm diam., hirsutu- 
lous with slender hairs 0.2-0.5 mm long, ebrac- 
teolate (but the outer sepals resembling bracts). 
Flowers with strongly unequal sepals, outer 3 se- 
pals 4-6 mm long, 2.5-5 mm wide, ovate with 
truncated or subcordate bases and resembling the 
leaves, ciliolate along the margin, inner sepals 



BURGER: FLORA COSTARICENSIS 



23 



narrowly triangular; corolla 6-9 mm long, pale 
blue to lavender or violet (white), slightly exsert- 
ed beyond the sepals, 4-lobed with the upper lobe 
emerginate; stamens 4, included, filaments 1-2 
mm long, glabrous, larger anthers 1.2-1.5 mm 
long; ovary ca. 1 mm long, narrowly ovoid, gla- 
brous, style 2-5 mm long, stigma 0.3 mm wide. 
Fruits ca. 3 mm long and 1 mm wide, narrowly 
ovoid-oblong; seeds many, small, reddish brown, 
reticulate. 

Plants of shallow water and moist open sunny 
sites at the edges of water in both evergreen and 
deciduous forest areas, 40-1700 m elevation. This 
is a common species of marshes, swamps, and wet 
depressions, growing both as a partly submerged 
aquatic, prostrate and mat-forming on wet mud, 
or erect among other wetland plants, flowering 
throughout the year. This species ranges from 
Mexico and the West Indies to southern Brazil. 

Bacopa salzmannii is the most commonly col- 
lected species of Bacopa in Costa Rica. It is rec- 
ognized by its often aquatic or repent habit in wet 
sites, the villous stems, sessile rounded leaves, 
and leaf-like (bract-like) outer sepals. The round- 
ed bases of the leaf blades may obscure the stem 
but they are not joined (interpetiolar, cf. B. re- 
pens) nor are they decurrent on the stem. 

Bacopa sessiliflora (Benth.) Pulle, Enum. PI. Su- 
rinam 415. 1906. Herpestis sessiliflora Benth. 
i/i J. D. Hook., Companion Bot. Mag. 2: 58. 
1836. B. sessiliflora (Benth.) Edwall, Bol. Com- 
miss. Geogr. Estado Sao Paulo 13: 176, 181. 
1897, incomplete combination fide D'Arcy 
1979. Caconapea conferta Pennell, Proc. Acad. 
Nat. Sci. Philadelphia 72: 153. 1920. C. sessi- 
liflora (Benth.) Pennell, loc. cit. 75: 11. 1923. 

Erect herbs to 40 cm tall, with short lateral 
branches, rooting only at the base, leafy stems 
0.8-3 mm diam., glabrous, with 2 opposite lon- 
gitudinal ridges. Leaves sessile, separate at the 
base and not forming an interpetiolar line, decur- 
rent on the stem, distal leaves smaller (5 mm) and 
bract-like; leaf blades 10-35 mm long, 2-5 mm 
wide, oblanceolate to linear-oblanceolate or nar- 
rowly elliptic-oblanceolate, apex acute, distal half 
of the margin with prominent teeth ca. 0.5 mm 
high, 1-2 mm long, cuneate to the base, drying 
yellowish brown, glabrous, punctate on both sur- 
faces, venation pinnate. Inflorescences fasciculate 
in distal spiciform arrangements or 1-4 flowers in 
leaf axils, distal spikes 1-3 cm long, ca. 1 cm 
diam., flowers sessile or subsessile on pedicels 
less than 1.5 mm long, bracteoles 1-2 mm long 



at apex of pedicel or absent. Flowers glabrous 
externally, calyx united at base, outer calyx lobes 
3-3.5 mm long, 1.3-2 mm wide, ovate, thin with 
3 prominent parallel veins; corolla 3-4 mm long, 
tubular-campanulate, exserted, bluish or white, 
lobes ca. 0.5 mm long; stamens 4, attached near 
middle of tube; ovary ca. 1 .5 mm long, style ca. 
1 mm long. Fruits 3-4 mm long, 1.5-2 mm 
diam., narrowly ovoid, surface with minute (0.1 
mm) capitate hairs; seeds ca. 0.3 X 0.2 mm, rect- 
angular with truncated ends, brown, longitudinal- 
ly reticulate/ridged. 

Rarely collected plants of open sunny sites in 
wet marshes and depressions. The plants are tol- 
erant of brackish water and can be found near the 
seashore. The species has not been collected in 
Costa Rica but is known from Bluefields, Nica- 
ragua, and is found in central Panama. The spe- 
cies ranges from Guatemala and Belize to Ecua- 
dor and the West Indies. 

Bacopa sessiliflora is distinguished by its erect 
habit, general lack of pubescence, sessile oblan- 
ceolate leaves serrulate along their distal margins, 
and sessile flowers often in axillary fascicles that 
may be arranged in terminal spike-like inflores- 
cences. Compare B. monnierioides. 



Benjaminia Martius 

REFERENCE L. B. Smith & J. M. Pires, An 
evaluation of Benjaminia Martius ex Benjamin. J. 
Wash. Acad. Sci. 46: 86. 1956. 

Herbs, aquatic and submerged, rooting at lower 
nodes, glabrous or puberulent, glandular punctate 
on vegetative parts. Leaves verticillate, united at 
the base, petiolate, the blades pinnatifid with slen- 
der linear segments. Inflorescences of solitary ax- 
illary flowers, bracts absent, pedicels elongating 
slightly in fruit, bracteoles absent at the base of 
the calyx. Flowers with calyx lobes united near 
the base, calyx lobes (sepals) 5 and subequal, nar- 
row, valvate in bud; corolla tubular and 2-lipped, 
upper lip slightly 2-lipped, lower lip 3-lobed; sta- 
mens 4, of 2 unequal pairs, included, anthers 
equal, thecae similar; ovary subtended by a ring 
of filaments, 2-locular, ovules many, style simple, 
stigma flat and slightly curved. Fruits thin-walled 
capsules, loculicidal, surface smooth; seeds many, 
oblong, surface longitudinally reticulate. 

A monotypic genus of unusual aquatic plants, 
this taxon is closely related to Bacopa but differs 
in having nearly equal sepals, unusual pinnatifid 
leaves, and ovary subtended by staminodial fila- 



24 



FIELDIANA: BOTANY 



ments (present in some Bacopa species). This ge- 
nus has been confused with the Old World Lim- 
nophila, but Benjaminia is distinct because of its 
estipitate anther-thecae and its two-lobed stigma. 
Also, Limnophila species usually have emergent 
leaves that are broad and dentate, whereas emer- 
gent leaves are never found in Benjaminia. For a 
short discussion of the complex nomenclature of 
the genus, see D'Arcy (1979, p. 194). 

Benjaminia reflexa (Benth.) D'Arcy, Ann. Mis- 
souri Bot. Card. 66: 194. 1979. Herpestis re- 
flexa Benth. in DC., Prodr. 10: 399. 1846. Be. 
utriculariaeformis Mart., Fl. Bras. 10: 256. 
1847. Quinquelobulus utriculariaeoides Benj., 
Linnaea 20: 316. 1847. Monnierea reflexa 
(Benth.) Kuntze, Rev. Gen. PI. 2: 463. 1891. 
Bacopa reflexa (Benth.) Edwall, Bol. Commiss. 
Geogr. Estado Sao Paulo 13: 176. 1897. Naia- 
dothrix longipes Pennell, Mem. Torrey Bot. 
Club 16: 105. 1920. Bacopa naias Standl., Field 
Mus. Bot. Ser. 11: 141. 1932. Limnophila cos- 
taricensis Suesseng., Bot. Jahrb. Syst. 72: 284. 
1942. L. costaricensis forma aquatica Sues- 
seng., loc. cit. L. costaricensis forma semiter- 
restris Suesseng., loc. cit. Figure 1. 

Submerged herbs, only the flowers and fruits 
extending above the water surface, branched, 
rooting at lower nodes, internodes 2-60 mm long, 
0.5-2 mm diam., glabrous or minutely and sparse- 
ly puberulent, vegetative parts glandular punctate. 
Leaves in verticels of 6 or 8/node (rarely 2 or 4), 
glabrous or sparsely puberulent, petioles 2-6 mm 
long (to the first pinna), 0.2-0.4 mm wide; leaf 
blades 4-35 mm long, 5-25 mm wide, with slen- 
der filiform pinnate lobes, lobes 4-15/side and in 
a single plane, central rachis 0.2-0.4 mm wide, 
lobes 0.05-0.15 mm wide, often with small (0.1- 
0.2 mm) punctate glands. Inflorescences of soli- 
tary axillary flowers, 1 flower/node, pedicels 5- 
9(-18) mm long, 0.2-0.4 mm diam., glabrous or 
sparsely and minutely puberulent. Flowers with 
sepals 2-4 mm long, 0.4-0.7 mm wide, linear- 
lanceolate to linear-oblong, apex usually slightly 
rounded; corolla 4-6 mm long, bluish white or 
purple, yellowish within the throat; anthers ver- 
satile and similar; ovary ca. 2 X 1 mm, narrowly 
ovoid, style ca. 1 mm long, slender. Fruits 2-3 
mm long, narrowly ovoid, smooth, with a persist- 
ing style ca. 1 mm long; seeds ca. 0.6 mm long, 
oblong-fusiform, reticulate. 

Submerged aquatic plants of shallow ponds and 
lakes, 0-800 m elevation. Only three collections 
from Costa Rica have been seen, all from the 



General Valley (near. Buenos Aires and between 
San Isidro and Rivas), flowering and fruiting in 
November (Crow 6176 & 6239), December (Nic- 
aragua), and February (Kupper 597). The species 
(in a wide sense) ranges from Mexico and Cuba 
to Brazil. 

Benjaminia reflexa is an unusual aquatic plant 
distinguished by being almost entirely submerged, 
having whorls of feather-like leaves with slender 
pinnate lobes, solitary little flowers with five near- 
ly equal narrow sepals, and sympetalous two 
lipped corollas. The flowers and fruits are borne 
above the water surface on stiff pedicels. The 
leaves can also be interpreted as being opposite, 
with each leaf having three or four primary axes 
and each of these having deeply divided pinnatifid 
divisions. These plants resemble the submerged 
portions of species of Cabomba (Nymphaeaceae) 
as well as aquatic Utricularia species (Lentibu- 
lariaceae). The only similar Neotropical species of 
Scrophulariaceae is Bacopa myriophylloides 
(Benth.) Pennell, which has palmately dissected 
leaves and differing flowers. This species was 
treated as Bacopa naias in the Flora of Guate- 
mala. Suessenguth's Limnophila costaricensis 
lacks a species description but has Latin descrip- 
tions of two forms, all based on a single collection 
number (Kupper 597, fragments at F). 



Buchnera Linnaeus 

REFERENCE D. Philcox, Revision of the New 
World species of Buchnera L. Kew Bull. 18: 275- 
316. 1965. 

Herbs, annual (in ours) or perennial, hemipar- 
asitic, erect, simple or branched, hairs with a 
broad base and stiff straight tip, often scabrous, 
drying dark. Leaves opposite or subopposite, 
sometimes alternate distally, sessile to short-peti- 
olate, leaf blades narrow, dentate or entire, mar- 
gins often revolute, usually scabrous, venation 
pinnate or palmate, with 1 or 3 (5) major veins. 
Inflorescences terminal compact or lax spikes, 
flowers sessile or subsessile, subtended by a bract, 
with 2 slender lateral bracteoles beneath the calyx. 
Flowers with a tubular calyx, slightly enlarging 
in fruit, with 5 or 10 prominent longitudinal veins, 
lobes 4 or 5, shorter than the tube, acute; corolla 
radially symmetric and usually salverform, tube 
longer than the calyx, cylindric and straight or 
slightly curved, blue to white or purple, glabrous 
(in ours) or puberulent externally, lobes 5 and 



BURGER: FLORA COSTARICENSIS 



25 



subequal, rotate-spreading, shorter than the tube, 
apex of the tube often with moniliform hairs; sta- 
mens 4, of 2 unequal pairs, inserted in the prox- 
imal hall of the tube, subsessile or with short fil- 
aments, anthers ovoid, versatile, 1-thecous; ovary 
ellipsoid to ovoid, 2-locular with many ovules, 
style slender and included, stigma clavate. Fruits 
dry capsules partly enclosed by the persisting ca- 
lyx, splitting loculicidal into 2 equal parts; seeds 
many, oblong or ellipsoid, curved or angled, lon- 
gitudinally reticulate. 

A genus of ca. 100 species in tropical and tem- 



perate climates throughout the world. Most of the 
species are found in the Old World tropics; there 
are about 1 6 species in the New World. The genus 
is distinguished by its slender few-branched habit, 
stiff, narrow, usually scabrous leaves, spicate in- 
florescences, tubular calyx, and one-thecous an- 
thers, all parts drying dark or black. All species 
are believed to be hemiparasites, attaching by 
haustoria to the roots of the host plants. The genus 
is placed in the tribe Buchnerae, along with many 
Old World genera. Its closest Neotropical relatives 
are Alectra and Escobedia. 



Key to the Species of Buchnera 

la. Stems hispid-scabrid, rough to the touch; larger leaves with prominent teeth; calyx pubescent, be- 
coming slightly thickened in fruit, but the areas between the longitudinal veins usually remaining 
flat and the minor venation not visible B. pusilla 

Ib. Stems usually smooth to the touch at distal internodes; larger distal leaves entire and linear; calyx 
glabrous, becoming conspicuously thickened and with raised minor venation between the longitu- 
dinal veins in fruiting stages B. weberbaueri 



Buchnera pusilla Kunth in H.B.K., Nov. Gen. 
Sp. 2: ed. quarto 340. 1818. B. tinctoria Bertol., 
Fl. Guatemala 26. 1840. B. major Polak., Lin- 
naea 41: 588. 1877. B. mexicana Hemsl., Biol. 
Centr. Am. Bot. 2: 457. 1881. Figure 4. 

Stiff erect herbs (6-)10-70(-150) cm tall, an- 
nual, unbranched or with 2-7 distal branches, dry- 
ing dark, stems 0.4-3 mm diam., terete, hispid 
with stiff whitish hairs 0.3-1.3 mm long. Leaves 
opposite or subopposite, sessile or with poorly 
differentiated petioles to 8 mm long; leaf blades 
15-50 mm long, 1-8 mm wide, linear to linear- 
oblong or linear-oblanceolate (oblong near the 
base), apex acute, margin with 1-5 prominent 
teeth (0.2-1 mm) or entire, gradually narrowed to 
the cuneate base, subcoriaceous, with scattered 
short scabrid hairs above, larger (0.2-0.5 mm) 
stiff hairs along the margin and veins beneath, 
with 1 or 3 major veins. Inflorescences 3-15 cm 
long, bracts 4-9 mm long, ovate to lanceolate, 
pedicels 0-1 mm long, with 2 shorter linear brac- 
teoles beneath the calyx, bracts and bracteoles 
scabrid and ciliate. Flowers with calyx 4-5 mm 
long, enlarging (to 8 mm) and becoming more 
scabrid in fruit, with 10 longitudinal veins, hispid 
only along the veins, lobes 1-2.5 mm long, equal 
or unequal, acute; corolla 8-14 mm long, salver- 
form, white to purplish white or lilac, tube 0.4- 
0.8 mm diam., lobes 1-5 mm long, obovate with 
narrowed base and broadly rounded apex; stamens 



inserted near the middle of the tube. Fruits 4-6 
mm long, ca. 2 mm wide, oblong, smooth, partly 
enclosed in the scabrid calyx; seeds 0.4-0.5 mm 
long, oblong or curved. 

Plants of open sunny sites in fields and savan- 
nas in deciduous and evergreen forest areas of the 
Pacific slope and central highlands, 10-1700 m 
elevation. Flowering and fruiting from October to 
early March. The species ranges from Mexico to 
Ecuador and Brazil. 

Buchnera pusilla is distinguished by its short, 
stiff, erect (usually few-branched) habit, hispid to 
scabrous vesture, spicate inflorescences, and sal- 
verform white to pinkish corollas, all parts turning 
dark when dried. This is a frequently collected 
species in Central America. We have included 
Herrera 801 within our concept of this species, 
which has been identified as B. longifolia Kunth 
by Wilcox (1965, see above). Buchnera pusilla 
and B. longifolia form a variable species complex, 
widely distributed in the lowland Neotropics. 
They are distinguished by the amount and type of 
pubescence on the calyx, but that character can 
vary with the age of the flower, the fruits becom- 
ing more scabrid as they mature. See the illustra- 
tion in Flora of Guatemala (Standley & Williams, 
1973, p. 337). 

Buchnera weberbaueri Diels, Bot. Jahrb. Syst. 
37: 430. 1906. B. leiantha Standl., Publ. Field 
Mus. Nat. Hist., Bot. Sen 22: 105. 1940. 



26 



FIELDIANA: BOTANY 



Stiff erect herbs 30-70 cm tall, annual, usually 
without lateral branches, leafy stems 1-2.5 mm 
diam., glabrous or with minute (0.1-0.2 mm) sca- 
brid hairs in longitudinal files or at lower inter- 
nodes, usually smooth to the touch. Leaves op- 
posite or subopposite, sessile and slightly clasping 
the stem, a petiole not differentiated; leaf blades 
2-7 cm long, 0.5-4 mm wide, linear to linear- 
lanceolate, apex acute, margins entire (except 
rarely on basal leaves), base gradually narrowed, 
drying black or brown, glabrous, with 1 central 
vein or tripliveined. Inflorescences 4-14 cm long, 
flowers separate or closely congested into a dense 
spike, abaxial bracts 2-3 mm long, acute, enlarg- 
ing to 5 X 3 mm in fruit, ciliolate along the mar- 
gin, pedicels 0-1 mm long. Flowers with calyx 
5-7 mm long, to 10 mm in fruit, sparsely puber- 
ulent with few white hairs, sepal lobes ca. 1 .5 mm 
long, acute, minor venation between the main 
veins not apparent until fruiting; corolla 7-13 mm 
long, salverform, lilac to pink-purple or white, 
tube ca. 0.6 mm diam., lobes 1.5-2 mm long, 
equal. Fruits 5-7 mm long, ca. 3 mm diam., sur- 
face smooth and glabrous, included within the 
stiff thickened calyx. 

Rarely collected plants of open sunny sites, 
200-1300 m elevation. In Costa Rica this species 
has only been collected in Guanacaste Province, 
flowering in January and September-October, 
with old fruit in February. The species has been 
collected in Belize, in Izabal (Guatemala), and on 
the Cerro de Espiritu Santo near Naranjo in Costa 
Rica; it ranges southward to Trinidad and Peru. 

Buchnera weberbaueri is recognized by its 
short, usually unbranched stems, very sparse pu- 
bescence, linear entire leaves, and sepals that be- 
come indurated and with many elevated veins in 
fruiting stages. Two rather different specimens are 
provisionally placed here: Herrera 801, with 
young flowers, and Williams & Williams 24514, 
with old fruits. Standley and Williams (1973) con- 
fused B. weberbaueri with the larger-flowered B. 
palustris (Aubl.) Spreng., which is found in low- 
land South America in wet areas. In addition to 
its larger flowers, B. palustris has longer pedicels 
and narrowly lanceolate bracts and bracteoles. 



Calceolaria Linnaeus 

RKFERENCES U. Molau, Scrophulariaceae, Part 
1. Calceolarieae. Fl. Neotropica, Monogr. 47: 1- 
326. 1988. L. R. Landrum & R. McVaugh, Cal- 
ceolaria mexicana and C. tripartita in Mexico. 
Contrib. Univ. Michigan Herb. 11: 273-309. 1978. 



Shrubs, vines or herbs, annual or perennial, 
usually confined to high-elevation habitats, stems 
terete, hairs simple. Leaves opposite (ternate in 
some Andean species), sessile or petiolate, leaf 
blades simple and without prominent lobes or be- 
coming deeply lobed or pinnatifid. margins usu- 
ally serrate, pinnatcly veined. Inflorescences ter- 
minal or axillary, often a compound thyrse of 
cymes with 3 or 4 flowers or of 1-3 axillary flow- 
ers, often subtended by smaller leaves, bracteoles 
absent at the base of the calyx. Flowers small to 
large, calyx deeply 4-parted. sepal lobes equal or 
unequal, valvate in bud, usually persisting in fruit; 
corolla strongly bilabiate with the lower lip usu- 
ally developed into a globose or slipper-like sac- 
cate form, the upper lip usually much smaller and 
arched or hooded and enclosing style and sta- 
mens, yellow to red or purple, sometimes spotted 
or mottled, glabrous on the exterior; stamens 2, 
attached near the base of the tube, filaments short, 
anther with 2 contiguous thecae opening by a lon- 
gitudinal slit, staminodes absent; ovary superior 
and conical (partly inferior in the temperate sub- 
genera Cheiloncos and Rosula), somewhat 2- 
lobed. 2-locular, ovules many, style short and of- 
ten recurved, stigma simple or capitate. Fruits 
dry capsules, septicidal and loculicidal, the 4 
valves opening from the apex; seeds many, small 
(0.3-1 mm), usually with longitudinal and trans- 
verse ridges or minute tubercles. 

A pantropical genus of ca. 300 species with a 
majority of the species found above 1000 m ele- 
vation in the Andes. A number of species range 
into the lowlands of temperate Chile and Argen- 
tina. The genus is classified with Jovellana and 
Porodittia in the tribe Calceolarieae. This tribe 
exhibits links between southern South America 
and New Zealand, duplicating the disjunction 
found in the tribe Veronicae. Molau (1988) pro- 
vided a fine treatment of this complex and fasci- 
nating group in the Neotropics; his discussions of 
biology, distribution, speciation, and phylogeny 
are especially noteworthy. Calceolaria is imme- 
diately recognizable because of its bright yellow 
bilabiate corollas with small upper lip and round- 
ed saccate or inflated slipper-like lower lip. The 
four-parted calyx, only two stamens, and restric- 
tion to moist high-elevation habitat (in Central 
America) are further distinctions. Some South 
American species have corollas that are somewhat 
S-shaped in lateral view. A number of hybrids are 
popular as potted ornamentals, called "slipper- 
worts" or "slipper flowers." 



BURGER: FLORA COSTARICENSIS 



27 



Key to the Species of Calceolaria 

la. Petioles broadly winged to the base and united across the node, the leaves perfoliate [larger blades 

triangular to sagittate, never pinnatifid: anther cells contiguous, dark brown; uncommon] 

C. perfoliata 

Ib. Petioles not broadly winged to the base, the leaves not perfoliate 2 

2a. Larger leaves lanceolate, without pinnate lobes or deep sinuses; anther-thecae contiguous, not sep- 
arated by an expanded connective; stems slightly woody and plants clambering to 2 m high (note 

that these 2 species are very similar) 3 

2b. Larger leaves usually with prominent pinnate lobes separated by deep sinuses; anther-thecae sepa- 
rated by the slender expanded connective; stems mostly herbaceous and semisucculent, plants to 1 

m high 4 

3a. Fruits puberulent; 2 veins of leaves not loop-connected in the distal half of the blade, usually 
strongly ascending, upper surface of the blade with very short hairs and glands, lower surface 
often with brown punctations; stems usually puberulent; seeds with prominent longitudinal 

ridges and smaller transverse ribs; common high-montane plants C. irazuensis 

3b. Fruits glabrous; 2 veins of leaves usually loop-connected in the distal half of the blade, upper 
surface lacking very short hairs or glands, lower surface not punctate; stems usually glabrous 
(rarely with a few hairs at nodes and petioles); seeds with prominent longitudinal ridges but no 

transverse ribs; rarely collected in Costa Rica C. microbe/aria 

4a. Corolla 5-15 mm long, mouth open; fruits 3-7 mm long, oblate; seeds 0.4-0.5 mm long; sinuses 

of larger leaves rarely reaching the midvein; lower anther-theca fertile C. mexicana 

4b. Corolla 10-20 mm long, mouth closed; fruits 6-9 mm long, ovoid; seeds 0.6-0.8 mm long; sinuses 
of larger leaves sometimes reaching the midvein; lower anther-theca sterile C. tripartita 



Calceolaria irazuensis J. D. Smith, Bot. Gaz. 20: 
292. 1895. C. costaricensis Kranzl., Ann. K. K. 
Naturhist. Hofmus. 22: 192. 1907. Figure 3. 

Subshrubs, erect or scandent, 0.2-1 (-2) m tall, 
leafy stems 1.2-5 mm diam., densely puberulent 
with slightly viscous hairs 0.05-0.3 mm long, old- 
er stems glabrescent. Leaves forming a line across 
the stem at their base, petioles 2-8(-15) mm long, 
0.5-1.8 mm diam.. puberulent with longer hairs 
adaxially; leaf blades 1.8-9(-12) cm long, 6-35(- 
45) mm wide, lanceolate to ovate-lanceolate or 
elliptic-lanceolate, gradually narrowed to the 
acute apex, margin serrate with teeth 0.3-1 mm 
high, 2-5 teeth/cm, base acute to obtuse or slight- 
ly rounded, asymmetric, drying much paler be- 
neath than above, minutely (0.1-0.2 mm) puber- 
ulent above, glabrous between the veins beneath, 
2 veins 4-9/side and strongly ascending. Inflo- 
rescences mostly of 2-4 long-pedunculate cymes 
subtended by a pair of smaller leaves (bracts) or 
with 2-4 flowers terminal on the distal leafy node, 
minutely viscid puberulent, peduncles 1.5-12 cm 
long, 1-2 mm diam., usually bearing 3 or 4 ped- 
icellate flowers, pedicels 1-4 cm long, 0.4-0.6 
mm diam. Flowers with calyx 6-9 mm long, ca- 
lyx lobes 4-9 mm long, broadly ovate-triangular, 
minutely puberulent along the margin; corolla 



12-24 mm long, minutely glandular-papillose on 
the exterior, upper lip 6-8 mm long, lower lip 1 5- 
25 mm long, 12-18 mm wide, bright yellow; fil- 
aments ca. 1.5 mm long, anthers with divaricate 
thecae to 3.5 mm wide; style 2-3 mm long, 
curved. Fruits 6-9 mm long, 5-8 mm wide, ovate 
with truncated base, minutely papillate-puberu- 
lent; seeds 0.4-0.7 mm long, 0.2-0.3 mm wide, 
oblong-ellipsoid, dark brown, with prominent lon- 
gitudinal ridges and minute transverse ribs (X50). 

Plants of evergreen high-montane forest for- 
mations, (1800-)2400-3500 m elevation. Flow- 
ering and fruiting throughout the year but with 
most collections made between November and 
May. The species ranges from Volcan Barva 
southward to the Chiriqui highlands of Panama. 

Calceolaria irazuensis is recognized by it most- 
ly lanceolate leaves, slightly viscid hairs, inflores- 
cences often with long peduncles and/or pedicels, 
and slipper-shaped yellow flowers. This species is 
frequently encountered in open sites at higher el- 
evations. Compare the closely similar but rarely 
collected C. microbe/aria. Common names are 
baton de oro and gallitos. 

Calceolaria mexicana Benth., PI. Hartw. 47. 
1840. C. trachelifolia Martens & Galeotii, 
Acad. R. Sci. Bruxelles 12: 16. 1845. C. urti- 



28 



FIELDIANA: BOTANY 



cina Kranzl., Feddes Repert. Spec. Nov. Regni 
Veg. 1: 82. 1905. Figure 3. 

Annual herbs, erect to decumbent, 10-60(- 
100) cm tall, leafy stems 0.6-3 mm diam., slightly 
succulent, often reddish in color, sparsely to 
densely puberulent with simple or viscid gland- 
tipped hairs 0.3-0.6 mm long. Leaves slightly 
clasping the stem and forming an interpetiolar 
ridge, petioles 2-40(-160) mm long, 0.4-1.4 mm 
diam., pubescence similar to that of the stem; leaf 
blades 1-12 cm long, 0.4-8(-13) cm wide, vary- 
ing from narrowly ovate-triangular in smaller 
blades to broadly ovate-triangular in outline with 
2-4 prominent pinnatifid lobes separated by deep 
sinuses in larger blades, apex acute, margins 
strongly dentate-serrate with intermixed larger (2- 
4 mm) and smaller (0.3-2 mm) teeth, base obtuse 
to subcordate, drying yellowish green to grayish 
green, upper surface with slender hairs 0.2-0.4 
mm long, lower surface with few thin hairs to 1 
mm long on the major veins, 2 veins 6-12/side. 
Inflorescences of solitary axillary flowers (21 
node) or resembling terminal cymes when sub- 
tended by reduced bract-like distal leaves, pedi- 
cels 4-14 mm long, elongating in fruit, 0.15-0.25 
mm diam., minutely puberulent with gland-tipped 
hairs. Flowers with calyx 4-6 mm long, calyx 
lobes 2-5 mm long, 1-3 mm wide at the base, 
acute, with glandular hairs at the base; corolla 5- 
15 mm long, 3-10 mm wide, the upper lip 1-3 
mm long, lower lip 5-14 mm long, 3-6 mm wide, 
slipper-shaped, yellow; anthers 2-3.5 mm wide, 
the 2 small thecae fertile, separated by the longer 
narrow connective; ovary 2.5 mm diam., papil- 
late-puberulent. Fruits 3-7 mm long, 3-6 mm 
wide, oblate or globose, thin-walled; seeds 0.4- 
0.5 mm long, ca. 0.3 mm thick, oblong, dark. 

Plants of wet sites near streams, moist depres- 
sions, and wet cliffsides, 1500-3200 m elevation. 
Flowering throughout the year, but with most 
Costa Rican collections made in April-August. 
This species is the commonest Calceolaria in 
Mexico and Central America; it ranges to Bolivia. 

Calceolaria mexicana is recognized by its pref- 
erence for wet montane habitats, diverse foliage 
with the large leaves having prominent pinnate 
lobes, small slipper-shaped yellow corollas, and 
stamens with two small but functional thecae. 
This species can be found growing together with 
the very similar C. tripartita, and the two are eas- 
ily confused. Both species have a strongly devel- 
oped connective separating the thecae. In C. mex- 
icana the corollas are open and the lower lip is 



slightly three-lobed. In addition, the larger leaves 
do not have the deep sinuses of C. tripartita 
leaves. 

Calceolaria microbefaria Kranzl., Ann. K K. 
Natur. Hofmus. Wien 22: 193. 1907. C. storkii 
Standl., Publ. Field Mus. Nat. Hist. Bot. Ser. 18: 
1103. 1938. 

Shrubs or subshrubs 0.3-3 m tall, erect or 
clambering, much branched, stems glabrous or 
with a few hairs at the nodes. Leaves with peti- 
oles 3-1 1 mm long, glabrous or with a few 'hairs 
along the adaxial margins; leaf blades 4-9(-13) 
cm long. l-2.3(-2.8) cm wide, lanceolate to nar- 
rowly oblong-lanceolate, apex acute, margin ser- 
rate with 5-7 gland-tipped mucronulatc teeth/cm, 
base acute, upper surface glabrous or minutely pa- 
pillate, lower surface often drying pale grayish or 
reddish, 2 veins 5-1 I/side, loop-connected dis- 
tally. Inflorescences 3-12 cm long, usually with 
2 or 3 pairs of 4-8 flowered cymes, peduncles 1- 
5 cm long, bracts often lacking at the base of the 
cymes, pedicels 0.5-3.4 cm long, glabrous to mi- 
nutely papillate-puberulent (to tomentose in South 
America). Flowers with sepals 3.3-6 mm long, 
2.8-4 mm wide at anthesis, minutely puberulent 
along the margins; corolla 10-20 mm long, bright 
yellow, the upper lip hooded or flattened and sub- 
circular, the lower lip projecting or pendant, sac- 
cate; anthers 2-3.7 mm wide. Fruits 5-9 mm 
long, 5-9 mm wide, broadly ovoid with truncated 
base, glabrous; seeds 0.7-0.9 mm long, ca. 0.3 
mm thick, with prominent longitudinal ridges, 
surface smooth. 

Plants of subparamo formations near the high- 
est point along the Interamerican Highway in the 
western part of the Talamanca range. 3100-3500 
m elevation. Flowering in March-August; fruiting 
in August. This species is known from only four 
collections in Costa Rica (Barringer el al. 2913, 
Grayum & Affolter 8190, Skutch 5188, and Stork 
3048, type of C. storkii). The species ranges from 
Venezuela to central Ecuador in the Andes, with 
a small disjunct population in Costa Rica. 

Calceolaria microbefaria is recognized by its 
lanceolate leaves, generally glabrous vegetative 
parts and fruits, bright yellow slipper-shaped co- 
rollas, and very limited range in our area. Molau 
(1988) divides this species into three geographi- 
cally and morphologically distinct subspecies, 
with the Costa Rican collections placed in subsp. 
microbefaria. In Costa Rica, this species is easily 
mistaken for the much more common C. irazuen- 



BURGER: FLORA COSTARICENSIS 



29 



sis; the characteristics used in the key to separate 
the two species usually allow confident identifi- 
cation. 

Calceolaria perfoliata L.f., Suppl. 86. 1781. Fa- 
gelia perfoliata (L.f.) Kuntze, Rev. Gen. PI. 2: 
460. 1891. C. sciadephora J. D. Smith, Bot. 
Gaz. 25: 151. 1898. Figure 6. 

Herbs or weak-stemmed subshrubs 0.5-1.5 m 
high (to 5 m long), often scandent on other plants, 
leafy stems 1.5-6 mm diam., often with 4 prom- 
inent longitudinal ridges, slightly succulent, with 
slender crooked multicellular hairs 0.3-1 mm 
long. Leaves sessile and perfoliate, opposing 
leaves united at the base with lateral tissue to 6 
mm wide, leaves subtending the inflorescences 
not perfoliate, petioles broadly winged and blade 
like, 0.5-6 cm long, 3-12 mm wide and expanded 
near the stem; leaf blades 2-11 cm long (beyond 
the blade-like petiole), 2-7 cm wide, narrowly tri- 
angular or sagittate, apex acute, margins serrate- 
dentate with larger (2-4 mm) and smaller (1-2 
mm) teeth intermixed, base truncated to the 
winged petiole-like part, drying yellowish brown, 
often silvery below (in life), upper surface with 
short (0.2 mm) scattered hairs, lower surface with 
thin hairs to 1 mm long, 2 veins 5-7/side. Inflo- 
rescences with 8-12 (4-24) flowers, borne on ax- 
illary peduncles 6-15 cm long, 1-1.5 mm diam., 
terminated by a pair of asymmetric leaves/bracts 
2-5 cm long, pedicels 1 .5-5 cm long, densely pu- 
berulent. Flowers with calyx 8-12 mm long, 
lobes 6-8 mm long, puberulent, broadly ovate, 
rounded at base, apex obtuse, enlarging slightly 
in fruit; corolla to 22 mm long, bright yellow, 
upper lip 4-9 mm long, lower lip 1 5-20 mm long, 
8-18 mm wide, anthers U- or C-shaped, 2-4 mm 
wide; ovary pubescent, style 3-5 mm long. Fruits 
5-7 mm long, ca. 5 mm diam., ovoid with trun- 
cated base, pubescent; seeds 0.5-0.7 mm long, 
0.1-0.2 mm wide. 

Infrequently collected plants of evergreen mon- 
tane forest formations, 2700-3300 m elevation (as 
low as 1800 m in Chiriqui). Flowering and fruit- 
ing appear to be restricted to January-February in 
Costa Rica. This species ranges from the Cordil- 
lera de Talamanca and the Chiriqui highlands to 
southern Ecuador. 

Calceolaria perfoliata is distinguished by its 
perfoliate leaves with broadly winged petioles and 
distal narrowly triangular or sagittate blades. The 
anthers are attached at the center and decurved 
laterally to produce a rounded C-shaped or horse- 



shoe-shaped form. The restricted flowering period 
may account for the relatively few Costa Rican 
collections. A related species, C. trilobata Hem- 
sley, is native to Mexico-Guatemala and Vene- 
zuela-Bolivia but is not known from southern 
Central America. That species has a more defi- 
nitely contracted petiole between the distal blade 
and the expanded perfoliate base, and the anthers 
are not horseshoe-shaped. 

Calceolaria tripartita R. & P., Fl. Peruv. Prodr. 
1: 14, tab. 22. 1798. C. heterophylla Wild., 
Enum. PI. 1: 29. 1809 (non C. heterophylla R. 
& P.). Figure 3. 

Herbs, erect or decumbent, 9-90 cm tall, stems 
1-4 mm diam., slightly succulent, puberulent with 
multicellular, often gland-tipped hairs 0.2-0.7 mm 
long. Leaves simple and serrate to deeply pinnat- 
ifid, slightly clasping the stem and forming a ridge 
across the node, petioles 1-5 cm long, 0.5-1.5 
mm wide, glandular puberulent; leaf blades 1-16 
cm long, 0.3-1 1 cm wide, ovate-lanceolate in 
smaller leaves to broadly ovate-triangular, larger 
blades with deep proximal sinuses separating 1-3 
pairs of subopposite lobes, apex acute, margin 
with larger (3-10 mm) and smaller (0.3-3 mm) 
teeth, base truncated in larger leaves, drying 
greenish or brown, upper surface with transparent 
multicellular hairs to 1 mm long. Inflorescences 
of terminal or axillary flowers, 2-4/node (if small 
distal leaves are interpreted as bracts, the sub- 
tending internodes can be interpreted as peduncles 
of thyrse-like inflorescences), pedicels 4-45 mm 
long, 0.2-0.4 mm diam., puberulent with simple 
or glandular hairs 0.2-0.4 mm long. Flowers with 
calyx 5-7 mm long, lobes ca. 4 mm long, with 
small glandular hairs; corolla 10-20 mm long, 
bright yellow, upper lip 2-5 mm long, lower lip 
10-25 mm long, 6-15 mm wide, slipper-shaped, 
unlobed; anthers 2-5 mm long, one theca sterile, 
reduced to a nob, connective slender. Fruits 6-9 
mm long, ovoid, rounded at the apex; seeds 0.6- 
0.8 mm long, ca. 0.4 mm diam., reddish brown, 
with longitudinal ridges. 

Plants of wet or moist open sites in evergreen 
montane forest formations, 1000-3200 m eleva- 
tion. Flowering throughout the year, but with most 
collections made between October and December. 
This species ranges from Mexico to Peru. 

Calceolaria tripartita is recognized by it high- 
land habitats, glandular viscid pubescence, larger 
leaves deeply pinnatisect (almost pinnately com- 
pound), yellow slipper-shaped corollas, and un- 



30 



FIELDIANA: BOTANY 



usual stamens. This species is similar to C. mex- 
icana (q.v.), and the two have often been con- 
fused, as in the Flora of Guatemala. In addition 
to the differences used in the key, the pedicels of 
C. tripartita usually have more conspicuous 
gland-tipped hairs. 



Capraria Linnaeus 

REFERENCES T. A. Sprague, A revision of the 
genus Capraria. Kew Bull. 1921: 205-212. 1921. 
C. Niezgoda & S. Tomb, Systematic palynology 
of the tribe Leucophylleae (Scrophulariaceae) and 
selected Myoporaceae. Pollen et Spores 17: 497- 
516. 1975. 

Herbs or subshrubs, annual or perennial, 
branches usually slender and terete, pubescent or 
glabrous. Leaves alternate, sessile or petiolate, 
blades usually serrate, glandular punctate (viewed 
by transmitted light). Inflorescences of 1-3 axil- 
lary flowers borne on slender pedicels at distal 
nodes (racemose when distal leaves are reduced 
and bract-like), bracteoles absent beneath the ca- 
lyx. Flowers radially symmetric, calyx united 
only near the base, sepals 5, equal or subequal, 
narrow, valvate in bud; corolla funnelform or 
campanulate, white, purple, or greenish yellow, 
glabrous externally, tube as long as the lobes or 
slightly shorter, lobes 5, throat sometimes bearded 
within; stamens 4 or 5, equal or subequal, includ- 
ed, borne proximally or distally within the tube, 
filaments glabrous, anthers sagittate, basifixed, 
versatile, introrse, thecae divergent but basal parts 
confluent; ovary 2-locular, ovules many, style 
slender, apically flattened with stigmas on the lat- 
eral faces. Fruits capsules, ellipsoid to ovoid, 
glandular-punctate, dehiscing loculicidally and 
secondarily septicidal, 4-vaIved, placenta remain- 
ing as a conspicuous column with pitted-reticulate 
surface; seeds many, small, yellow to brown. 

Capraria is a genus of five species ranging 
from the southern United States to Peru; one spe- 
cies is adventive in West Africa. The narrow al- 
ternate leaves, flowers that are radially symmetric, 
and fruit with persisting reticulated column (pla- 
centa) make these plants distinctive. The campan- 
ulate five-lobed corolla and five stamens are un- 
usual in the Scrophulariaceae, so that the genus 
has never been satisfactorily classified within the 
family. Niezgoda and Tomb (reference above) 
showed that the pollen of Capraria is diorate and 
more closely related to types found in the My- 



oporaceae than to the pollen of other Scrophular- 
iaceae. 

Capraria biflora L., Sp. PI. 628. 1753. C. biflora 
var. pilosa Griseb., Fl. Brit. W.I. 427. 1861. C. 
biflora form hirta Loes., Bull. Herb. Boissier 2, 
3: 284. 1903. Figure 7. 

Herbs or subshrubs. perennial, erect or clam- 
bering, 0.5-2 m tall, distal stems usually few- 
branched, leafy stems 0.7-5 mm diam., densely 
puberulent to glabrescent with thin erect whitish 
hairs 0.2-0.7 mm long. Leaves alternate, gradu- 
ally becoming smaller on distal stems, petioles to 
8 mm long but usually not clearly differentiated; 
leaf blades 1.5-11 cm long, 6-30 mm wide, el- 
liptic to elliptic-oblanceolate or oblanceolate, 
apex acute and sharp-tipped, distal half of the 
margin with sharp-pointed teeth 0.5-3 mm high. 
1-6 teeth/cm, gradually narrowed to the cuneate 
base, drying grayish green, both surfaces with 
thin white hairs 0.1-0.4 mm long, 2 veins 3 or 
4/side, strongly ascending. Inflorescences of 2 or 
3 flowers in distal leaf axils, pedicels 4-12 mm 
long, ca. 0.2 mm diam.. minutely puberulent, not 
articulated at the base of the calyx. Flowers with 
calyx 4-6 mm long, sepals 1-1.5 mm wide at the 
base, narrowly triangular to linear, to 7 mm long 
in fruit, with short (0.3 mm) thin whitish hairs; 
corolla 7-10 mm long, 4-7 mm wide at the 
mouth, tubular-campanulate, white, lobes 3-5 mm 
long, 2.5-3.5 mm wide at base, triangular; fila- 
ments attached at the base of the tube. 3-4 mm 
long, anthers ca. 3 mm long. Fruits 4-6 mm long. 
2.5-4 mm wide, ovoid-ellipsoid and bisulcate, 
surface smooth, yellowish brown and punctate; 
seeds 0.3-0.4 mm long, oblong or angular. 

Plants of open sunny sites in lowland situations 
in both the Caribbean (evergreen) and northern 
Pacific (deciduous) areas of Costa Rica, 0-500 m 
elevation (to 1000 m in Nicaragua). Flowering 
and fruiting throughout the year but collected pri- 
marily in July-January. The species ranges from 
Florida, Mexico, and the West Indies to Argenti- 
na; it is found on Cocos Island and has become 
naturalized in West Africa. 

Capraria biflora is recognized by its long slen- 
der stems, alternate leaves that are smaller distal- 
ly, oblanceolate leaf blades serrate in the distal 
half, flowering nodes with two or three flowers on 
slender pedicels, and flowers with narrow sepals 
and radially symmetric campanulate white corol- 
la. The glandular punctations of the leaves can 
best be seen by transmitted light. This species is 



BURGER: FLORA COSTARICENSIS 



31 



uncommon in Central America. In Panama it is 
used to make a tea that may be a dangerous de- 
pressant in large quantities (D'Arcy, 1979). 



Castilleja Mutis ex Linnaeus filius 

RHFKRHNCE N. H. Holmgren, Castilleja (Scro- 
phulariaceae) in Costa Rica and Panama. Brittonia 
30: 182-194. 1978. 

Herbs or subshrubs, annual or perennial, erect, 
branching mostly from the base, hemiparasitic on 
roots of hosts, drying dark. Leaves alternate, cau- 
line, sessile or the petioles poorly differentiated, 
leaf blades entire to deeply dissected or pinnately 
lobed. Inflorescences terminal spikes or racemes, 
spikes usually with flowers hidden by colorful 
bracts, the racemes often with conspicuous flow- 
ers arranged along one side (secund), subtending 
leaves intergrading with bracts, distal bracts often 
more conspicuous than the flowers, bracteoles ab- 
sent at the base of the calyx. Flowers bilaterally 
symmetric, calyx tubular with 4 equal or unequal 
lobes or united into 2 entire lateral lobes, slightly 
enlarging in fruit; corolla bilaterally symmetric 
and strongly 2-lipped, greenish yellow to brightly 
colored, tube usually elongate and narrow, upper 
(adaxial) lip entire with united lobes hooded and 
forming a beak-like galea, enclosing the anthers, 
lower lip slightly 3-saccate, with 3 rudimentary 



teeth or petaloid lobes; stamens 4, of 2 unequal 
pairs, attached near or above the middle of the 
corolla tube, anther sacs (thecae) unequally placed 
on the connective, the outer longer and attached 
near the center, the inner theca smaller and borne 
on the apex of the connective; ovary 2-locular, 
ovules many, style slender, stigma capitate or 
slightly 2-lobed. Fruits dry capsules, usually 
asymmetric, ovate to globose, opening loculici- 
dally; seeds many, with a loose reticulate exotesta. 
A genus of ca. 200 species, ranging from North 
America into higher elevation Central America 
and the Andes, and with a few northern Asian and 
European species. These plants are easily recog- 
nized because of their colorful inflorescences in 
which bracts, sepals, and petals may all be vari- 
ously colored and showy. The calyx tube is some- 
what gibbous at the base (abaxially) and often 
with only two large lateral lobes; the narrow, 
curved corolla has a beaked upper lip that is usu- 
ally much longer than the lower lip. In many spe- 
cies all parts become blackish on drying. Many 
species of Castilleja have the reputation of being 
very variable and difficult to distinguish, making 
the delimitation of some species quite arbitrary. 
This problem manifests itself in Costa Rica, 
where C. irasuensis, C. quirosii, and C. talaman- 
censis form a closely related complex that de- 
serves further study. Our treatment is based on the 
publication (1978, cited above) and determina- 
tions of Noel Holmgren. 



Key to the Species of Castilleja 

la. Calyx 9-12 mm long, corolla 9-12 mm long; flowers sessile and difficult to see within the densely 
congested distal portion of the spicate inflorescence; specimens drying pale grayish green to dark 
gray; 700-2500 m elevation in Costa Rica C. arvensis 

Ib. Calyx 13-24 mm long, corolla 18-34 mm long; flowers short- to long-pedicellate and usually easily 
seen among the bracts; specimens drying grayish to blackish; 1000-3800 m in Costa Rica .... 2 

2a. Leaves to 2 cm long and nearly as wide, pinnatifid with narrow (1-2 mm) central rachis and slender 
filiform (1 mm) pinnate lobes; clefts of the calyx tube more nearly the same (adaxial 4-6.5 mm 
deep, abaxial 8-12 mm deep); rarely collected, 1 100-1600 m C. tayloriorum 

2b. Leaves much longer than wide, usually lanceolate to linear, the pinatifid lobes short or > 1 mm 
wide when present; clefts of the calyx tube differing greatly in length (adaxial 1-4 mm deep, abaxial 
9-17 mm deep); rare to common, 1400-3500 m 3 

3a. Leaves lanceolate, mostly 3-6 cm long; upper galeate lip equal to the corolla tube or up to 1.5 
times as long; rarely encountered at 1400-1700 m elevation C. lentii 

3b. Leaves mostly linear or pinnatifid, 1-4 cm long; upper galeate lip 2-3 times as long as the corolla 
tube; not collected from below 1600 m elevation [the following three species are part of a closely 
related complex] 4 

4a. Plants usually densely puberulent with stiff slender whitish hairs in most parts, the stems terete or 
with poorly defined longitudinal ridges, parts drying dark grayish; corolla tube 7-10 mm long; 
uncommon in Costa Rica, 1600-3500 m C. quirosii 



32 



FIELDIANA: BOTANY 



4b. Plants glabrous to sparsely pubemlent with slender whitish hairs, the stems usually with well-defined 
longitudinal ridges continuous with the leaf bases, parts usually drying black; corolla tubes 7-16 
mm long; commonly collected in Costa Rica, 2600-3800 m 5 

5a. Le.aves mostly entire, with short lobes near the apex; calyx red to brownish purple with yellow 
distal tips; 2600-3800 m in the Cordillera de Talamanca C. talamancensis 

5b. Leaves usually with prominent pinnate lobes in the distal half; calyx reddish throughout; 3000- 
3700 m on Volcan Irazu, Volcan Turrialba, and rarely collected in the Cordillera de Talamanca . . 

C. irasuensis 



Castilleja arvensis Schldl. & Cham., Linnaea 5: 
103. 1830. C. communis Benth. in DC., Prodr. 
10: 529. 1846. C. agrestis Pennell, Fieldiana, 
Bot. 28: 519. 1953. Figure 7. 

Erect herbs, annual, 1 5-80 cm tall, unbranched 
or with few lateral branches, leafy stems 0.7-4 
mm diam., sparsely to densely hirsutulous with 
thin whitish hairs 0.2-1.5 mm long, gland-tipped 
hairs also present. Leaves sessile and lacking a 
clearly defined petiole, gradually smaller distally 
and intergrading with the floral bracts; leaf blades 
15-60(-90) mm long, 4-14(-22) mm wide, ob- 
lanceolate to narrowly elliptic-obovate or elliptic, 
apex bluntly acute, margin entire, gradually nar- 
rowed to the cuneate base, drying greenish gray 
or dark gray, with thin hairs 0.2-0.5 m long on 
both surfaces, tripliveined. Inflorescences 4-15 
cm long, 2-3 cm diam., flowers subsessile and 
densely congested (becoming more distant in 
fruit), pubescent throughout with villous and glan- 
dular hairs, bracts 12-20 mm long, leaf-like to 
ovate-oblong, often green tipped with red or yel- 
low, enlarging slightly in fruit, pedicels 0-1 mm 
long. Flowers hidden within the bracts, calyx 9- 
12 mm long, ovate-oblong and slightly curved, 
viscid-villous, adaxial and abaxial clefts subequal, 
3-5 mm deep; corolla 9-12 mm long, greenish 
yellow, usually hidden within the calyx, tube 6- 
8.5 mm long, galeate upper lip 2.5-5 mm long, 
lower lip reduced, with 3 lanceolate lobes; ovary 
glabrous. Fruits 6-8 mm long, 3.5-5 mm wide, 
ovoid-oblong, dark brown, producing many seeds; 
seeds 0.6-1.2 mm long, 0.2-0.3 mm wide, wedge- 
shaped to narrowly rectangular with translucent 
truncated ends. 

Weedy plants of moist open sunny sites in ev- 
ergreen or partly deciduous forest areas, 700- 
2500 m elevation (to 3000 m in Guatemala). 
Flowering and fruiting throughout the year, but 
collected most often in November-March. The 
species ranges from central and northeastern Mex- 
ico to Paraguay; it is adventive in Hispaniola and 
Hawaii. 



Castilleja arvensis is distinguished by its an- 
nual growth, alternate grayish leaves (when 
dried), densely congested spikes in which the co- 
rollas are difficult to see, green bracts often red- 
dish at the tips, and the subsessile fruit producing 
numerous wedge-shaped seeds. The entire, nar- 
rowly elliptic leaves give this species a quite dif- 
ferent appearance from Costa Rica's other Castil- 
leja species. Small (3 mm) fleshy tubercle-like 
leaves may be present at the base of the stems 
near ground level. This is a well-defined species 
and the widest ranging of the genus; it belongs to 
section Epichroma. 



Castilleja irasuensis Oersted, Vidensk. Meddel. 
Dansk Naturhist. Foren. Kjobenhavn 1853: 27. 
1854. Figure 7. 

Herbs or subshrubs, 10-50 cm tall, branching 
mostly from the woody base, lateral branches usu- 
ally short, leafy stems 1-2 mm diam., mostly gla- 
brous, with longitudinal ridges. Leaves sessile, ar- 
ticulated at the base, with fewer lobes distally and 
becoming bract-like in the inflorescence; leaf 
blades 6-30 mm long, the central portion 1-2.5 
mm wide, linear and entire to pinnatifid with 1-3 
pairs of lobes 0.5-8 mm long and ca. 0.8 mm 
wide (the longest pair often also pinnately lobed), 
apex bluntly acute, margin usually revolute, base 
with parallel margins to the stem, drying black, 
minutely papillate puberulent above, with larger 
(0.1-0.2 mm) hairs beneath, venation obscure. 
Inflorescences 3-15 cm long, racemose and 1 -sid- 
ed, rachis hispid-villous, proximal bracts little dif- 
ferentiated from the leaves, distal bracts oblanceo- 
late, ca. 2 cm long, bright red, pedicels 2-6(-ll) 
mm long, ca. 0.3 mm diam., pubescent. Flowers 
with calyx 14-20 mm long, 4-6 mm wide, ovoid- 
tubular, adaxial cleft 1-4 mm deep, abaxial cleft 
9-15 mm deep, lobes rounded and entire, becom- 
ing red, puberulent; corolla 22-30 mm long, tube 
8-12 mm long, galeate upper lip 12-19 mm long, 
puberulent and green above, red along the mar- 



BURGER: FLORA COSTARICENSIS 



33 



gins; anthers ca. 2 mm long. Fruits 7-12 mm 
long. 

Plants of open habitats in high-montane forest 
and paramo formations, 3000-3700 m elevation. 
Flowering and fruiting throughout the year. The 
species is endemic to the eastern volcanoes (Irazu, 
Turrialba) and the western part of the Cordillera 
de Talamanca. 

Castilleja irasuensis is recognized by its re- 
striction to high-elevation habitats, alternate pin- 
natifid leaves, and colorful racemes. Plants often 
differ because they grow on volcanic ash, open 
disturbed sites, or moist depressions, and the spe- 
cies exhibits considerable variation. More impor- 
tant, the characters used to separate this species 
from C. talmancensis vary greatly within and be- 
tween populations. Also, the two species may hy- 
bridize in the Talamanca Mountains, and it ap- 
pears that the separation of the two species may 
be an arbitrary distinction. This problem is worthy 
of careful study in the field. The name gallito has 
been recorded for this species. 



Castilleja lentii N. Holmgren, Brittonia 30: 191. 
1978. Figure 7. 

Erect herbs, 20-60 cm tall, simple or 
branched, leafy stems 0.7-3 mm diam., with thin 
whitish hairs to 0.8 mm long or glabrescent, terete 
and with longitudinal ridges. Leaves sessile, often 
slightly thickened (articulated) at the base; leaf 
blades (1.4-)3-6 cm long, 2-7 mm wide, linear- 
oblanceolate to narrowly elliptic, apex acute or 
slightly rounded, distal half of the margin with 1- 
4 short (0.7-3 mm) pairs of lateral lobes, gradu- 
ally narrowed to the cuneate base, drying black, 
mostly glabrous above, with thin whitish hairs be- 
neath, tripliveined from near the base. Inflores- 
cences 4-9 cm long, terminal 1 -sided racemes, 
proximal bracts leaf-like and green, distal bracts 
oblanceolate to obovate and distally reddish, ped- 
icels 3-12 mm long, glabrous or pubescent. Flow- 
ers with calyx 12-24 mm long, 4-6 mm wide, 
glabrous, adaxial cleft 2-4 mm deep, abaxial cleft 
12-17 mm deep, lobes usually entire, pale green 
below, yellow or reddish distally; corolla 27-33 
mm long, tube 11-15 mm long, curved, galeate 
upper lip 12-18 mm long, 1-2 mm wide, green 
above and reddish along the sides, lower lip with 
short (1 mm) teeth. Fruits 9-12 mm long, 4-5 
mm diam., narrowly obovoid; seeds not seen. 

Plants of open sunny wet sites or moist cliff- 
sides in wet evergreen cloud forest formations. 



1400-1700 m elevation. Flowering material has 
been collected in April, August, and November. 
The species is only known from near Cachi (Car- 
tago Prov.) and the Rio Cascajal (San Jose Prov.). 
The species is endemic to the Caribbean slope of 
central Costa Rica. 

Castilleja lentii is recognized by its lower ele- 
vation habitat (among our species of Castilleja), 
the alternate oblanceolate leaves with short pin- 
nate distal lobes, glabrous sepals, and long slender 
upper corolla lip reddish along the lateral margins. 

Castilleja quirosii Standl., Publ. Field Mus. Nat. 
Hist., Hot. Sen 18: 1104. 1938. C. aurantiaca 
Pennell, Ann. Missouri Hot. Card. 27: 338. 
1940. C. chiriquiensis Pennell, loc. cit. 338. 
1940. C. seibertii Pennell, loc. cit. 339. 1940. 
C. bicolor Pennell, loc. cit. 340. 1940. Figure 7. 

Herbs or subshrubs, 30-90 cm tall, main stems 
with few to many branches, leafy stems 0.7-3 mm 
diam., terete or with poorly defined longitudinal 
ridges, densely hirsutulous with straight or curved 
hairs 0.2-0.5 mm long. Leaves sessile, a petiole 
not clearly differentiated, slightly thickened at the 
base (abaxially); leaf blades 6-30(-40) mm long, 
0.7-3 mm wide (not including the lobes), linear 
to linear-oblong, rounded at the apex, margin en- 
tire or with 1 or 2 pairs of short ( 1-2 mm) narrow 
(0.3-0.7 mm) lobes distally, base parallel-cuneate, 
drying dark grayish green, densely to sparsely pu- 
bescent with stiff hairs ca. 0.2 mm long, often 
scabrous, venation obscure. Inflorescences 4-12 
cm long, 1 -sided racemes, rachis and bracts mi- 
nutely (0.3 mm) puberulent, bracts ca. 25 mm 
long, 1-2 mm wide, proximal bracts leaf-like, 
marked with red or orange, pedicels 2-12 mm 
long, puberulent (sometimes with gland-tipped 
hairs). Flowers with calyx 17-20 mm long, 35 
mm wide, minutely puberulent, red to orange or 
yellowish at the tip, adaxial cleft 1-3 mm deep, 
abaxial cleft 11-16 mm deep; corolla (18-)22- 
28(-32) mm long, yellow to yellowish green with 
red margin, tube 7-10 mm long, curved, galeate 
upper lip 15-23 mm long, ca. 1.5 mm wide, 
densely puberulent on the dorsal (adaxial) margin; 
anthers ca. 1.7 mm long. Fruits 8-13 mm long, 
3-6 mm diam., ovoid-oblong, terminated by the 
persisting style base. 

Plants of open sunny sites or partly shaded sites 
in evergreen montane and open paramo forma- 
tions, 1600-3500 m elevation. The species ap- 
pears to do best in volcanic soils and burnt areas. 
Flowering and fruiting throughout the year. The 



34 



FIELDIANA: BOTANY 



species ranges from Volcan Irazu eastward to the 
Chiriqui highlands of Panama. 

Castilleja quirosii is recognized by its generally 
dense .short pubescence, small alternate linear 
leaves with few short distal lobes, colorful flowers 
in one-sided racemes, and unusual corollas. 
Among our species of Castilleja, this species has 
the widest altitudinal range within Costa Rica, al- 
though it is not frequently collected. This species 
and its close congeners should be studied in the 
field; it seems possible that C. quirosii might be 
included within a more broadly defined C. ira- 
suensis. 

Castilleja talamancensis N. Holmgren, Brittonia 
30: 187. 1978. 

Herbs and shrubs, erect or clambering, 0.1- 
1 .5(-2) m tall, simple or with many distal branch- 
es, woody at the base, stems 0.8-3.5 mm diam., 
glabrous or minutely (0.1 mm) puberulent, with 
prominent longitudinal ridges. Leaves sessile or 
subsessile, slightly articulated at the base, bract- 
like at the base of the inflorescence; leaf blades 
5-18(-25) mm long, 0.5-2 mm wide, linear to 
linear-oblong, often with short distal lateral lobes 
0.5-1.5 mm long, apex rounded, margins entire or 
with small lobes near the apex, base decurrent on 
the stem, drying black, glabrous or minutely (0.05 
mm) puberulent beneath, venation obscure. Inflo- 
rescences 2-8 cm long, 1 -sided racemes, rachis 
minutely hispidulous, proximal bracts leaf-like, 
distal bracts obovate with rounded apex, marked 
with red, purple, or green, pedicels 2-1 1 mm 
long, 0.2-0.4 mm diam., with erect thin hairs 0.1- 
0.2 mm long. Flowers with calyx 14-20 mm 
long, 3-4 mm diam., adaxial cleft 1-3.5 mm 
deep, abaxial cleft 9-17 mm deep, rounded at the 
apices, minutely puberulent along the veins, red 
to brownish purple with yellow distal margin; co- 
rolla 20-32 mm long, tube 7-12 mm long, hood- 
ed upper lip 13-22 mm long, green above, mar- 
gins red to orange, lower lip reduced with 3 nar- 
row teeth ca. 1 mm long. Fruits 6-9 mm long, 
4-5 mm diam., dark brown and smooth; seeds ca. 
2 mm long, C-shaped. 

Plants of open or partly shaded sites in high- 
montane forest and paramo formations, 2600- 
3800 m elevation. Flowering and fruiting through- 
out the year. The species is endemic to the Cor- 
dillera de Talamanca of Costa Rica. 

Castilleja talamancensis is recognized by its re- 
striction to higher elevations, narrow linear-ob- 
long leaves that may have three small lobes at the 



apex, and the colorful inflorescences. The distal 
bracts subtending undeveloped flowers are often 
bright red. Plants in protected sites become large 
and shrubby, while those on exposed ridges or 
open sites are small and have few branches. This 
species is particularly successful along the dis- 
turbed margins of the Carretera Interamericana. 
However, there is so much variation in this spe- 
cies, and within C. irasuensis and C. quirosii as 
well, that there is the strong possibility that all 
three would be better considered a single variable 
species, with C. irasuensis having priority. Care- 
ful field work with the living populations should 
help resolve this problem. The name ./for de Indio 
has been used for this species. 

Castilleja tayloriorum N. Holmgren, Brittonia 
30: 193. 1978. 

Slender herbs, annual or short-lived perennials, 
erect, 25-150 cm high, stems 0.4-1.5 mm diam., 
glabrous or with few minute (0.1 mm) appressed 
whitish hairs. Leaves sessile or appearing to have 
winged petioles (to the first lobes), deeply pin- 
natisect from a narrow (0.7-2 mm) rachis; leaf 
blades 5-20 mm long, 3-14 mm wide, with a 
slender (petiole-like) proximal half and the distal 
rachis bearing 2 or 3 pairs of narrow (1-2 mm) 
lateral lobes, the proximal lobes often with small- 
er distal lobes, apices rounded, margin entire and 
revolute, drying dark grayish, glabrous, venation 
obscure. Inflorescences 3-7 cm long, racemes 
(often 1 -sided), lower bracts leaf-like and green, 
distal bracts shorter and reddish, often deciduous, 
pedicels 3-8 mm long, ca. 0.4 mm diam., mi- 
nutely puberulent. Flowers with calyx 13-17 mm 
long, glabrous, adaxial cleft 4-6.5 mm deep, ab- 
axial cleft 8-12 mm deep, major lobes usually 
rounded and yellowish; corolla 22-25 mm long, 
curved, tube 8-12 mm long, ca. 1 mm diam., 
curved, galeate upper lip 13-15 mm long, finely 
puberulent and green dorsally. margins reddish, 
lower lip short. Fruits 7-10 mm long, ca. 4 mm 
wide, obovoid or oblong with abruptly truncated 
apex. 

Plants of open sites or cliffsides in wet ever- 
green cloud forest formations, 1 100-1600 m ele- 
vation. Flowers were collected in August and Oc- 
tober, fruiting in October. The species is known 
from only two collections: G6mez-Laurito 10180, 
from near the highway tunnel at Zurqui, and J. 
Taylor & C. Taylor 11901, from below Bajo La 
Hondura. Presently the species is only known 
from the upper Rio Zurqui drainage in Braulio 



BURGER: FLORA COSTARICENSIS 



35 



Carrillo National Park on the Caribbean slope of 
central Costa Rica. 

Castilleja tayloriorum is recognized by its low- 
er elevation habitat, small pinnatifid leaves with 
slender lateral lobes, glabrous calyx, slender 
curved corolla tube, and long narrow upper lip. 
Castilleja tayloriorum is probably most closely 
related to species of section Epichroma from 
south-central Mexico. 



Cymbalaria Hill 

REFERENCES G. Cufodontis, Die Gattung 
Cymbalaria Hill. Nachtrage und Zusammenfas- 
sung. Bot. Not. 1947: 135-156. 1947. D. Sutton, 
A Revision of the Tribe Antirrhineae. British Mu- 
seum (Natural History) & Oxford Univ. Press. 
1988. 

Weak-stemmed herbs, creeping or twining, pe- 
rennials, simple or much-branched. Leaves op- 
posite or alternate, simple, long-petiolate, leaf 
blades reniform to suborbicular, palmately lobed 
or toothed, venation palmate. Inflorescences ra- 
cemes, cymes, or solitary flowers in leaf axils, 
pedicels well developed, elongating in fruit, brac- 
teoles absent at the base of the calyx. Flowers 
small, calyx united only at the base, sepals 5, im- 
bricate (valvate) in bud, enlarging slightly in fruit; 
corolla strongly bilabiate, usually blue or purple, 
tube with a short, backward-pointing abaxial spur 
at the base, upper lip with 2 lobes, lower lip with 
3 lobes and elevated transverse ridge (upper pal- 
ate) restricting access to the throat; stamens 4, of 
2 unequal pairs, included, thecae separate and par- 
allel; ovary 2-locular, ovules many, stigma 2- 
lobed. Fruits dry capsules, opening by 2 pore-like 
slits, the 2 valves often splitting into 3; seeds el- 
lipsoid or crested, rugulose. 

A genus of nine species native to western Eu- 
ropean and the Mediterranean. It is classified in 
the tribe Antirrhineae, and its species were once 
placed in Linaria, but they differ in having soli- 
tary axillary flowers and leaves with palmate ve- 
nation. Several species are used in ornamental 
horticulture as groundcover and in hanging bas- 
kets. One adventive species is sometimes found 
on moist walls in Mexico and Central America. 

Cymbalaria muralis P. Gaertn., B. Meyer & J. 
Sherb., Oekon. Fl. Wetterau 2: 397. 1800. An- 
tirrhinum cymbalaria L., Sp. PI., 612. 1753. 



Linaria cymbalaria (L.) Miller, Gard. Diet., ed. 
8, #17. 1768. 

Herbs with slender creeping or pendent stems, 
often rooting from the nodes, internodes 0.5-10 
cm long, 0.5-1.7 mm diam., glabrous. Leaves 
mostly alternate, glabrous, petioles 10-45 mm 
long, 0.3-0.5 mm diam.; leaf blades 10-35 mm 
long, 9-40 mm wide, orbicular-oblate to reniform 
in outline, apex obtuse to rounded with apiculate 
tip, margins entire with usually 5 (3, 7) rounded 
lobes separated by short narrow sinuses, base sub- 
cordate to deeply cordate, drying dark greenish or 
grayish, with usually 3 major veins. Inflorescenc- 
es of solitary axillary flowers, pedicels 1 2-40 mm 
long, 0.4-0.7 mm diam., glabrous, elongating to 
6 cm and negatively phototropic after anthesis. 
Flowers glabrous externally, sepals 0.8-1.5 mm 
long, 0.5-0.8 mm wide, lanceolate to narrowly 
oblong, elongating to 2.5 mm long in fruit; co- 
rolla 7-9 mm long, lilac or pale violet to blue 
with yellow interior, tube 3.5-4 mm long (includ- 
ing the spur to 2 mm long), lower lip ca. 3 mm 
long. Fruits 2.7-4 mm long, 3-4 mm wide, glo- 
bose or rounded-oblong, thin-walled. 

Cymbalaria muralis, as its Latin epithet im- 
plies, is usually found on damp shaded walls. A 
native of Europe, this species has only been col- 
lected in San Jose and Tres Rios (1100-1300 m) 
in Costa Rica. The slender stems, small thin-tex- 
tured five-lobed cordate leaves, and solitary flow- 
ers with strongly bilabiate bluish corolla and 
backward-oriented spur make these plants quite 
distinctive. In addition, the pedicels are negatively 
phototropic after anthesis, moving the capsules 
into dark corners as they elongate. They are some- 
times planted as cover plants in gardens and are 
called Kenilworth ivy, Coliseum ivy, and penny- 
wort. 



Darcya Turner & Cowan 

REFERENCE B. Turner & C. Cowan, Darcya 
(Scrophulariaceae), a new genus from Central and 
South America. Phytologia 74: 267-270. 1993. 

Herbs or weak-stemmed subshrubs to 1 m tall, 
annual or perennial, glabrous or puberulent, dry- 
ing greenish to dark brown. Leaves opposite, sim- 
ple, petiolate, blades with serrulate margins, ve- 
nation pinnate or subpalmate with 3-5 major 
veins from near the base. Inflorescences terminal 
racemes with 2 flowers/node separated by well- 



36 



FIELDIANA: BOTANY 



developed internodes (flowers solitary in axils of 
distal leaves or bracts), pedicels slender, bracte- 
oles absent at the base of the calyx. Flowers with 
calyx deeply 5-parted, calyx lobes equal or sub- 
equal, valvate in bud, enlarging slightly in fruit; 
corolla tubular and strongly 2-lipped (bilaterally 
symmetric), white to blue or purple, tube shorter 
than the lips, lower (abaxial) lip 3-lobed and larg- 
er, upper (adaxial) lip 2-lobed; stamens 4, sube- 
qual or unequal, filaments borne at 2 levels on the 
corolla tube, anthers with straight white hairs on 
dorsal side; ovary ovoid, style short. Fruits thin- 
walled capsules, dehiscence loculicidal and sep- 



ticidal producing 4 valves (or irregular); seeds 
small, oblong to trapezoidal, testa reticulate. 

Darcya is a genus of three narrowly endemic 
species, restricted (respectively) to Costa Rica, 
Panama, and Colombia. These plants resemble 
species of Stemodia but differ in having pubescent 
anthers, very short styles, more clearly racemose 
inflorescences, three or five prominent veins from 
the base of the blade, and oblong trapezoidal 
seeds. The decurrent leaf bases merge with the 
longitudinal ridges (angles) of the stem in a dis- 
tinctive manner. This genus also resembles the 
Asiatic genus Limnophila. which has become an 
invasive weed in some areas. 



Key to the Species of Darcya 

la. Calyx and pedicels glabrous; leaf blades 15-45 mm long, ovate-triangular to ovate-elliptic; stamens 
4, subequal; plants of the wet Caribbean slope in central Costa Rica D. costaricensis 

Ib. Calyx and pedicels puberulent; leaf blades 17-60 mm long, ovate-lanceolate; stamens 4 in 2 unequal 
pairs; collected in the area of Boquete, Chiriquu Panama D. reliquiarum 



Darcya costaricensis (B. L. Turner) B. L. Turner, 
Phytologia 74: 268. 1993. Stemodia costaricen- 
sis B. L. Turner, Phytologia 73: 253. 1992. Fig- 
ure 3. 

Erect or sprawling herbs to 1 m tall with many 
internodes 3-6 cm long, leafy stems 0.7-2.5 mm 
diam., glabrous, terete but with 4 longitudinal 
ridges, nodes without an interpetiolar ridge. 
Leaves opposite, petioles 2-10 mm long, 0.4-1.2 
mm wide, lateral margins decurrent on the stem 
and continuous with the stem ridges; leaf blades 
15-45 mm long, 10-25 mm wide, ovate-triangu- 
lar to ovate-elliptic, apex acute, margin with 4-6 
teeth/cm, often revolute (dried), base obtuse to 
truncate, minutely punctate, glabrous or with few 
minute (0.1 mm) hairs beneath, subpalmate with 
3 prominent basal veins, 2 veins 2 or 3/side. In- 
florescences 5-10 cm long, glabrous racemes, 
basal flower pair subtended by reduced leaves 
(12-16 mm long), distal flowers subtended by ses- 
sile lanceolate or linear bracts 2-6 mm long, ped- 
icels 6-16 mm long, glabrous. Flowers glabrous 
externally, calyx lobes 2.5-3 mm long, to 4 mm 
in fruit, 0.3-0.5 mm wide, narrowly oblong, apex 
truncated and thickened, with 3 prominent parallel 
veins; corolla 4-7 mm long, blue or purple, tube 
3-4 mm long, puberulent internally near the 
mouth, central lobe of lower lip 3-6 mm long; 
stamens 4, subequal, anthers similar; stigma in- 



cluded. Fruits 3-4 mm long, 1.5-2 mm wide, 
narrowly ovate to oblong, persisting style 0.4-0.8 
mm long; seeds 0.5-0.7 mm long, yellowish 
brown, reticulate with longitudinal ridges separat- 
ed by rounded pits. 

Plants of open sites and along stream edges in 
very wet lower montane forest formations of the 
Caribbean slope, 1400-1600 m elevation. Flow- 
ering and fruiting throughout the year. This spe- 
cies is known only from along the upper Rio 
Grande de Orosf, Tapantf Refuge, Cartago Prov- 
ince, central Costa Rica. 

Darcya costaricensis is recognized by its weak- 
stemmed herbaceous habit, opposite leaves with 
broad serrulate blades, racemose inflorescences, 
narrow calyx lobes, two-lipped blue or purple co- 
rollas with short tube, and restricted geographical 
range. This material had earlier been placed in 
Stemodia reliquiarum (see the following species). 

Darcya reliquiarum (D'Arcy) B. L. Turner & C. 
C. Cowan, Phytologia 74: 269. 1993. Stemodia 
reliquiarum D'Arcy, Ann. Missouri Bot. Card. 
66: 258. 1979. 

Herbs with sprawling stems to 40 cm tall, leafy 
stems 0.7-3 mm diam., with 2 prominent ridges, 
glabrous except near the nodes with erect 5-celled 
hairs, glabrescent, drying brown. Leaves oppo- 
site, petioles 0-4(-8) mm long, with narrow lat- 



BURGER: FLORA COSTARICENSIS 



37 



eral margins dccurrent on the stem and poorly dif- 
ferentiated from the blade; leaf blades 1 .7-6 cm 
long, 10-26 mm wide, ovate-lanceolate to nar- 
rowly ovate, apex acute, margins serrulate distally 
with 10-15 teeth/side, base obtuse or truncate, 
punctate on both surfaces, glabrous, subpalmate 
with 3 major veins from the base, midvein with 2 
or 3 lateral veins/side. Inflorescences terminal 
open racemes, rachis 0.5-0.8 mm diam., bracts 
leaf-like to scale-like, mostly linear (ca. 3 X 1 
mm), ciliolate along the edge with multicellular 
gland-tipped hairs 0.1-0.3 mm long, pedicels 4- 
12 mm long, puberulent. Flowers small, calyx 
lobes 2-4 mm long, ca. 0.5 mm wide, minutely 
puberulent externally, with 3 longitudinal veins, 
drying brown; corolla 4-6 mm long, salverform, 
blue to lavender or purple, tube 3-4 mm long, 
slightly exceeding the calyx, subglabrous exter- 
nally, lobes 2-3 mm long, subequal, minutely pu- 
berulent at the base within; stamens 4, the lower 
2 filaments ca. 0.5 mm long and with reduced 
thecae. the upper filaments 0.7-1 mm long, in- 
serted near the middle of the tube, one theca ses- 
sile on the connective with the other on a stipe- 
like arm of the connective; ovary conical, style 
0.5 mm long, stigma club-like. Fruits 4-5 mm 
long, 2.4-3 mm wide, dehiscing apically into 4 
valves, placenta broad, unwinged; seeds many, 
0.4-0.6 mm long, oblong to trapezoidal, longitu- 
dinally reticulate, dark brown. 

Plants of evergreen montane cloud forests 
around Boquete, 1500-2500 m elevation. Proba- 
bly flowering throughout the year. Endemic to the 
Chiriqui highlands of western Panama. 

Darcya reliquiarwn is recognized by its her- 
baceous sprawling habit, opposite serrulate leaves 
with subpalmate venation, racemose puberulent 
inflorescences, narrow calyx lobes, strongly 2- 
lipped blue or purple corolla, pubescent anthers, 
and restricted geographical range. 



Digitalis Linnaeus 

REFERENCE K. Werner, Zur Nomenclature und 
Taxonomie von Digitalis L. Bot. Jahrb. Syst. 79: 
218-254. 1960. 

Erect herbs or rarely shrubs, biennial or peren- 
nial, stems simple or branched from the base. 
Leaves alternate, sessile or petiolate, entire to 
dentate, pinnately veined, drying greenish or 
grayish. Inflorescences terminal racemes, the 
flowers often aligned along one side, bracts sub- 



tending the pedicels, lacking bracteoles at the base 
of the calyx. Flowers large and colorful, calyx 
with 5 sepals united only near the base, imbricate 
in bud, enlarging slightly in fruit; corolla tubular 
to campanulate, longer than the calyx, purple to 
yellow or white, bilaterally symmetric with the 
lower lip slightly longer than the upper, upper lip 
entire or 2-cleft, lower lip 3-lobed, lateral lobes 
exterior in bud; stamens 4 in 2 similar pairs, in- 
cluded within the tube, anthers with 2 divergent 
thecae; ovary 2-locular, ovules many, style sim- 
ple, stigma 2-lobed. Fruits dry capsules, dehisc- 
ing septicidally; seeds many, minute, rugulose. 

Digitalis is a genus of about 20 species ranging 
from western Europe into central Asia. They are 
classified in the subtribe Digitalieae of the the 
tribe Veroniceae and are distinguished by their 
erect habit, one-sided racemes, and large tubular 
corollas. The common foxglove is found in gar- 
dens and as an escape in the cooler highlands of 
Central America. 

Digitalis purpurea L., Sp. PI. 621. 1753. Figure 8. 

Stout herbs, 0.5-1.8 m tall, biennial, leafy 
stems 2-8 mm diam., minutely puberulent with 
thin hairs 0.1-0.3 mm long. Leaves becoming 
progressively shorter from base to inflorescence, 
petioles 1-11 cm long but not well differentiated 
from the blade, 2-12 mm wide with winged mar- 
gins; leaf blades 2.5-20 cm long, 1-8 cm wide, 
narrowly elliptic to ovate-elliptic or elliptic-ob- 
ovate, apex acute to obtuse, margin serrate with 
5-10 teeth/cm, gradually narrowed to the cuneate 
base, minutely puberulent on both surfaces, 2 
veins 3-5/side, strongly ascending. Inflorescenc- 
es 15-40 cm long, flowers mostly along 1 side, 
bracts 8-18 mm long, lanceolate, sessile, pedicels 
5-10 mm long, 0.4-0.7 mm diam., puberulent. 
Flowers with calyx 8-16 mm long, sepals sepa- 
rate to the base, 4-7 mm wide, broadly ovate to 
ovate-oblong, ciliolate along the edge; corolla 
20-50 mm long, tube 1.3-2 cm wide, tubular- 
campanulate, pink to purple (white), lower lip ex- 
tending 3-8 mm beyond the upper lip, usually 
with dark spots within the throat; thecae divergent 
and equal, ca. 3 mm long. Fruits 9-1 1 mm long, 
9-10 mm diam., ovoid with broadly truncated 
base, minutely puberulent; seeds 0.4-0.7 mm 
long, narrowly rectangular. 

Digitalis purpurea, native to Europe, occurs as 
a garden plant and in small naturalized popula- 
tions in high-montane areas, 1800-3300 m ele- 
vation. This species is distinctive, with its alter- 



38 



FIELDIANA: BOTANY 



nate leaves gradually narrowed into the long pet- 
iole, conspicuous (often one-sided) racemes of 
large colorful flowers, and the open corolla tubes 
marked with dark spots within. Many varieties are 
used in ornamental horticulture. The plants are 
poisonous to livestock and contain a number of 
potent compounds. This species is the principal 
source of digitalin, an important drug in treating 
some kinds of heart ailments. Common names are 
digital, manga de la Senora, and foxglove. 



Escobedia Ruiz & Pav6n 

REFERENCES F. Pennell, Escobedia, a Neotrop- 
ical genus of Scrophulariaceae. Proc. Acad. Nat. 
Sci. Philadelphia 83: 411-426. 1931. J. Thieret, 
The Scrophulariaceae Buchnereae of Central 
America. Ceiba 8: 92-101. 1961. 

Perennial herbs, stems simple or branched, gla- 
brous or puberulent, striate or angled, roots yel- 
low to orange. Leaves opposite, sessile or sub- 
sessile, base cuneate to clasping, coriaceous, en- 
tire to serrulate, venation pinnate to subpalmate, 
often scabrous. Inflorescences racemose or with 
solitary flowers in axils of distal leaves, pedicels 
(peduncles) with 2 bracteoles distally or lacking 
bracteoles. Flowers large, calyx tubular to fun- 
nelform, with 5-10 longitudinal veins, lobes 3-6, 
shorter than the tube, triangulate to rounded; co- 
rolla salverform with a long slender tube, usually 
white, externally glabrous or puberulent, lobes 5, 
small to large, equal and rounded; stamens 4, sub- 
equal, inserted in the middle of the tube, filaments 
ciliate or glabrous, anthers glabrous and aristate; 
ovary 2-locular, ovules many, style elongate, stig- 
mas linear on the side of the style apex. Fruits 
dry capsules, ellipsoid, hard, included within the 
persisting calyx; seeds many, narrowly conical to 
linear-oblong, surface reticulate. 

Escobedia is a genus of six to eight species 
ranging from Mexico to Bolivia and Brazil. The 
genus is classified in the tribe Buchnereae, sub- 
tribe Melasmineae. These plants are distinctive 
among Neotropical Scrophulariaceae because of 
their large salverform white corollas with long 
narrow tubes. The roots have been used by indig- 
enous peoples as a source of a yellow dye for food 
coloring. Three species are found in northern Cen- 
tral America (Standley & Williams, 1973), but 
only one of these has been found in Costa Rica. 
Escobedia laevis Schldl. & Cham, (with long lin- 
ear leaves and calyx 4-7 cm long with narrow 



calyx lobes) ranges from southern Mexico to cen- 
tral Nicaragua. 

Escobedia grandiflora (L.f.) O. Kuntze, Rev. 
Gen. PI. 3: 231. 1893. Buchnera grandiflora 
L.f., Suppl. PI. 287. 1781. . scabrifolia Ruiz 
& Pav6n, Syst. Veg. Peruv. Chil. 159. 1798. 
Micalia grandiflora (L.f.) Raf., Fl. Tell. 2: 104. 
1837. E. curialis Pennell. Proc. Acad. Nat. Sci. 
Philadelphia 83: 417. 1931. E. longiflora Pen- 
nell, loc. cit. 423. 1931. E. reticulata Pennell. 
loc. cit. 420. 1931. Figure 6. 

Erect perennial herbs, 0.6-1. 5(-2) m tall, leafy 
stems 1.5-7 mm ilium., with stiff whitish hairs 
0.1-0.3 mm long, smooth or scabrous, roots 
bright yellow-orange. Leaves opposite or subop- 
posite, gradually becoming smaller distally, sub- 
sessile with petioles 1-4 mm long, ca. 2 mm 
wide; leaf blades 5-12 cm long, 1.5-3 cm wide, 
lanceolate to narrowly triangular or ovate-elliptic, 
apex acute, margin serrate with 1-3 teeth/cm, 
base rounded and truncate to auriculate. drying 
pale to dark gray and coriaceous, scabrous above 
and below with short (0.1-0.2 mm) hairs, tripli- 
veined from the base or palmately 5-veined. In- 
florescences of solitary flowers (2/node) in axils 
of smaller distal leaves (or the inflorescences ra- 
cemose if the smaller leaves are interpreted as 
bracts), pedicels 11-35 mm long, 0.7-2 mm 
diam., slightly curved, often with linear bracteoles 
to 4 mm long in the upper third. Flowers with 
calyx 34-44 mm long, 6-10 mm diam., scabrous, 
lobes 2-5 mm long, triangular; corolla 7-12 cm 
long, 6-8 cm wide at the mouth, salverform with 
a long narrow tube to 10 cm long, 3-8 mm diam., 
lobes 1-3 cm long and broadly rounded, white, 
minutely puberulent; stamens inserted on the mid- 
dle of the tube; ovary glabrous, style 5-8 cm long. 
Fruits 20-28 mm long, ca. 10 mm wide, smooth; 
seeds 3-4 mm long, 0.3-0.5 mm wide, linear-rect- 
angular, translucent except for the center. 

Uncommon plants of open sites, in marshy or 
wet situations of lower montane evergreen forest 
formations, 1000-1500 m elevation. Collected in 
flower in July-February. The species ranges from 
Mexico to Brazil. 

Escobedia grandiflora is recognized by its op- 
posite scabrous coriaceous subsessile leaves with 
three (five) major veins, distal nodes with two 
large opposite flowers, and large white corollas 
with long narrow tubes and widely flaring limb. 
The color, form, and size of the corolla suggest 
nocturnal pollination by hawkmoths. Escobedia 



BURGER: FLORA COSTARICENSIS 



39 



reticiilata (from near Canas Gordas, Pitiier 11118 
holotvpe t S) was distinguished by Pennell and 
Thicrct on the basis of a pustulate calyx and the 
size and position of the bracteoles. This is most 
likely a local variation and not deserving of spe- 
cies rank (D'Arcy, 1979, p. 220). 

Gerardia species are now placed in Agalinis 
(q.v.). 

Gibsoniothamnus is now placed in the Schle- 
geliaceae. 



Hemichaena Bentham 

REFERENCE J. Thieret, Synopsis of Hemichae- 
na, including Berendtiella (Scrophulariaceae). 
Fieldiana, Bot. 34: 89-99. 1972. 

Herbs or shrubs, simple or branched, glabrous 
or viscid-pubescent, drying brownish. Leaves op- 
posite or fasciculate, sessile or subsessile, margins 
dentate, surface often rugose, venation pinnate. 
Inflorescences axillary, of solitary flowers or 1 or 
2 cymes/axil, usually pedunculate with bracts sub- 
tending the well-developed pedicels, bracteoles 
absent at the base of the calyx. Flowers showy, 
calyx campanulate to tubular, with 5 prominent 
longitudinal costae, 5-lobed or 5-toothed, lobes 
unequal, shorter than the tube; corolla funnelform 
to tubular-campanulate, 2-lipped, bright yellow to 
orange or red, tube exceeding the calyx, lobes 
equal to or shorter than the tube, upper lip 2-lobed 
or emarginate, lower lip 3-lobed; stamens 4, sub- 
equal or of 2 unequal pairs, included or exserted, 
inserted at the middle or on the lower half of the 
tube, anthers 2-thecous, parallel but becoming di- 
vergent, staminode absent; ovary 2-locular, pla- 
centas bilamellate, ovules many, style slender, 
with 2 flattened stigmatic areas or lobes at the 
apex. Fruits dry capsules, ovoid to oblong, lo- 
culicidally dehiscent and secondarily septicidal; 
seeds very many, minute, linear-oblong to fusi- 
form, testa reticulate with thin translucent walls. 

A genus of five species, ranging from northern 
Mexico to the Talamanca mountains of Costa 
Rica. Hemichaena is classified in the tribe Do- 
dartieae, with Mimulus, Mazus, Leucocarpus, and 
Berendtiella. The genus was monotypic, but 
Thieret (1972, cited above) broadened the concept 
to include the four species of Berendtiella. Ge- 
neric delimitations within the tribe are problem- 
atic. If characters of the fruit are emphasized, then 



Hemichaena and Berendtiella appear closely re- 
lated, but Hemichaena can be distinguished by its 
ampliate corolla tube, included stamens that are 
attached near the base of the corolla, and cordate- 
amplexicaul leaves. The generic classification of 
the tribe Dodartieae will remain obscure until Mi- 
mulus is revised and the classification of the entire 
group is compared. Only the following disjunct 
species is found south of northern Nicaragua. 

Hemichaena fruticosa Ik-nib.. PI. Hartw. 78. 
1841. Leucocarpus fruticosus (Benth.) Benth. 
in DC., Prodr. 10: 336. 1846. 

Herbs or subshrubs to 2 m tall, distal stems 
with few or no lateral branches, leafy stems 2-1 1 
mm diam., terete or slightly quadrangular, densely 
pubescent with gland-tipped hairs 0.5-1.3 mm 
long. Leaves opposite, sessile, of similar size 
along the stems, equal or subequal at the node; 
leaf blades 5-17 cm long, 2-5 cm wide, lanceo- 
late to narrowly elliptic-lanceolate or oblanceo- 
late, apex acute or acuminate, margin serrate with 
36 small or prominent teeth/cm, base rounded 
and auriculate to cordate-amplexicaul, drying 
dark above and paler beneath, minutely glandular 
puberulent with hairs 0.2-0.6 mm long, 2 veins 
4-9/side. Inflorescences 3-8 cm long, usually of 
1 or 2 axillary cymes (1-4/node), peduncles 10- 
28 mm long, bracts 6-22 mm long, 1-5 mm wide, 
lanceolate, viscid puberulent with hairs ca. 0.5 
mm long, pedicels 5-18 mm long. Flowers with 
calyx 11-16 mm long, 4-6 mm diam., densely 
puberulent with gland-tipped hairs, lobes 3-8 mm 
long, narrowly triangular and acute; corolla 25- 
45 mm long, 14-28 mm wide at the mouth, tu- 
bular-campanulate, tube 6-11 mm wide at the 
mouth, minutely puberulent externally, lobes 5-9 
mm long; stamens subequal, inserted in the lower 
4 of the tube, filaments 12-15 mm long, glabrous, 
anthers 3.3-3.8 mm long; style 15 mm long, stig- 
mas flattened. Fruits 13-17 mm long, 4-6 mm 
diam., oblong, becoming brown; seeds 0.6-1 mm 
long, 0.3-0.4 mm wide, narrowed at each end. 

Plants of open moist to wet sunny sites along 
streams, on steep slopes, and in disturbed sites in 
montane evergreen forest formations, 1600-3100 
m elevation. Probably flowering and fruiting 
throughout the year, but collected most often in 
December-March. The species is found in south- 
ern Mexico and Guatemala and in the western 
portion of the Cordillera de Talamanca. 

Hemichaena fruitcosa is recognized by its stout 
viscid puberulent stems, sessile opposite serrate 



40 



FIELDIANA: BOTANY 



leaves, axillary cymes, bright yellow corollas that 
are only slightly bilabiate, and its restricted geo- 
graphical range (in Costa Rica). It is surprising 
that this species, which is so common in the west- 
ern half of the Cordillera de Talamanca, has not 
been collected elsewhere in Costa Rica. D'Arcy 
(1979) placed this species in synonomy under 
Leucocarpus perfoliatus (H.B.K.) Benth., but that 
species has smaller flowers, smaller leaves, and 
baccate white fruits. 



Lamourouxia Kunth in H.B.K. 
Nomen conservandum 

REFERENCE W. Ernst, Floral morphology and 
systematics of Lamourouxia (Scrophulariaceae: 
Rhinanthoideae). Smithson. Contrib. Bot. 6: 1-63. 
1972. 

Perennial herbs or subshrubs, usually woody at 
the base, stems erect or scandent, distal stems of- 
ten few-branched and arched, glabrous or pubes- 
cent, gland-tipped hairs often present, probably 
hemiparasitic on roots. Leaves opposite (verticil- 
late), sessile or short-petiolate, often smaller and 
bract-like distally, blades serrate to deeply dis- 
sected, drying brownish or black, pubescent with 
simple, branched, or gland-tipped hairs. Inflores- 
cences of solitary flowers in distal leaf axils (1 or 
2/node) or appearing racemose (paniculate or cor- 
ymb-like) when distal leaves are bract-like, pedi- 
cels ebracteolate at the apex. Flowers large and 
showy, calyx tubular-campanulate, 4-lobed, lobes 
subequal or of 2 unequal pairs (rarely cleft to 



base), obtuse to deltoid or linear, with 10 promi- 
nent longitudinal veins; corolla tubular and bila- 
biate, red to orange or white, usually puberulent 
on the exterior, upper lip often erect (rarely hood- 
like), entire or bilobed, lower lip usually shorter, 
narrow, 3-lobed, biplicate within below the apex; 
stamens 4 in 2 subequal pairs or with 2 fertile 
stamens and 2 staminodes, attached near base of 
corolla tube, filaments swollen and puberulent at 
the base, included in the upper lip, fertile anthers 
basi fixed and pilose; ovary 2-locular, ovules 
many, stigma terminal, exserted and 2-lipped. 
Fruits dry capsules, ovoid to ellipsoid with per- 
sisting style-base, opening loculicidally into 2 en- 
tire valves; seeds many, ellipsoid to oblong, mi- 
nutely bullate to reticulate. 

Lamourouxia is a genus of ca. 24 species found 
in Neotropical highlands from Mexico to Peru. 
The center of species richness is in Mexico, with 
only three species found in southern Central 
America and two species in the Andes. Although 
a very distinctive genus, the individual species 
can be difficult to separate from each other. In 
Costa Rica the plants are restricted in distribution 
and not often collected. The large showy flowers 
with bright red or orange narrowly tubular corol- 
las, spicate or racemose inflorescences, four-lobed 
calyx, subsessile leaves, and preference for open 
habitats at mid-elevations help distinguish this ge- 
nus. Our species belong to section Hemispadon, 
according to Ernst's fine monograph (1972, cited 
above). Lamourouxia is in the subfamily Rhin- 
anthoideae. Every member of this subfamily that 
has been tested is hemiparasitic, perhaps explain- 
ing why these lovely plants are not seen in gar- 
dens. 



Key to the Species of Lamourouxia 

la. Calyx glabrous; young stems glabrous except for longitudinal lines of minute hairs; leaves linear- 
lanceolate; 1600-2800 m elevation L. lanceolata 

Ib. Calyx puberulent externally; young stems pubescent; leaves narrowly elliptic to ovate-triangular; 
900-1800 m elevation 

2a. Stems and flowers with simple (non-gland-tipped) hairs, leaves 7-24 mm long; flowering mostly 
November-February L. gutierrezii 

2b. Stems and flowers with gland-tipped hairs; leaves 10-110 mm long; flowering in July-December 

. L. viscosa 



BURGER: FLORA COSTARICENSIS 



41 



Lamourouxia gutierrezii Oerst. in Benth. & 
Ocrst., Vidcnsk, Meddel. Dansk Naturhist. For- 
en. Kjobenhavn 1853: 29. 1853. L scabra See- 
mann, Bot. Voy. Herald 177: pi. 33. 1854. Fig- 
ure 5. 

Herbs or subshrubs, 0.5-1. 3(-2) m tall, distal 
branches arched and often with short lateral 
branching, leafy stems 1-4 mm diam., sparsely to 
densely puberulent with thin straight or retrorse 
hairs 0.2-0.5 mm long. Leaves opposite (verticil- 
late), sessile or subsessile with poorly differenti- 
ated petioles to 2 mm long; leaf blades 7-24 mm 
long, 2-10 mm wide, elliptic to elliptic-oblong or 
ovate-elliptic, apex obtuse or rounded, margin 
with 4-6 rounded teeth/cm, base cuneate, drying 
dark grayish green, slightly scabrous above and 
below with short stiff hairs ca. 0.2 mm long, 2 
veins 3-5/side. Inflorescences 5-20 cm long, spi- 
cate or racemose, bracts 8-12 mm long, 5-6 mm 
wide, leaf-like and lanceolate to ovate, sometimes 
caducous and the flowers appearing ebracteate, 
pedicels 2-5 mm long, ca. 0.7 mm diam., puber- 
ulent. Flowers with calyx 6-9 mm long, enlarging 
to 12 mm in fruit, 2-4 mm diam., cupulate, 5- 
lobed, lobes 2-4 mm long, triangular to lanceo- 
late; corolla 30-44 mm long, 4-6 mm diam., tu- 
bular, bright red to orange-red, densely puberulent 
externally, upper lip ca. 14 mm long; functional 
stamens 2, anthers connivent and positioned 
against the upper lobe, 3-4 mm long with hairs 
1-2 mm long; ovary glabrous, style 2-4 cm long. 
Fruits 8-13 mm long, 4-8 mm diam., ovoid, 
short-beaked, dark and smooth; seeds 0.8-1 mm 
long, 0.3-0.4 mm thick. 

Infrequently collected plants of open sites in 
lower montane evergreen forest formations, 900- 
1900 m elevation. Probably flowering throughout 
the year, with most collections made in Novem- 
ber-February and June-July. The species ranges 
from the western part of the Meseta Central in 
Costa Rica to the Chiriqui highlands in Panama. 

Lamourouxia gutierrezii is recognized by its 
smaller narrow scabrous leaves, simple pubes- 
cence, and puberulent calyx. The structure and 
position of the staminodes suggest that they may 
function as a barrier to keep visiting insects from 
the ovary. This is the only species of the genus 
endemic to southern Central America. 

Lamourouxia lanceolata Benth. /// DC., Prodr. 
10: 542. 1846. L. longiflora var. lanceolata 
(Benth.) L. O. Williams, Fieldiana, Bot. 34: 
121. 1972. Figure 5. 



Herbs or shubshrubs to 1 .5(-3) m tall, erect or 
scandent, stems often arching and with multiple 
branching, leafy stems 1-3 mm diam., with mi- 
nute (0.2 mm) crooked hairs in 2 opposing lon- 
gitudinal lines along the stems. Leaves sessile or 
subsessile with poorly defined petioles to 2 mm 
long, not conspicuously smaller distally; leaf 
blades 1 1-48(-57) mm long, 2-8(-14) mm wide, 
narrowly lanceolate to narrowly linear-lanceolate 
or narrowly elliptic-oblong, apex acute, margin 
with 4-6 teeth/cm, base narrowly cuneate, drying 
brownish to blackish, glabrous or with minute 
(0.1 mm) papillate-puberulent hairs on the mid- 
vein above, 2 veins 4-7/side. Inflorescences 3- 
15 cm long or of flowers axillary to distal leaves, 
flowers subtended by leaves very similar to those 
lower on the stems, pedicels 3-9(-14) mm long, 
0.4-0.8 mm diam., glabrous or with a line of hairs 
along one side. Flowers with calyx 7-18 mm 
long, 3-5 mm diam., glabrous externally, veins 
elevated, lobes 6-15 mm long and becoming re- 
flexed; corolla 25-47 mm long, 4-7 mm diam., 
tubular, bright red to red-orange, densely puber- 
ulent externally, upper lip 14-18 mm long; sta- 
mens 2, with larger pubescent anthers often co- 
herent, staminodes 2, borne above the fertile sta- 
mens, thickened at the apex. Fruits 8-14 mm 
long, 5-8 mm diam., ovoid-rounded, beaks 1-2 
mm long; seeds 1-1.3 mm long, oblong to wedge 
shaped, surface reticulate. 

Infrequently collected plants of open sites in 
evergreen montane forest formations, 1600- 
2800(-3200) m elevations. Flowering in Decem- 
ber-March. In Costa Rica this species is known 
only from the western portion of the Cordillera de 
Talamanca. The species ranges from central Mex- 
ico to Costa Rica. 

Lamaourouxia lanceolata is recognized by its 
narrow opposite leaves, glabrous calyx with re- 
flexed lobes, narrow bright red corolla tubes, an- 
droecium of two stamens with pubescent anthers 
and two staminodes, and higher elevation habitats. 
This species has been confused with L. longiflora 
Benth. (Williams, 1972; Standley & Williams, 
1973), but L. longiflora has four fertile stamens 
and the calyx is puberulent on the exterior. 

Lamourouxia viscosa Kunth in H.B.K., Nov. 
Gen. Sp. 2 ed., folio 272. 1817; ed. quarto 338. 
1818. L. veijensis Oerst. in Benth. & Oerst., Vi- 
densk. Meddel. Dansk. Naturhist. Foren. Kjob- 
enhavn 1853: 28. 1853. 



42 



FIELDIANA: BOTANY 



Erect perennial herbs 0.5-1. 3(-3) m tall, leafy 
stems 1 .5-7 mm diam., terete, densely puberulent 
with gland-tipped hairs 0.2-0.7 mm long. Leaves 
gradually diminishing in size distally, sessile or 
subsessile with petioles ca. 1 mm long, appearing 
amplexicaul; leaf blades l-7(-ll) cm long, 6- 
28(-53) mm wide, ovate-triangular to narrowly 
triangular-oblong, apex acute, margin serrate with 
2-6 teeth/cm, base truncate to cordate-auriculate 
and amplexicaul, drying dark grayish to dark 
brown and coriaceous, with short (0.3 mm) hairs 
above and more densely puberulent beneath, 2 
veins 3-7/side. Inflorescences 10-20 cm long, ra- 
cemes or spicate, glandular puberulent, flowers 
crowded or separated by internodes to 12 mm 
long, bracts 5-16 mm long, 3-1 1 mm wide at the 
base, ovate to lanceolate, sessile, pedicels l-3(-6) 
mm long. Flowers glandular-puberulent external- 
ly, calyx 4-8 mm long, 3-5 mm diam., lobes 2- 
4 mm long, triangular and acute; corolla 14-38(- 
60) mm long, 3.5-6(-10) mm diam., tubular, dark 
red to red-orange (pink or purple), upper lip 6-12 
mm long, lower lobes ca. 2 mm long; functional 
stamens 2; thecae glabrous. Fruits 6-12 mm long, 
4-7 mm diam., ovoid, dark; seeds 0.8-1.2 mm 
long, 0.4-0.6 mm thick, wedge shaped, with a 
deeply reticulated surface. 

Plants of open sites in lower montane evergreen 
or partly deciduous formations, 1000-1800 m el- 
evation (500-2800 m in Mexico). Flowering pri- 
marily in July-December in Costa Rica (flowering 
and fruiting throughout the year in Nicaragua). In 
Costa Rica this species has been collected only at 
the eastern and western edges of the Meseta Cen- 
tral and in the region between Paraiso and Orosi. 
This species ranges from northern Mexico to Pan- 
ama. 

Lamourouxia viscosa is recognized by its mi- 
nute gland-tipped hairs on stems and flowers, stiff 
sessile opposite leaves that often surround the 
stem at their cordate bases, and lower montane 
habitats. This species is most closely related to 
the Mexican species L smithii Pringle and L 
rhinanthifolia Kunth in H.B.K. Costa Rican col- 
lections of L. viscosa do not possess leaves and 
flowers as large as some found in Mexico. 



Leucocarpus D. Don in Sweet 

Herbs, stems erect, quadrangular in cross-sec- 
tion and often with 4 longitudinal wings, glabrous 
or sparsely puberulent. Leaves opposite, sessile, 
narrow and serrulate, cordate-amplexicaul at the 



base, venation pinnate, drying yellowish brown. 
Inflorescences cymose. solitary in distal leaf ax- 
ils, peduncles with bracteoles subtending the ped- 
icels or secondary inflorescence branches, brac- 
teoles absent at the base of the calyx. Flowers 
with cupulate thin-walled calyx, 5-veined. per- 
sisting in fruit, calyx lobes narrow and tooth-like; 
corolla tubular or tubular-campanulate. somewhat 
bilaterally symmetrical with subequal upper and 
lower lips, glabrous externally, upper lip exterior 
in bud, 2-lobed, lower lip 3-lobed, puberulent at 
the mouth within; stamens 4 in 2 unequal pairs 
(or with only 2 fertile stamens), anthers with 2 
slightly diverging thecae, staminodes rarely pres- 
ent; ovary conical, 2-locular. ovules many, style 
thickened at the apex, stigma 2-lobed. Fruits 
fleshy indehiscent berries, white, pericarp thin, 
style base forming a small beak; seeds immersed 
in pulp, minutely reticulate. 

Leucocarpus is a monotypic genus, ranging 
from Mexico to Bolivia. It is included in the tribe 
Dodartieae, with Hemichaena, Mimulus. and Ma- 
zus. At one time, Bentham included Hemichaena 
in Leucocarpus, but the genera are now consid- 
ered distinct. 

Leucocarpus perfoliatus (Kunth in H.B.K.) 
Benth. in DC., Prodr. 10: 335. 1846. Mimulus 
perfoliatus Kunth in H.B.K., Nov. Gen. Sp. 2: 
371. 1817. Conobea alata Graham, Edinburgh 
New. Philos. J. 10: 168. 1830. L alatus (Gra- 
ham) D. Don in Sweet, Brit. Fl. Card. 2: tab. 
124. 1833. Figure 6. 

Erect herbs or shrubs 0.5-2.5 m tall, flowering 
stems usually unbranched. leafy stems 3-8 mm 
wide, with 4 prominent longitudinal wings (ridg- 
es), wings 0.3-2 mm wide, usually glabrous. 
Leaves of the same node slightly united at the 
base to form a short interpetiolar ridge, sessile and 
amplexicaul; leaf blades 8-23(-28) cm long, 1- 
4.5(-6) cm wide, narrowly lanceolate to very nar- 
rowly elliptic-oblong, tapering gradually to the 
acute or acuminate apex, margin serrulate with 3- 
5 teeth/cm, cuneate in the lower V4 but slightly 
expanded at the rounded-auriculate base, drying 
thin-chartaceous, usually glabrous. 2 veins 10- 
14/side. Inflorescences 2-6 cm long, axillary 
cymes or with 2 branching, with 2-12 flowers, 
peduncles 1-3 cm long, 0.7-1.3 mm diam., gla- 
brous or minutely puberulent with thin hairs 0.1- 
0.2 mm long, bracts 3-8 mm long, lanceolate, 
pedicels 4-14 mm long, minutely puberulent. 
Flowers with calyx 5-9 mm long, 2.5-3.5 mm 



BURGER: FLORA COSTARICENSIS 



43 



diam.. campunulate, with narrow acute lobes 1-3 
mm long, sparsely minutely puberulent; corolla 
(12-)14-17(-23) mm long, tubular with slightly 
expanded lobes, yellow to white, marked with 
yellow and barbate within, tube 2-5 mm diam., 
lobes slightly unequal; filaments 5-10 mm long; 
style 5-7 mm long. Fruits globose, 7-10 mm 
diam.. white, fleshy; seeds 0.4-0.5 mm long, 0.2- 
0.3 mm wide, ellipsoid-oblong, yellowish brown, 
reticulation minute (visible at 50X). 

Plants of open moist to wet sites in lower mon- 
tane evergreen forest formations, 800-2200 m el- 
evation. Probably flowering and fruiting through- 
out the year, but with most Costa Rican collec- 
tions made in December-January and April-June. 
The species ranges from Veracruz, Mexico, to the 
highlands of western Panama and Bolivia. 

Leucocarpus perfoliatus is recognized by the 
unbranched flowering stems with winged margins, 
sessile leaves with auriculate bases clasping the 
stems, axillary inflorescences, always shorter than 
the leaves, smaller tubular yellow corollas, and 
baccate fruit. Superficially, these plants are simi- 
lar to Hemichaena fruticosa (q.v.), but that spe- 
cies has larger flowers and dry dehiscent fruits. 



Limosella Linnaeus 

REFERENCE H. Gliick, Limosella studien. Bei- 
trage zur Systematik, Morphologic, und Biologic 
der Gattung Limosella. Bot. Jahrb. Syst. 66: 490- 
563. 1934. A. Lourtieg, Etude sur Limosella. 
Comite Nacional Fran^ais des Researches Antarc- 
tiques: Biologic 1(10): 165-173. 1964. 

Very small herbs, annual or perennial, growing 
in shallow water or on moist soil, caespitose or 
with short stolon-like stems rooting from the 
nodes, glabrous or puberulent. Leaves from the 
apex of the rootstock or fasciculate at the nodes, 
densely clustered, sessile or with poorly differ- 
entiated petioles, blades linear to obovate, entire, 
flat or cylindric, drying brown or yellowish, gla- 
brous, venation obscure or with a midvein. Inflo- 
rescences of 1 to several flowers from the leaf 
axils, glabrous, bracts absent, pedicels usually 
shorter than the leaves, bracteoles absent at the 
base of the calyx. Flowers very small, glabrous 
externally, calyx campanulate, usually thin- 
walled, equal to or slightly shorter than the corolla 
tube, lobes 5 (4) with the posterior lobe exterior; 
corolla funnelform to campanulate, white to blu- 
ish or pinkish, tube short, lobes 3-5, subequal and 



imbricate in bud, equal to or shorter than the co- 
rolla tube; stamens 4 (2, 5), included, filaments 
simple, borne near the middle of the tube, fila- 
ments of lower stamens crossing over the fila- 
ments of upper stamens, anther thecae confluent 
(1-thecous), not mucronate; ovary ovoid to ellip- 
soid, 2-locular near the base, style short, stigma 
capitate. Fruits capsules or indehiscent, usually 
bivalvate; seeds small, numerous, ovoid, striate 
and reticulate. 

A genus of ca. 1 1 species distributed widely 
over temperate and tropical montane regions but 
most diverse in the Southern Hemisphere. The 
very small size of the plants, their small flowers, 
and their aquatic or moist habitats help distinguish 
the genus. The genus is classified in the tribe Gra- 
tioleae and is often placed in the subtribe Limo- 
sellineae. 

Limosella acaulis Sesse & Mocino, Fl. Mex. ed. 
2: 143. 1894. L. americana Gliick, Notizbl. Bot. 
Gart. Berlin Dahlem 12: 75. 1934. L. americana 
f. submersa Gliick, loc. cit. 75. 1934. L. amer- 
icana f. natans Gliick, loc. cit. 76. 1934. L. 
americana f. terrestris Gliick, loc. 76. cit. 1934. 
Figure 1. 

Small herbs to 5 cm tall, stems to 6 cm long 
or little developed (and plants acaulescent), root- 
ing at the nodes, glabrous. Leaves rosulate or fas- 
ciculate, sessile or with poorly differentiated pet- 
ioles to 4 cm long, clasping the stem at their base, 
glabrous throughout; leaf blades 4-15 mm long 
(-11 cm when linear), 0.7-4 mm wide, linear to 
narrowly oblong to spatulate or oblanceolate, 
apex usually rounded, margins entire, gradually 
narrowed at the base, drying brown or yellowish, 
midvein usually visible only near the base. Inflo- 
rescences of 1-3 flowers in leaf axils (but difficult 
to see among the crowded leaf bases), pedicels 
10-30 mm long, ca. 0.4 mm wide, glabrous. 
Flowers glabrous externally, calyx 1.5-2 mm 
long, lobes 5, 0.3-0.7 mm long, narrowly trian- 
gular to oblong, dark coloring sometimes present 
between the lobes distally; corolla ca. 4 mm long, 
campanulate or subrotate, slightly zygomorphic 
(bilaterally symmetric), white, tube ca. 2.5 mm 
long, ca. 1.3 mm diam., lobes 3-5, 1-1.5 mm 
long, rotate; stamens 4 in 2 opposing pairs. Fruits 
1.54 mm long, 1-3 mm diam., ovoid-ellipsoid, 
glabrous; seeds 0.5-1 mm long, 0.2-0.5 mm wide, 
striate. 

Rarely collected plants in shallow water or wet 
soils, found only above 3300 m elevation in Cen- 



44 



FIELDIANA: BOTANY 



tral America. Probably flowering and fruiting 
throughout the year. Known in Costa Rica from a 
single collection made in the Valle de los Conejos 
on Chirjipo Grande (3400 m) in February (Weston 
12360). The species ranges disjunctly from Mex- 
ico to Venezuela and Bolivia. 

Limosella acaulis is distinguished by its moist 
and very high altitude habitat, the small size of 
the plants, mostly fasciculate linear sedge-like 
leaves, and very small flowers with white corollas 
that have three to five lobes. When seen from 
above, the flowers may be reminiscent of Hous- 
tonia (Rubiaceae), with four corolla lobes held in 
a single horizontal plane. Species are variable and 
individual populations are often isolated, so local 
variation is common. Also, the plant's habit 
changes with water depth and length of immer- 
sion. When submersed, leaves develop to be lon- 
ger and more linear than spatulate; flowers remain 
closed and are cleistogamous. The Costa Rican 
collection describes the plants as forming small 
mats in a creek bed. Lourtieg (1964, cited above) 
placed most taxa as synonyms of L australis R. 
Br., but we follow the decisions of Louis Williams 
regarding the circumscription of this species (Fiel- 
diana, Bot. 34: 121, 1972). 



Linaria Miller 

REFERENCE D. Sutton, A. Revision of the 
Tribe Antirrhineae. British Museum (Natural His- 
tory) & Oxford Univ. Press, 1988. 

Herbs or subshrubs, annual, biennial or peren- 



nial, erect, branching, mostly from the base, gla- 
brous or less often glandular-puberulent, drying 
grayish to dark brown. Leaves alternate, some- 
times opposite or verticellate near the base, leaf 
blades usually sessile, entire, linear to reniform, 
pinnately veined. Inflorescences colorful racemes 
or spikes (rarely flowers solitary in leaf axils), al- 
ternate along the rachis, bracts usually small, 
bracteoles absent at base of calyx. Flowers usu- 
ally glabrous, calyx campanulate, with 5 sepals or 
5 calyx lobes, equal or subcqual, imbricate in bud, 
persisting in fruit; corolla tubular and strongly bi- 
labiate, tube with a backward-pointing abaxial 
spur, upper (adaxial) lip bilobed and exterior in 
bud, lower (abaxial) lip 3-lobed and with a prom- 
inent raised palate near the entrance to the tube; 
stamens 4 in 2 pairs, included, filaments attached 
near the base of the tube, anthers with 2 parallel 
or divergent thecae, a staminode sometimes pre- 
sent; ovary 2-locular, ovules many, style slender, 
stigma small, 2-lobed or capitate. Fruits thin- 
walled capsules, ovoid to globose, dehiscing lo- 
culicidally into 2 valves or forming distal pores; 
seeds ovoid to discoid or C-shaped, testa thin, 
smooth to rugose. 

Linaria is a genus of ca. 100 species, native to 
north temperate regions and the Mediterranean 
area. They are often grown in gardens for their 
colorful and complex flowers or their foliage. The 
corollas with backward-pointing spur and strongly 
two-lipped lobes are distinctive. The genus is 
classified in the tribe Antirrhineae and is closely 
related to Cymbalaria (q.v.). Although not known 
to be naturalized in Costa Rica, the following two 
species are likely to be present. 



Key to the Species of Linaria 

la. Flowers yellow, corollas 20-30 mm long; planted in gardens and rarely escaping . . . L. vulgaris 

Ib. Flowers blue to purple, corollas 5-15 (20) mm long; weedy plants of cooler climates 

. L canadensis 



Linaria canadensis (L.) Dumort., Bot. Cult. 2: 
96. 1802. Antirrhinum canadensis L., Sp. PI. 
618. 1753. L. texana Scheele, Linnaea 21: 761. 
1848. 

Herbs, annual or biennial, flowering stems 
erect from a basal rosette of leafy stems, 15-50 
cm tall, branching from the base, leafy stems 0.5- 
2.5 mm diam., glabrous. Leaves opposite or 
whorled near the base, alternate on erect stems. 



sessile; leaf blades of erect stems 5-20 mm long, 
1-2 mm wide, linear, apex acute to bluntly acute, 
margin entire, base cuneate, venation obscure. In- 
florescences racemes (or appearing spicate), usu- 
ally making up the distal half of erect stems, gla- 
brous, bracts ca. 2 mm long, pedicels 1-4 mm 
long. Flowers glabrous externally, calyx 2-3.5 
mm long, sepals 5, 0.5-1 mm wide; corolla 5- 
15(-20) mm long, blue to purple, spur 2-9 mm 
long, tube strongly 2-lipped, the throat open, 



BURGER: FLORA COSTARICENSIS 



45 



white: stamens 2-3 mm long; ovary 1-2 mm long, 
ovoid, style 1-2 mm long. Fruits 2-3.5 mm long, 
subglobose to rounded oblong, with persisting 
style 0.5 mm long; seeds 0.3-0.5 mm long, an- 
gled, smooth. 

Linaria canadensis, native from southern Can- 
ada to northern Mexico, is a weed of open sunny 
sandy sites in cooler temperate or montane trop- 
ical climates (ca. 2000 m elevation in Mexico). 
Although not yet recorded for Central America, it 
is naturalized in South America and is likely to 
become established in our area. The unbranched 
erect stems, linear leaves, slender racemes, and 
blue corolla with spur help to distinguish this spe- 
cies. 

Linaria vulgaris Miller, Card. Diet. ed. 8. 1768. 
Antirrhinum linaria L., Sp. PI. 616. 1753. 

Erect perennial herbs 0.2-0.8 m tall, spreading 
by underground stems to form persisting clumps, 
erect stems usually unbranched, 1-3 mm diam., 
usually glabrous. Leaves alternate and numerous 
along the stem, glabrous, sessile; leaf blades 3-5 
cm long, 1-6 mm wide, linear to linear-lanceolate, 
secondary veins obscure. Inflorescences spicate 
racemes with crowded flowers, bracts ca. 1 cm 
long, similar to the leaves, pedicels 1-4 mm long, 
pedicels and flower oriented upward at an acute 
angle to the stem. Flowers with glabrous calyx, 
sepals 2-3.5 mm long, narrowly ovate, acute; co- 
rolla 2-3 cm long, yellow with orange near the 
mouth, tube with a spur equaling the tube and 
lobes in length, mouth of the corolla strongly bi- 
labiate, upper (adaxial) lip 8-12 mm long, lower 
(abaxial) lip 6-9 mm long, with a puberulent 
rounded palate. Fruits 5-10 mm long, glabrous; 
seeds ca. 2 mm long, discoid and winged. 

Linaria vulgaris, a native of Eurasia, is occa- 
sionally grown in gardens for its unusual yellow 
flowers, but it is more likely to be seen as a weed 
in cooler climates of the world. It is recognized 
by its often linear alternate leaves and the unusual 
two-lipped yellow corolla with basal spur. This 
species has not been recorded as naturalized in 
Central America. These plants sometimes produce 
abnormal flowers that are radially symmetric (not 
two-lipped) and have five spurs or none. Linnaeus 
mistakenly erected the genus Peloria for such 
plants, and the word peloria is now used to de- 
scribe abnormal radially symmetric forms of flow- 
ers that are normally strongly bilaterally symmet- 
ric (zygomorphic). 



Lindernia Allioni 

REFERENCES D. Miranda, Flavonoid and mor- 
phological studies of Lindernia Allioni (Scrophu- 
lariaceae) in South America. Bot. J. Linn. Soc. 75: 
47-67. 1977. D. Philcox, Revision of the Male- 
sian species of Lindernia All. (Scrophulariaceae). 
Kew Bull. 22: 1-72. 1968. T. Yamazaki, Revision 
of the Indo-Chinese species of Lindernia All. J. 
Fac. Sci. Univ. Tokyo, Bot. 13: 1-64. 1981. 

Small annual herbs, terrestrial or semiaquatic, 
erect to prostrate or creeping and rooting at the 
nodes, stems simple or branched, glabrous or pu- 
berulent, usually 4-angled, drying yellowish or 
brown. Leaves opposite, simple, small, sessile or 
petiolate, blades entire or denticulate, base often 
truncated to cordate, often glandular-punctate, ve- 
nation pinnate or 3-5-veined from the base. Inflo- 
rescences of solitary flowers in the axils of distal 
leaves, less often terminal or axillary racemes or 
umbel-like clusters, bracts and bracteoles absent, 
pedicels slender, usually longer than the calyx. 
Flowers small, calyx tubular with 5 short spread- 
ing lobes or deeply parted to the base with 5 se- 
pals, the sepals or lobes subequal, glabrous or pu- 
berulent externally, tube with 5 longitudinal ridg- 
es (sometimes winged); corolla tubular to cam- 
panulate, longer than the calyx (equaling the calyx 
and not opening in cleistogamous flowers), blue 
to purple or white, bilabiate, upper lip usually 
shorter and erect, entire or 2-lobed, lower lip 3- 
lobed and spreading; stamens 4 or with 2 fertile 
and 2 staminodes present, filaments often inserted 
at 2 levels in the distal half of the tube, anterior 
filaments often with a distinct spur near the base, 
upper filaments often bent just near the apex, an- 
thers free or united in pairs, 2-thecous, thecae of- 
ten widely divergent and X-shaped; ovary gla- 
brous, 2-locular, ovules many, style slender, stig- 
mas 2-lobed. Fruits dry capsules, ovoid to ellip- 
soid, globose or cylindric, dehiscing septicidally 
and 2-valved, the placentae winged; seeds many, 
oblong-elliptic, smooth to reticulate or alveolate 
with prominent transverse or longitudinal raised 
ridges. 

A genus of 50 to 70 species widespread in trop- 
ical and temperate areas throughout the world, 
with the majority of species in tropical parts of 
Africa, Asia, and Australia. The genus is classi- 
fied in the tribe Gratioleae and placed in subtribe 
Lindernieae with Torenia. It seems likely that L. 
crustacea is indigenous to Costa Rica and that the 



46 



FIELDIANA: BOTANY 



other two species have been introduced. These 
plants are generally found in marshes, at the edges 
of standing water, or in moist depressions, in open 
sunny or partly shaded sites. The preference for 



moist soils, the short slender stems, small opposite 
leaves, small bluish flowers, tubular bilabiate co- 
rollas, four or two fertile stamens, and appenda- 
ged anther filaments help distinguish the genus. 



Key to the Species of Lindernia 

la. Flowers subsessile, pedicels < 3 mm long, flowers solitary in leaf axils; corollas white or yellowish 

to pale lilac; leafy stems usually with puberulent longitudinal ridges; fruits 6-10 mm long 

L. diffusa 

Ib. Flowers borne on pedicels 3-23 mm long (longer pedicels at distal nodes), flowers often in terminal 
cyme-like groupings; corollas bluish to lilac, rose, or white marked with yellow; leafy stems gla- 
brous or puberulent; fruits 3-4 mm long 2 

2a. Calyx 5-lobed, united ca. 50% to form a tube (but splitting after anthesis and appearing as separate 
sepals in fruit); fruits ovoid, seeds 0.4-0.5 mm long; leaves petiolate L Crustacea 

2b. Calyx with 5 sepals united only at the base; fruits ellipsoid, seeds 0.3-0.4 mm long; leaves sessile 

. / diihni 



Lindernia Crustacea (L.) F. v. Muell., Syst. Cen- 
sus Aust. PL 1: 97. 1882. Capraria Crustacea 
L., Mant. PL 1: 87. 1767. Torenia Crustacea 
(L.) Cham. & Schldl., Linnaea 2: 570. 1827. 
Vandelila Crustacea (L.) Benth., Scroph. In. 35: 
1835. Pyxidaria Crustacea (L.) Kuntze, Rev. 
Gen. PL 2: 464. 1891. Figure 2. 

Prostrate or procumbent herbs to 15 cm tall, 
rarely rooting at distal nodes, leafy stems 0.2-1.3 
mm diam., longitudinally ribbed, glabrous or with 
thin crooked hairs 0.1-1 mm long. Leaves op- 
posite, petioles 1-7 mm long, 0.4-0.7 mm wide, 
glabrous or sparsely puberulent, leaf bases not 
united across the stem; leaf blades 6-16 mm long, 
4-15 mm wide, ovate to ovate-rounded or ovate- 
triangular, apex obtuse or rounded, margin serrate 
with 4-6 teeth/cm, base truncate to rounded-sub- 
cordate, drying membranaceous and yellowish 
green, minutely punctate, glabrous or very sparse- 
ly puberulent beneath and along margin, 2 veins 
3-5/side. Inflorescences solitary axillary flowers 
or terminal cymes, pedicels 4-18(-25) mm long, 
0.1-0.2 mm diam., glabrous or with few thin hairs 
to 0.4 mm long. Flowers with calyx 2-4 mm 
long, 1-1.8 mm diam., lobes 0.7-1.5 mm long, 
triangular and acute, glabrous or very sparsely pu- 
berulent; corolla 4-7 mm long, blue to violet or 
blue-purple (white), tube ca. 5 mm long, 1-1.5 
mm diam.; fertile stamens 4 in 2 pairs, anthers 
connivent; ovary glabrous. Fruits 3-4 mm long, 
2-2.5 mm wide, rounded-ovoid, central column 
(placenta) to 2 mm wide with smooth lateral 
wings and central reticulated area; seeds 0.4-0.5 



mm long, 0.3-0.4 mm diam., oblong, yellowish, 
obscurely reticulate. 

Uncommon plants of moist open sunny or part- 
ly shaded sites in lowland evergreen forest for- 
mations, 0-300(-1000) m elevation. It is also 
found along stream edges in seasonally dry areas. 
Flowering and fruiting throughout the year. This 
species is a widespread weed, now found in trop- 
ical and warm-temperate climates throughout the 
world. 

Lindernia crustaced is recognized by its small, 
mostly prostrate habit, mostly glabrous parts, 
small ovate-serrate leaves, small blue flowers on 
prominent slender pedicels, calyx lobes united for 
half their length, small rounded fruit, and unusual 
persisting placenta with smooth lateral wings. In 
addition, the lower pair of anthers is inserted on 
the lower corolla lip. The persisting and expanded 
calyx often splits along the thin intercostal areas 
to produce separate sepals, appearing very differ- 
ent from the calyx in anthesis. 

Lindernia diffusa (L.) Wettst. in Engl. & Prantl, 
Naturlichen Pflanzenfam. 4, 3b: 79. 1891. Van- 
delia diffusa L., Mant. PL 1: 89. 1767. Pyxi- 
daria diffusa (L.) Kuntze, Rev. Gen. PL 2: 464. 
1891. Figure 2. 

Prostrate or decumbent herbs 2-15 cm tall, 
sometimes forming small mats to 25 cm wide, 
often rooting at the nodes, leafy stems 0.4-1.3 
mm diam., quadrangular with 4 puberulent lon- 
gitudinal ribs, the thin whitish hairs 0.2-0.3 mm 
long. Leaves opposite, united at the base to form 



BURGER: FLORA COSTARICENSIS 



47 



an interpetiolur line or ridge, subsessile or with 
petioles to 4 mm long; leaf blades 6-25 mm long, 
4-24 nun \vide. ovate to broadly ovate or rhom- 
bic, apex obtuse to rounded, margin serrate with 
6-10 teeth/cm, base broadly obtuse to rounded 
and truncate, minutely punctate, glabrous above, 
with minute (0.1-0.2 mm) hairs along the veins 
beneath, venation subpalmate with 2-3 2 veins/ 
side. Inflorescences of solitary axillary flowers, 
usually 2/node, pedicels 1-3 mm long, ca. 0.3 mm 
diam., minutely puberulent. Flowers with calyx 
4-7 mm long, 1-1.5 mm diam., lobes 2-4 mm 
long, linear-lanceolate, glabrous or minutely pu- 
berulent on the major veins and margins of lobes; 
corolla 6-9 mm long, white, yellowish, or pale 
lilac with yellow interior; fertile stamens 4; ovary 
glabrous, style ca. 3 mm long. Fruits 6-10 mm 
long (including the 1-2 mm beak), 2.3-4 mm 
diam., ellipsoid or narrowly ovoid-ellipsoid; seeds 
0.5-0.6 mm long ca. 0.3 mm diam., oblong, with 
minute spines and a reticulate-pitted surface. 

Plants of open sunny sites in evergreen forest 
formations, 10-1200 m elevation. Probably flow- 
ering and fruiting throughout the year. Probably 
originating in the Old World, this species is now 
widespread in warmer climates. 

Lindernia diffusa is recognized by its small 
size, stems with usually four puberulent ridges, 
small subsessile serrate leaves with broad blades, 
solitary axillary flowers on very short pedicels, 
and white corollas marked with yellow. 

Lindernia dubia (L.) Pennell, Acad. Nat. Sci. 
Philadelphia Monogr. 1: 141. 1935. Gratiola 
dubia L., Sp. PI. 17. 1753. Capraria gratiolo- 
ides L., Syst. Veg. ed. 10: 1117. 1759. G. in- 
aequalis Walt., Fl. Carol. 61. 1788. G. anagal- 
lidea Michx., Fl. Bor. Am. 1: 6. 1803. Ilysan- 
thes gratioloides (L.) Benth. in DC, Prodr. 10: 
419. 1846. L. gratioloides (L.) Lloyd & Fouc., 
Fl. Quest Fr. ed. 4: 246. 1886. L. inaequalis 
(Walt.) Pennell, Torreya 19: 149. 1919. /. dubia 
(L.) Barnhart, Bull. Torrey Bot. Club. 26: 376. 
1899. L. anagallidea (Michx.) Pennell, Acad. 
Nat. Sci. Philadelphia Monogr. 1: 152. 1935. 
Figure 4. 

Herbs, stems procumbent or erect, 5-30 cm 
tall, simple or much-branched, rooting mostly at 
the base, leafy stems 0.2-1 mm diam., usually 
with 4 longitudinal ridges, glabrous. Leaves op- 
posite, sessile, free or united at the base and form- 
ing an interpetiolar line; leaf blades 4-14(-25) 
mm long, 3-8(-10) mm wide, narrowly ovate to 



narrowly ovate-triangular or lanceolate, apex 
acute to obtuse, margin entire or with 2 or 3 teeth/ 
side, base acute to somewhat rounded, drying 
membranaceous, glabrous, venation usually pal- 
mate with 3 major veins. Inflorescences of soli- 
tary axillary flowers or cymes subtended by leaf- 
like bracts, pedicels 3-23 mm long, ca. 0.2 mm 
diam., glabrous, distal pedicels much longer than 
those at lower nodes. Flowers with calyx deeply 
5-parted, sepals 1.8-2.5 mm long, 0.3-0.4 mm 
wide, linear-oblong, becoming 3-5 mm long in 
fruit but remaining narrow; corolla 5-10 mm 
long, blue or white with purplish throat or lilac to 
rose; fertile stamens 2, staminodes 2 and bifid at 
the apex; ovary glabrous, style 3 mm long. Fruits 
3-6 mm long, ca. 2 mm wide, narrowly ovoid- 
oblong to oblong-ellipsoid; seeds 0.3-0.4 mm 
long, 0.2-0.3 mm diam., oblong, with longitudinal 
rows of pits. 

Plants of open wet sites and marshes in ever- 
green or seasonally deciduous areas, 5-1100 m 
elevation. Probably flowering primarily at the end 
of the rainy season (November-January). Rarely 
encountered in Costa Rica, this species ranges 
from southern Canada and the eastern United 
States to southern South America and the West 
Indies. 

Lindernia dubia is recognized by its small stat- 
ure, moist habitats, lack of pubescence, opposite 
sessile leaves, distal flowers on longer pedicels, 
white and blue two-lipped corollas, and separate 
sepals. There is considerable diversity of leaf 
form, with some plants having narrowly ovate 
leaves (van anagallidea (Michx.) Cooperrider) 
and other plants with more elongate narrow leaves 
(var. dubia). Holmgren (Flora of the Great Plains, 
1 986, p. 769) found that L. anagallidea cannot be 
effectively separated from L. dubia. Likewise, in 
1984 D. A. Quails annotated many North Amer- 
ican collections as L. dubia. We follow their de- 
cision to consider the two as elements of the same 
species. Lindernia microcalyx Pennell & Stehle is 
very similar but has corollas 9-14 mm long; it 
has been collected along the Caribbean shore of 
Guatemala and Honduras (determinations by D. 
A. Quails, 1984). 



Lophospermum D. Don 

REFERENCES W. Elisens, Monograph of the 
Maurandyinae (Scrophulariaceae Antirrhineae). 
Syst. Bot. Monogr. 5: 1-97. 1985. D. Sutton, A 
Revision of the Tribe Antirrhineae. British Mu- 



48 



FIELDIANA: BOTANY 



seum (Natural History) & Oxford Univ. Press. 
1988. 

Perennial herbs or climbers, stems often scan- 
dent or clambering, branches often arising from a 
woody base, sparsely puberulent to glandular-vil- 
lous, drying grayish or greenish. Leaves alternate, 
petioles well developed and often bending to fa- 
cilitate climbing; leaf blades deltoid to reniform, 
apex acute to mucronate, margins dentate to 
broadly crenate, venation palmate. Inflorescences 
of solitary flowers in axils of leaves, pedicels hor- 
izontal to ascending, bracts and bracteoles absent. 
Flowers large, calyx urceolate or campanulate 
and expanded, sepals free or united at the base, 
narrowly to broadly ovate, glandular puberulent, 
often enlarging in fruit; corolla tubular-campan- 
ulate, somewhat bilabiate and open-throated, red- 
dish to violet or purple distally (whitish or pale 
near the base), 5-lobed, upper 2 lobes recurved, 
lower 3 lobes projecting forward; stamens 4, sub- 
equal or in 2 pairs, included, connective often ex- 
panded, a staminode usually present; ovary gla- 
brous or glandular puberulent, 2-locular, ovules 
many, stigma recurved or straight, forked and di- 
vergent (rarely conical and lobed). Fruits dry 
capsules, ovoid to globose, symmetric or asym- 
metric, dehiscing irregularly or poricidal, bi- 
valved; seeds rounded with a wing around the 
margin. 

A genus of 6 or 20 species, depending on 
whether the Mexican genus Rhodochiton Zucc. is 
included (following Elisens, 1985) or excluded 
(following Sutton, 1988). Species range from 
north-central Mexico to Guatemala; a few species 
are planted as ornamental climbers. These plants 
are characterized by the scandent or clambering 
habit, alternate leaves, twining petioles, deltate to 
cordiform blades, large corollas, and round seeds 
with thin peripheral wing. The one species found 
in Costa Rica was formerly placed in the genus 
Maurandya; both are placed in the tribe Antirrhi- 
neae. 

Lophospermum erubescens D. Don in Sweet, 
Brit. Fl. Card, sen 2, 1: tab. 68 and after tab. 
75. 1830. Maurandya erubescens (D. Don) 
Gray, Proc. Am. Acad. Arts 7: 377. 1868. Asar- 
ina erubescens (D. Don) Pennell, Proc. Acad. 
Nat. Sci. Philadelphia 99: 174. 1947. Figure 8. 

Climbing herbs, leafy stems 1-4 mm diam., 
pubescent with thin straight or crooked multicel- 
lular hairs 0.3-0.8 mm long, some hairs with 



gland tips. Leaves alternate, petioles 3-6 cm long, 
0.6-1.2 mm diam.. often bent or curving (some- 
times helping to support the stem), pubescent; leaf 
blades 3-7(-15) cm long, 2-6.5(-15) cm wide, 
triangular to triangular-rhombic or sagitate, apex 
acute, margin prominently crenate-serrate with 
teeth 0.5-6 mm high and 1-8 mm wide, base cor- 
date, with thin whitish hairs 0.3-0.5 mm long 
above and below, major veins 3 or 5. Inflores- 
cences of solitary axillary flowers, pedicels 2-7(- 
11) cm long, 0.7-1.4 mm diam., puberulent, not 
bent or twining. Flowers puberulent externally, 
calyx 15-24 mm long, sepals subequal, 10-14 
mm wide, broadly ovate; corolla 6-7 cm long, 
tubular-campanulate, bright pink to red distally. 
tube constricted above the base, whitish within 
and with yellow hairs in the throat, ca. 2 cm wide 
at the mouth, lobes 10-14 mm long, 12-16 mm 
wide, rounded distally. Fruits 15-20 mm long, 
subtended by persisting sepals, surface with thin 
multicellular hairs; seeds ca. 2.5 X 2.5 mm, wings 
lateral with a narrow notch at apex and truncated 
base, body of the seed ca. 1 .5 X 0.8 mm, tuber- 
culate. 

Lophospermum erubescens is characterized by 
its vining habit, twisted petioles, triangular and 
coarsely dentate leaf blades, broad sepals, and 
large, tubular, slightly asymmetric pink corollas. 
This species is native to the oak forests of the 
Sierra Madre Oriental of Mexico and is now com- 
monly grown in gardens as a climbing ornamen- 
tal. It has been collected as an escape at Monte- 
verde, where it was flowering in January and Sep- 
tember at ca. 1400 m, and north of San Isidro del 
General at 1500 m elevation, where it was flow- 
ering in April. Compare Maurandya barclaiana 
and M. scandens. 



Maurandya Ortega 

RHFERHNCKS W. Elisens, Monograph of the 
Maurandyinae (Scrophulariaceae Antirrhineae). 
Syst. Bot. Monogr. 5: 1-97. 1985. D. Sutton, A 
Revision of the Tribe Antirrhineae. British Mu- 
seum (Natural History) & Oxford Univ. Press. 
1988. 

Scandent herbs, annual or perennial, with thin 
flexible stems from a fibrous base or taproot, gla- 
brous. Leaves alternate, petioles long and often 
twisting to support the twining habit; leaf blades 
hastate to sagittate or rarely cordiform, margins 
usually entire, glabrous, venation palmate. Inflo- 



BURGER: FLORA COSTARICENSIS 



49 



rescences of solitary flowers in leaf axils, pedi- 
cels long, terete or winged near the base, glabrous 
or glandular-puberulent distally, bracts and brac- 
teoles absent. Flowers with 5 sepals united only 
at the base, equal, lanceolate, imbricate near the 
base ( urceolate), margins entire, glabrous or glan- 
dular-pubescent; corolla tubular and prominently 
2-lipped, with open or closed throat, pink to red, 
violet, or blue, often whitish near the base, gla- 
brous to glandular-puberulent externally, upper lip 
with 2 recurved lobes, lower lip with 3 recurved 
or projecting lobes, often closing the mouth with 
the well-developed palate; stamens 4 in 2 pairs, 
included, inserted near the base of the tube, fila- 
ments villous at the base; ovary 2-locular, locules 
subequal or unequal, glabrous or with glandular 
trichomes, ovules many, stigma conical and re- 
curved. Fruits dry 2-valved capsules, ovoid to 



globose, dehiscence irregular, irregular-transverse 
or poricidal; seeds rectangular with tuberculate- 
cristate surface (ovoid with a peripheral wing and 
minute tuberculate surface sculpturing in M. wis- 
lizeni), dark brown. 

As revised by Elisens (1985, cited above), 
Maurandya is a genus of four species ranging 
from the southwestern United States to central 
Mexico. Sutton (1988, cited above) recognizes 
only two species in the genus, transferring two 
species found in the southwestern United States 
to splinter genera. The genus is closely related to 
Lophospermum, and they are placed in the tribe 
Antirrhineae. Two species are likely to be found 
as garden ornamentals in Central America, and 
one has recently been collected as an escape. 
These plants have also been placed in the genus 
Asarina. 



Key to the Species of Maurandya 

la. Seeds with lateral wings; corolla to 6 cm long (see Lophospermum erubescens) 

Ib. Seeds without wings; corolla to 4 cm long 2 

2a. Calyx covered with short gland-tipped hairs; corolla usually blue-violet M. barclaiana 

2b. Calyx glabrous or with few gland-tipped hairs; corolla usually pink M. scandens 



Maurandya barclaiana Lindley, Hot. Reg. 13, 
tab. 1108. 1827. Figure 8. 

Vines with slender herbaceous climbing stems 
and twisting petioles, leafy stems 0.6-1 mm diam., 
glabrous or minutely papillate-puberulent at the 
nodes. Leaves alternate, petioles 12-32 mm long, 
0.3-0.5 mm diam., glabrous, often coiling around 
objects for support; leaf blades 16-24(-35) mm 
long, 14-35 mm wide, triangular-hastate or sagit- 
tate, apex acute, usually with 2 basal lobes, gla- 
brous, venation subpalmate with 3 (5) major veins 
from the base. Inflorescences of solitary flowers 
in leaf axils, pedicels 15-55 mm, ca. 0.5 mm 
diam., straight or curved, glabrous except near the 
calyx. Flowers with calyx 8-16 mm long, 2-3 mm 
wide at base, lobes triangular with long narrow 
apex, covered with gland-tipped hairs 0.3-0.5 mm 
long; corolla 2.5-4 cm long, ca. 15 mm wide, 
blue- violet, lobes 6-10 mm long, filaments ca. 18 
and 14 mm long. Fruits 11-17 mm long, ovoid; 
seeds with angular projections. 

Maurandya barclaiana is a vining ornamental 
often planted in tropical gardens. It is recognized 
by its colorful tubular two-lipped corollas, glan- 
dular-puberulent calyx, often sagittate leaves, and 
twisting petioles. 



Maurandya scandens (Cav.) Pers., Syn. PI. 2: 
160. 1806. Usteria scandens Cav., Icon. 2: 15, 
tab. 116. 1793. Maurandya semperflorens Or- 
tega, Nov. PI. Descr. Dec. 21. 1797. Reichardia 
scandens (Cav.) Roth, Catal. bot. 2: 65. 1800. 
Asarina scandens (Cav.) Pennell, Proc. Acad. 
Nat. Sci. Philadelphia 99: 175. 1947. Figure 8. 

Herbaceous climbers with slender twining 
stems, often with adventitious roots, leafy stems 
0.5-2 mm diam., glabrous. Leaves alternate, pet- 
ioles 8-35(-42) mm long, 0.4-0.8 mm diam., gla- 
brous, often bent or twisted along their length; 
leaf blades 2-5(-6) cm long, l-3.5(-4.5) cm 
wide, sagittate to hastate or triangular, apex acute, 
margin entire, base cordate and usually with 
prominent lateral lobes, drying membranaceous, 
glabrous or minutely papillate-puberulent near the 
petiole attachment, major veins 5 or 7, the 3 cen- 
tral veins reaching the middle of the blade. Inflo- 
rescences of solitary axillary flowers, pedicels 2- 
6(-8.5) cm long, 0.4-1 mm diam., slightly thick- 
ened near the base, glabrous. Flowers with calyx 
9-15 mm long, sepals 2-4 mm wide near the 
base, lanceolate, glabrous (rarely sparsely glan- 
dular puberulent); corolla to 4 cm long, tubular- 



50 



FIELDIANA: BOTANY 



campanulate, pink to pale purple, tube 2-3 cm 
long, lobes 6-10 mm long, rounded; filaments 16 
and 12 mm long, lower filaments shorter; ovary 
glabrous or with glandular hairs near apex. Fruits 
9-12 mm long, ovoid to oblong, locules subequal; 
seeds with rounded projections. 

Maurandya scandens, a native of central Mex- 
ico, is planted as an ornamental climber in Central 
America. The thin twining stems, lack of pubes- 
cence, often twisted petioles, sagittate-triangular 
leaves, solitary axillary flowers, and tubular two- 
lipped pinkish corollas make the plants distinc- 
tive. 



Mazus Loureiro 

Small annual herbs, diffusely branched from a 
basal rosette, stems with longitudinal ridges, pu- 
bescent or subglabrous. Leaves opposite and 
crowded near the base, alternate distally, petioles 
poorly differentiated; leaf blades obovate with 
crenulate or coarsely dentate margins, venation 
pinnate. Inflorescences terminal racemes, flowers 
alternate and solitary along the rachis (often on 1 
side), bracts minute or absent, pedicels well de- 
veloped, lacking bracteoles. Flowers with cam- 
panulate or tubular calyx, 5-lobed, tube usually 
equaling the lobes in length, lobes equal in size 
and shape, glabrous or puberulent, slightly ex- 
panded and enclosing the fruit; corolla 2-lipped, 
light blue to white or lavender, tube short, upper 
lip 2-lobed, lower lip 3-lobed, larger than the up- 
per and spreading; stamens 4 in 2 pairs, filaments 
inserted at the base of the tube, anthers divaricate, 
thecae contiguous, a staminode absent; ovary 2- 
locular, ovules many, style longer than the ovary, 
stigma broadly bilabiate. Fruits capsules, globose 
or slightly compressed, loculicidal, producing 2 
entire valves; seeds, ovoid to oblong or angled, 
rugulose or reticulate, black. 

Mazus is an Asian-Australian genus of 30 to 40 
species that is placed in the tribe Dodartieae with 
Mimulus and Leucocarpus. The following species 
is now an occasional weed in temperate and trop- 
ical montane climates. 

Mazus pumilus (Burm.f.) Steenis, Nova Guinea 
N.S. 9: 31. 1958. Lobelia pumila Burm.f., Fl. 
Ind. 186, t. 60, f. 3. 1768. M. japonicus 
(Thunb.) Kuntze, Rev. Gen. PI. 1: 462. 1891. 
Lindernia japonica Thunb., Fl. Jap. 253. 1784. 
M. rugosus Lour., Fl. Cochinch. 385. 1790. 



Annual herbs 3-15 cm tall, ascending or de- 
cumbent, sometimes forming small mats, stems to 
20 cm long, leafy stems 0.8-1.5 mm diam., 
sparsely puberulent with hairs to 0.5 mm long. 
Leaves rosulate or crowded at the base, opposite 
leaf bases united across the stem, distal leaves 
smaller and alternate, petioles 2-15 mm long, 
with lateral margins not clearly differentiated 
from the blades; leaf blades 6-60 mm long, 4- 
20 mm wide, obovate to cuneate-oblong or oblan- 
ceolate, apex obtuse or rounded, margins of larger 
leaves with prominent lobes 0.5-3 mm long (sep- 
arated by wide sinuses), base cuneate and decur- 
rent, sparsely puberulent with hairs to 0.4 mm 
long, 2 veins 2 or 3/side. Inflorescences 3-8 cm 
long, racemes with 3-13 flowers separated by in- 
ternodes to 12 mm long, bracts 1-2 mm long, lin- 
ear-filiform, pedicels 2-6(-l 1) mm long, sparsely 
to densely papillate puberulent. Flowers with ca- 
lyx 4-7 mm long, with 5 prominent veins, tube 
1-2 mm diam., lobes 2-3 mm long, narrowly tri- 
angular or oblong, with 5 prominent longitudinal 
veins, glabrous or minutely papillate puberulent; 
corolla 7-10 mm long, pale violet and white, tu- 
bular, upper lobes erect, lower lip ca. 2 mm long 
with 2 raised longitudinal ridges adaxially, palate 
yellow or whitish; stamens included; stigma lobes 
closing after pollination. Fruits 2-4 mm long, 2- 
3 mm diam., obovoid and bisulcate, surface gla- 
brous, smooth, enclosed within the persisting ca- 
lyx cup; seeds oblong, minutely reticulate. 

Mazus pumilus, native to eastern Asia, is rarely 
collected in the Neotropics. It is often found as a 
weed in gardens and open places (1000-2000 m). 
The species is recognized by its small size, nar- 
rowly obovate opposite leaves with crenulate mar- 
gins and long-decurrent blades, few-flowered ra- 
cemes with long pedicels, partly united calyx, and 
blue or pink two-lipped corolla. These plants may 
resemble species of Veronica, but that genus lacks 
the strongly two-lipped flowers and four stamens. 



Mecardonia Ruiz & Pav6n 

RHFHRKNCE F. Pennell, Reconsideration of the 
Bacopa-Herpestis problem. Proc. Acad. Nat. Sci. 
Philadelphia 98: 83-98. 1946. R. Rossow, Revi- 
si6n del gnero Mecardonia (Scrophulariaceae). 
Candollea 42: 431-474. 1987. 

Erect or prostrate herbs, annual or perennial, 
stems with few to many branches, mostly gla- 
brous, drying brown or blackish. Leaves opposite, 



BURGER: FLORA COSTARICENSIS 



51 



sessile or short-petiolate, blades with serrate mar- 
gins, usually glabrous, glandular-punctate, vena- 
tion pinnate with most veins diverging in the 
proximal half of the blade. Inflorescences of sol- 
itary axillary flowers (2/node), 2 small linear 
bracts subtending the pedicel, bracteoles absent at 
the base of the calyx, pedicels slender. Flowers 
with deeply 5-parted calyx, sepals unequal in 
width with 3 broad outer ones; corolla tubular- 
campanulate, slightly 2-lipped and sometimes 
with a prominent palate (personate), yellow or 
white, upper lip slightly 2-lobed, lower lip 3- 
lobed; stamens 4 in 2 pairs, inserted near the base 
of the tube, thecae stipitate on short arms of the 
connective, a short staminode sometimes present; 
ovary 2-locular, ovules many, stigma flattened, 2- 
lobed, slightly bent. Fruits thin-walled capsules, 
dehiscence loculicidal; seeds oblong, surface re- 
ticulate and ridged. 

A tropical and warm-temperate American ge- 
nus of ca. 15 species, the majority in South Amer- 
ican. The species have sometimes been united 
with those of Bacopa, but Pennell (1946) sepa- 
rated them because of the stipitate anther-thecae 
and the yellow pigmentation of the corolla. The 
anthers, the gland-dotted foliage, the slightly bi- 
lobed deflexed stigma, and the loculicidal capsule 
strengthen the distinction and indicate a close re- 
lationship with Stemodia. In Costa Rica, Mecar- 
donia differs from Stemodia in having yellow co- 
rollas and pedicels with small bracts at the base. 
However, these distinctions do not hold over the 
entire range of all species, and it may be neces- 
sary to reevaluate the genera. 

Mecardonia procumbens (Mill.) Small, Fl. S.E. 
U.S. 1065 & 1338. 1903. Erinus procumbens 
Mill., Card. Diet. ed. 8. 1768. Lindernia dianth- 
era Sw., Prodr. 92. 1788. Herpestis caparioides 
Kunth i/i H.B.K., Nov. Gen. Sp. 2: 368. 1818. 
H. peduncularis Benth., Bot. Mag. 2: 56. 1836. 
Bacopa procumbens (Mill.) Greenman, Publ. 
Field Columb. Mus. Bot. Ser. 2: 261. 1907. Fig- 
ure 2. 

Herbs, prostrate to procumbent, stems to 40 cm 
long, distal stems few-branched, sometimes root- 
ing at the proximal nodes, leafy stems 0.4-2 mm 
diam., glabrous, with 4 longitudinal ridges. 
Leaves opposite, subsessile or with poorly de- 
fined petioles to 4 mm long; leaf blades 6-24 mm 
long, 4-16 mm wide, ovate-elliptic to broadly or 
narrowly elliptic, apex obtuse, margin with 6-9 
teeth/cm, base obtuse to cuneate, drying charta- 



ceous and often blackish, glabrous, 2 veins 2 or 
3/side. Inflorescences of solitary axillary flowers, 
linear bracts ca. 2 mm long at base of pedicel, 
pedicels 2-16(-24) mm long, 0.2-0.4 mm diam., 
glabrous. Flowers glabrous externally, calyx 5-9 
mm long, outer 3 sepals 2.7-4 mm wide, ovate to 
lanceolate; corolla 8-10 mm long, tube ca. 7 mm 
long, yellow, purplish in the throat, barbate in the 
mouth, lobes 1-2 mm long; stamens 4, filaments 
borne on the lower half of the tube, a staminode 
sometimes present. Fruits 4-6 mm long, ovoid, 
dehiscent from the apex, placenta spongy; seeds 
0.4-0.5 mm long, ca. 0.2 mm diam., oblong or 
ovoid, minutely reticulate, light brown. 

Plants of open sunny or partly shaded sites on 
wet soils or wet sand in evergreen or deciduous 
areas, 0-1200(-2400) m elevation. Flowering and 
fruiting throughout the year. The species is partic- 
ularly common in areas supporting evergreen low- 
land rain forest formations. The species ranges 
from Mexico to Uruguay and Argentina. 

Mecardonia procumbens is recognized by its 
short herbaceous stems, general lack of pubes- 
cence, subsessile opposite serrate leaves, solitary 
axillary flowers on prominent pedicels, calyx with 
wide outer sepals and narrow inner sepals, and 
yellow tubular two-lipped corolla. These plants 
resemble species of Bacopa, Lindernia, and Ste- 
modia, both in growth habit and in preference for 
moist open sites. This species can be separated 
from those in the other genera by the bright yel- 
low corolla, glabrous gland-dotted foliage and 
bracteolate pedicels. This species is very closely 
related to M. montevidensis (Spreng.) Kuntze of 
southern South America. 



Micranthemum Michaux 

REFERENCE L. O. Williams, Tropical Ameri- 
can plants XII. Fieldiana, Bot. 34: 101-132. 1972 
(Micranthemum pp. 122-124). 

Small slender stemmed herbs, aquatic or grow- 
ing in wet places, diffuse and much-branched, 
nodes often with adventitous roots. Leaves op- 
posite, sessile, small, blades with entire margins, 
thin, usually glabrous, venation palmate. Inflores- 
cences of solitary (rarely 2) axillary flowers, usu- 
ally with 1 flower/node, bracts and bracteoles ab- 
sent, pedicels short. Flowers minute, calyx with 
4 or 5 lobes or 4 or 5 sepals, lobes equal, tube 
short or absent, glabrous or sparsely puberulent; 
corolla 2-lipped, tube very short, adaxial lip short 



52 



FIELDIANA: BOTANY 



and 2-lobed, abaxial lip prominent and 3-lobed; 
stamens 2, filaments inserted at the mouth of the 
tube abaxially, anthers 2-thecous, staminodes ab- 
sent; ovary 1- or 2-locular, ovules many, style 
short, stigma 2-lobed. Fruits dry capsules, glo- 
bose or ovoid, splitting irregularly; seeds oblong, 
with longitudinal ridges. 

Micranthemum is a genus of perhaps 10 to 12 
species, ranging from the southeastern United 
States to southern South America. It is most di- 
verse in Cuba. The genus is recognized by its 
small slender-stemmed aquatic or semiaquatic 
habit, small sessile rounded leaves, minute soli- 
tary flowers, and small two-lipped corolla with 
two distal stamens. These delicate little plants are 
sometimes grown in freshwater aquaria. Although 
rarely collected in Central America, it is repre- 
sented by the following species in Costa Rica. 

Micranthemum umbrosum (J. F. Gmelin) Blake, 
Rhodora 17: 131. 1915. Anonymous umbrosa 
Walter, Fl. Carol. 63. 1788. Globifera umbro- 
sum (Walter) J. F. Gmelin, Syst. Nat. 32. 1791. 
Figure 1. 

Herbs, aquatic or in wet sites, creeping or often 
forming small mats, stems to ca. 1 2 cm long, leafy 
stems 0.2-0.8 mm diam., glabrous, with 2 longi- 
tudinal ridges. Leaves sessile, base of opposing 
leaves often forming a line or ridge across the 
stem; leaf blades 3-8 mm long, 2-7 mm wide, 
ovate-orbicular to rounded-obovate, apex obtuse 
to rounded, margins entire, obtuse or rounded at 
the contracted base, drying greenish and translu- 
cent, glabrous, with 3 major veins and 2 lateral 
veins. Inflorescences of solitary axillary flowers, 
usually with 1 flower/node, pedicels 0.5-1 mm 
long, 0.2-0.4 mm diam., glabrous. Flowers with 
calyx 0.8-1.3 mm long, sepals 4, equal and sep- 
arate nearly to the base, narrowly oblong, with 
few minute hairs or glabrous; corolla ca. 1 .5 mm 
long, white, tube very short; stamens 2. attached 
at the base of the abaxial sinuses. Fruits ca. 1 
mm diam., globose, thin-walled, 1-locular; seeds 
0.3-0.4 mm long, ca. 0.2 mm diam., oblong, yel- 
lowish, with minutely ribbed longitudinal ridges 
(50X). 

Micranthemum umbrosum, ranging from the 
southeastern United States to South America, is 
rarely collected in Central America, but the very 
small size of the plants and their aquatic or wet 
habitats may have resulted in the species being 
overlooked. The species has been collected in 
Honduras, Nicaragua, and the Tortuguero region 
of northeastern Costa Rica. The slender glabrous 



stems, small opposite .sessile rounded leaves, mi- 
nute solitary flowers, white two-lipped corolla 
with two stamens, and thin-walled capsules with 
one locule help distinguish these plants. Micran- 
themum standleyi Williams, an endemic of Gua- 
temala, has smaller flowers, ovoid fruits, and pi- 
lose calyx lobes. Micranthemum pilosum Ernst, 
from Venezuela, is also similar, but it is possible 
that these are only variants of M. umbrosum in a 
broad sense. 



Mimulus Linnaeus 

Ri H:RKNCK A. Grant, A monograph of the ge- 
nus Mimulus. Ann. Missouri Bot. Gard. 11: 98- 
388. 1924. 

Annual or perennial herbs (rarely shrubs), of- 
ten preferring moist sites, decumbent or erect, 
with viscid pubescence or glabrous. Leaves op- 
posite, simple, sessile or petiolate, blades entire or 
dentate, venation pinnate or palmate. Inflores- 
cences of solitary flowers in leaf axils or forming 
terminal racemes, bracts absent or leaf-like, ped- 
icels without bracteoles at the apex. Flowers 
showy, calyx tubular to campanulate, almost as 
long as the corolla, with 5 prominent veins, tube 
longer than the 5 short lobes or teeth, lobes usu- 
ally unequal; corolla tubular or narrowly campan- 
ulate, 2-lipped, lips shorter than the tube, adaxial 
lip 2-lobed, abaxial lip 3-lobed. lobes subequal. 
usually with 2 yellowish protuberances in the 
throat; stamens 4 in 2 unequal pairs, included or 
exserted, filaments attached near the base of the 
tube, linear, anthers 2-thecous with the thecae di- 
varicate and confluent at the apex; ovary 2-locu- 
lar, ovules many, style included, stigma flattened 
and 2-lobed. Fruits capsules, oblong to linear, 
thin- or thick-walled, dehiscence loculicidal, en- 
closed within the persisting and enlarged calyx 
tube; seeds small, smooth or reticulate. 

Mimulus is a genus of ca. 150 species ranging 
from the Americas to Africa and Asia but with 
most of the species in western North America. 
Colorful species (monkey flowers) are grown in 
temperate gardens. No specimens of this genus 
have been collected in Costa Rica, but the follow- 
ing species may occur. 



Mimulus glabratus Kunth in H.B.K., Nov. Gen. 
Sp. 2: ed. quarto 370. 1818. Figure 1. 

Perennial herbs, procumbent or prostrate, 4-40 
cm long, rooting at many lower nodes, leafy stems 



BURGER: FLORA COSTARICENSIS 



53 



0.3-4 mm diam. (dried), succulent in life, gla- 
brous, nodes with an interpetiolar line formed by 
the clasping leaf bases, rooting at lower nodes. 
Leaves opposite, subsessile near the apex of 
stems or with petioles 2-18 mm long, 0.4-2 mm 
wide, with lateral margins continuous with the 
blade margins; leaf blades 8-45 mm long, 4-35 
mm wide, ovate to ovate-oblong or ovate-orbic- 
ular, apex obtuse to rounded, margins with 3-5 
prominent teeth/cm, base obtuse or truncated, dry- 
ing grayish green, glabrous, venation palmate 
with 3-5 major veins. Inflorescences of solitary 
flowers axillary to foliage leaves (2/node), pedi- 
cels 12-42 mm long, 0.4-0.8 mm diam., gla- 
brous. Flowers with calyx 5-10 mm long (14 mm 
in fruit), 2.5-4 mm diam., tubular, upper lobe 
larger (ca. 4 mm) than lower lobes (ca. 2 mm), 
glabrous; corolla 7-18 mm long, yellow with red 
spots within the throat, tube 2.5-4 mm diam., 
lobes 1-2 mm long. Fruits 6-10 mm long, 5-6 
mm diam., included within the calyx tube; seeds 
0.4-0.5 mm long, ca. 0.3 mm diam., oblong, 
brown, with poorly defined longitudinal ridges. 

Mimulus glabratus ranges from the southern 
United States and Mexico to Nicaragua and from 
Colombia to Chile. In tropical environments it 
grows in wet sites at 100-2200 m elevation. 
These plants are recognized by their wet habitat, 
creeping stems with adventitous roots, lack of pu- 
bescence, opposite petiolate leaves, solitary axil- 
lary flowers on slender pedicels, tubular-plicate 
calyx with unequal lobes, and yellow two-lipped 
corolla with short lobes. Although not collected 
between Nicaragua and Colombia, this species 
may occur in our area. 



Penstemon Mitchell 

REFERENCE R. Straw, The penstemons of 
Mexico. II. Penstemon hartwegii, Penstemon gen- 
tianoides, and their allies. Bol. Soc. Bot. Mexico 
27: 1-25. 1962. 

Perennial herbs or subshrubs, mostly erect, 
stems simple or branching from near the base, 
glabrous or pubescent. Leaves opposite, the lower 
leaves often petiolate, distal leaves often sessile 
and clasping the stem or bract-like; leaf blades 
serrate or entire, acute to rounded, venation pin- 
nate. Inflorescences terminal panicles or thyrses 
(often racemiform), the partial inflorescences usu- 
ally cymose, pedunculate with dichotomous 
branching, subtended by leaf-like or small bracts, 



glabrous or puberulent, bracteoles present or ab- 
sent. Flowers small or large, calyx deeply 5-part- 
ed, sepals imbricate in bud, equal or subequal; 
corolla tubular to tubular-campanulate, often nar- 
rowed at the base and abruptly expanded, 2- 
lipped, mostly blue to purple, sometimes red or 
white, upper (adaxial) lip 2-lobed and often erect, 
lower (abaxial) lip spreading or reflexed and 3- 
lobed; stamens 4 in 2 pairs, shorter than the co- 
rolla, bases glandular and nectariferous, filaments 
attached near the base of the tube, anthers gla- 
brous or villous, thecae distinct or confluent, a 
staminode present and conspicuous; ovary ovoid, 
2-locular, ovules many, style slender, stigma 
slightly 2-lobed or subcapitate. Fruits dry cap- 
sules, ovoid to globose, dehiscence septicidal and 
often loculicidal, valves entire or bifid; seeds 
many, dark, angular, rugulose to smooth, rarely 
winged along the margins. 

Penstemon is one of the largest genera of Scro- 
phulariaceae, with ca. 300 species ranging from 
temperate North America into the mountains of 
Mexico and Guatemala. The greatest number of 
species are found in the mountains of the western 
United States. The genus is identified by the tu- 
bular to campanulate flowers with a conspicuous 
staminode oriented on the lower (abaxial) side of 
the tube. Several species are colorful ornamentals, 
grown widely in temperate or tropical montane 
gardens. The following species is likely to be seen 
in Costa Rican gardens. 

Penstemon gentianoides (Kunth in H.B.K.) Poir- 
et, Diet. Sci. Nat. 38: 385. 1825. Chelone gen- 
tianoides Kunth in H.B.K., Nov. Gen. Sp. 2: 
363. tab. 172. 1818. P. skutchii Straw, Bol. Soc. 
Bot. Mexico 27: 13. 1962. Figure 8. 

Erect perennial herbs or subshrubs 0.6-1.5 m 
tall, branched mostly in the lower half, leafy 
stems 2-6 mm diam., terete, glabrous or with thin 
straight hairs 0.1-0.2 mm long, nodes with inter- 
petiolar ridges. Leaves opposite (but often with 
axillary short-shoots and giving a verticillate ap- 
pearance), sessile or subsessile; leaf blades 4-12 
cm long, 5-20 mm wide, lanceolate to linear-lan- 
ceolate, apex acute, margin entire, base cuneate to 
rounded, drying stiffly chartaceous, usually gla- 
brous (except at the base), punctate above and be- 
low, 2 veins often obscure. Inflorescences 10- 
35 cm long, racemiform thyrses with axillary 
groups of (l-)3-9 flowers, bracts (reduced leaves) 
1 2-30 mm long, usually lanceolate, peduncles 4- 
12 mm long, often terminated by opposite brac- 



54 



FffiLDIANA: BOTANY 



teoles 3-8 mm long, pedicels 3-12 mm long, mi- 
nutely puberulent or subglabrous. Flowers with 
calyx 6-10 mm long, sepals 3-5 mm wide, broad- 
ly imbricate, glabrous or puberulent; corolla 2.5- 
4 cm long, tube 7-10 mm diam., narrowed near 
the base, deep purple to rose, red. or white, gla- 
brous or minutely puberulent, lobes 6-10 mm 
long, throat white; filaments glabrous, anthers in- 
cluded, staminode ca. 22 mm long; ovary 5-7 mm 
long. Fruits 8-12 mm long, 5-6 mm wide, ovoid, 
style base persisting, septicidal and loculicidal; 
seeds black, angular, rugulose. 

Penstemon gentianoides, native to the higher 
(2400-4000 m) mountains of Mexico and Gua- 
temala, is sometimes cultivated in gardens of the 
Meseta Central and at higher elevations. The spi- 
cate or racemiform arrangement of verticillate 
groups of large colorful flowers, with tubular co- 
rollas slightly expanded above the base, and the 
prominent staminode help to distinguish these 
plants. 



Russelia Jacquin 

REFERENCE M. Carlson, Monograph of the ge- 
nus Russelia. Fieldiana, Bot. 29: 231-292. 1957. 

Perennial shrubs or herbaceous subshrubs from 
a woody base, erect or with pendent or scandent 
stems, distal branches few to many, stems angular 
or terete, puberulent or glabrous. Leaves opposite 
or whorled (caducous in R. equisetiformis), sub- 



sessile or short-petiolate, blades usually ovate to 
lanceolate, margin entire or dentate to deeply in- 
cised, glandular (resinous) peltate scales (dots) of- 
ten present, venation pinnate. Inflorescences di- 
chotomous cymes, 1-3 in leaf axils or subtended 
by reduced bract-like leaves, often verticillate. 
subsessile to long pedunculate, linear bracts sub- 
tending the slender pedicels, bracteoles absent at 
the base of the calyx. Flowers mostly small (1-2 
cm), calyx deeply 5-lobed or with 5 sepals, ovate 
to lanceolate, imbricate in bud, with or without 
peltate glands; corolla tubular to funnelform, 
slightly 2-lipped. red to pink (white), the 5 lobes 
slightly unequal, the 2 (upper) adaxial lobes out- 
side in bud, lower lip 3-lobed; stamens 4 in 2 
unequal pairs, included, filaments borne in the 
lower half of the tube, thecae divaricate and con- 
fluent, a short staminode usually present; ovary 2- 
locular, ovules many, style slender, stigma slightly 
thickened. Fruits dry capsules, ovoid to subglo- 
bose, glabrous, dehiscence septicidal (secondarily 
loculicidal); seeds small, slightly rugulose, devel- 
oping among translucent fragmenting hairs (ela- 
ters) produced by the placenta. 

Russelia is a genus of 51 species centered in 
Mexico and Guatemala, with one species reaching 
Cuba and Colombia. The genus has been placed 
in its own tribe because of the unusual hair-like 
fragments in the capsule. The small tubular red 
flowers (only slightly two-lipped) in axillary 
cymes and the peltate glands (when present) also 
help to distinguish members of this genus. Two 
species are found in Costa Rica. 



Key to the Species of Russelia 

la. Common wild plants of seasonally dry habitats; leaves ovate, subsessile, 1-9 cm long; stems with 
opposite branches; inflorescences > 2-flowered; corollas 10-16 mm long R. sarmentosa 

Ib. Plants grown in gardens; leaves falling off early and stems and verticillate branches leafless; inflo- 
rescences 2-flowered; corollas 15-30 mm long R. equisetiformis 



Russelia equisetiformis Schltdl. & Cham., Lin- 
naea 6: 377. 1831. R. juncea Zucc., Flora 15, 
Beibl. 2: 99. 1832. 

Erect subshrubs to 1 m tall, much-branched, 
lower nodes with whorls of 4-8 branches, distal 
nodes with 2 branches or peduncles at each node, 
longitudinal ridges 2, 4, or 6, distal stems usually 
leafless, 0.7-6 mm diam., glabrous or minutely 
puberulent near the nodes with thin hairs 0.1-0.3 
mm long. Leaves opposite or verticillate, usually 



caducous, and absent at distal stems, petioles 3- 
4 mm long and often remaining appressed to the 
stems when blades fall; lower larger leaf blades 
8-15 mm long. 6-9 mm wide, ovate to elliptic. 
Inflorescences 2-16 cm long, often resembling 
open racemose panicles, usually with 2-4 pedun- 
cles/node, each peduncle with 1 or 2 flowers, bas- 
al peduncles 0.5-1.5 mm diam., glabrous, with 2 
or 4 prominent longitudinal ridges, bracteoles 1- 
2 mm long, linear, pedicels 6-15 mm long, 0.2- 
0.3 mm diam. Flowers glabrous externally, calyx 



BURGER: FLORA COSTARICENSIS 



55 



2-3 mm long, imbricate, acute or acuminate, gla- 
brous; corolla 15-30 mm long, 2-4 mm diam., 
tubular-funnelform, bright red, glabrous within; 
stamens 18 and 20 mm long, anthers near mouth 
of the tube, staminode present; ovary 2 mm long, 
ovoid, style 15 mm long, stigma minute. Fruits 
3-6 mm diam., globose; seeds light brown, 
among white hairs within the capsule. 

Russelia equisetiformis is a popular garden 
plant because of its slender green leafless stems 
and bright red flowers. It is distinctive because of 
the ridged stems and verticillate arching branches, 
which resemble those of Equisetum. Common 
names in Central America are coral and lluvia de 
coral. The species is native to Mexico and is an 
occasional roadside escape in the Meseta Central. 

Russelia sarmentosa Jacq., Enum. PI. Syst. 6: 25. 
1760. R. colombiana Pennell, Proc. Acad. Nat. 
Sci. Philadelphia 72: 186. 1920. R. flavoviridis 
Blake, Contrib. U.S. Natl. Herb. 24: 22. 1922. 
R. tabacensis Lundell, Contrib. Univ. Michigan 
Herb. 6: 59. 1941. R. oxyphylla Lundell, loc. 
cit. 7: 51. 1942. Figure 5. 

Herbs or weak-stemmed shrubs 0.5-2 m tall, 
erect or spreading, base woody, distal flowering 
stems usually unbranched, leafy stems 1-5 mm 
diam., strongly 4- or 6-angled with raised longi- 
tudinal ridges, puberulent with thin whitish hairs 
0.1-0.3 mm long or glabrous (except at the 
nodes). Leaves 2 or 3 at each node, subsessile 
with petioles 0.5-4 mm long, often forming an 
interpetiolar line across the stem, lateral margins 
puberulent; leaf blades l-7(-9) cm long, l-5(-6) 
cm wide, ovate to broadly ovate or ovate-trian- 
gular, apex acute, margin with 2-5 teeth/cm, teeth 
0.5-2 mm long, obtuse or truncated at the base, 
drying dark or grayish, glabrous or puberulent 
above, usually more pubescent beneath with hairs 
0.2-0.4 mm long, peltate glands (ca. 0.2 mm 
diam.) often present on both surfaces, 2 veins 3- 
5/side and strongly ascending. Inflorescences 2- 
3 cm long, congested verticels of cymes in distal 
leaf axils, 12-20 flowers/cyme, peduncles 2-18 
mm long, resembling the stems, bracts 3-4 mm 
long, linear, pedicels 5-8 mm long. Flowers with 
calyx 3-4 mm long, ca. 1.3 mm diam., lobes 2- 
3 mm long, with thin lateral margins and acumi- 
nate apex, often with peltate glands; corolla 10- 
14(-16) mm long, 1.5-3 mm diam., tubular, deep 
red, puberulent within, lobes subequal, 1 .3-3 mm 
long; stamens ca. 8 and 6 mm long, inserted near 
the middle of the tube, anthers white; ovary 1-2 



mm long, style 4-8 mm long. Fruits 4-5 mm 
long (not including the 0.5-4 mm style base), 4- 
5 mm diam., rounded-ovoid to subglobose, drying 
dark; seeds 0.4-0.5 mm long, ca. 0.2 mm wide, 
dark, among yellowish hairs. 

Common plants of open weedy sites and forest 
edges in deciduous and partly deciduous (rarely 
evergreen) forest formations, 0-1300 m elevation. 
Flowering and fruiting throughout the year but 
collected most often in July-August and Decem- 
ber-January. The species ranges from Mexico to 
Cuba and Colombia. 

Russelia sarmentosa is recognized by the often 
shrub-like habit, stiff few-branched distal stems 
with two or three subsessile leaves at each node, 
short cymes often forming compact verticels of 
flowers and fruits, and bright red tubular corollas. 
The prominent longitudinal ridges on stems, pel- 
tate glands on leaf surfaces and calyx (when pre- 
sent), and sepals broadly overlapping at the base 
are also helpful in recognizing this common spe- 
cies. We agree with Margery Carlson's delimita- 
tion (1957, cited above) of this wide-ranging and 
variable species, but we do not believe that the 
recognition of forms and varieties is useful. The 
common name is coralillo. 



Schistophragma Bentham ex Endlicher 

Small annual herbs, erect or procumbent, stems 
quadrangular and with longitudinal ridges, gla- 
brous or sparsely minutely puberulent, drying yel- 
lowish brown. Leaves opposite, petiolate, blades 
deeply pinnatisect with few opposite or alternate 
narrow lobes (smaller leaves sometimes linear-ob- 
lanceolate), margins entire or with a few distal 
teeth, venation pinnate. Inflorescences of solitary 
flowers axillary to distal leaves, bracts and brac- 
teoles absent, pedicels slender. Flowers small, ca- 
lyx divided to near the base, sepals or calyx lobes 
5, narrow, acute, subequal; corolla tubular and 
slightly 2-lipped, upper (adaxial) lip with 1 emar- 
ginate lobe, lower (abaxial) lip 3-lobed; stamens 
4, in 2 subequal pairs, included; ovary narrowly 
ellipsoid, 2-locular, ovules many, style slender, di- 
lated at apex. Fruits linear capsules, somewhat 
compressed and bisulcate, septicidal; seeds ob- 
long, surface with compressed spiral ridges. 

The delimitation of Schistophragma and its dis- 
tinction from Leucospora Nuttall and Conobea 
Aubl. is the subject of disagreement. B. L. Turner 
(Turner & Cowan, Phytologia 74: 61-103, 1993) 
recognizes three species of Schistophragma and 



56 



FIELDIANA: BOTANY 



includes Leucospora within Stemodia. D. Keil 
(pers. comm.) includes the species of Schisto- 
phragma in his concept of Leucospora, which he 
retains a.s distinct from Stemodia. Schistophragma 
is placed in the tribe Gratioleae, subtribe Stemo- 
diinae. The spirally striate seeds are very unusual 
in the Scrophulariaceae, but seeds of this type are 
sometimes found in the Gesneriaceae. The small 
size of the plants, narrow or pinnatisect leaves, 
small tubular flowers, and very narrow capsules 
make these plants distinctive. The linear fruits re- 
semble those of some Brassicaceae. Among 
American Scrophulariaceae, only Mabrya Elisens, 
from Mexico, has similar linear fruits. 

Schistophragma mexicana Benth. ex D. Dietr, 
Syn. PI. 3: 513. 1842. S. pusilla Benth. in DC, 
Prodr. 10: 392. 1846. Conobea pusilla (Benth.) 
Benth. & Hook, ex Jackson, Index Kew 1, fasc. 
1: 596. 1893. Stemodia siliguosa Sesse" & Mo- 
cino, PI. Nov. Hisp. ed. 1: 98. 1887-1890, ed. 
2: 91. 1893. Figure 4. 

Erect to decumbent annual herbs 6-25 cm tall, 
with few or many branches, leafy stems 0.3-1 mm 
diam., with 4 longitudinal ridges (quadrangular in 
cross-section), glabrous or with few minute hairs, 
nodes lacking interpetiolar lines. Leaves opposite, 
deeply pinnatifid, petioles 2-7 mm long (to first 
lobe), undifferentiated from the narrow rachis, 0.2- 
0.5 mm wide, sulcate adaxially; leaf blades 4-18 
mm long, to 12 mm wide, deeply pinnatisect with 
3-7 narrow lobes (including the distal extension of 
the rachis), lobes 0.3-2 mm wide, entire or the 
larger with 1 or 2 lateral teeth, glabrous or mi- 
nutely puberulent on the veins beneath, proximal 
lobes rarely with 1 or 2 2 lobes (bipinnatifid). In- 
florescences of solitary axillary flowers, usually 2/ 
node, pedicels 3-5 mm long, ca. 0.15 mm diam., 
glabrous or subglabrous. Flowers with calyx 2-4 
mm long, sepals united only at the base, ca. 0.3 
mm wide, glabrous; corolla 5-6.5 mm long, violet 
to purple or bluish purple, tube ca. 4 mm long, 1 .5 
mm diam.; ovary glabrous, style 2-3 mm long. 
Fruits 8-12 mm long, 0.9-1.2 mm wide, linear, 
smooth and bisulcate, drying blackish, apex acute, 
style deciduous or sometimes persisting; seeds 0.7- 
0.9 mm long, 0.3-0.4 mm diam., cylindric with 
truncated ends, brown, the closely spaced ridges 
forming spirals around the surface. 

Rarely collected plants of moist areas in open 
sunny sites, 0-1 100 m elevation in Central Amer- 



ica. Probably flowering primarily in the wet sea- 
son, June-December. Only two Costa Rican col- 
lections have been seen: A. Jimenez 984, from 
near Playa Coco, Guanacaste, and Tonduz 13790, 
from Nicoya, Puntarenas. The species ranges from 
Mexico to northwestern Costa Rica and has been 
collected in Colombia. 

Schistophragma mexicana is very distinctive 
because of its small stature, small opposite pin- 
natifid leaves with narrow lobes, small flowers 
with tubular purple or violet corollas, and dark 
linear fruits. This is the only Costa Rican species 
of Scrophulariaceae with linear fruits and one of 
the few with pinnatifid leaves (compare Benja- 
min ia and Castilleja). 



Scoparia Linnaeus 

Herbs or subshrubs, annual or perennial, pro- 
fusely branched, leafy stems slender, 4- to 6-an- 
gled. Leaves opposite or verticillate, 2-4/node, 
sometimes caducous or reduced to scales, petioles 
little differentiated from the blade, blades dentate 
to pinnatifid or entire, base cuneate, often glan- 
dular-punctate, venation pinnate. Inflorescences 
of 1 or 2 flowers in leaf axils (1-8/node), bracts 
and bracteoles absent, pedicels filiform. Flowers 
small, calyx 4- or 5-parted, sepals ovate to lan- 
ceolate, 1 often larger than the others; corolla 
subrotate with very short tube and 4 spreading or 
reflexed lobes (upper lip emarginate), white to 
blue, rose, or yellow, with a ring of hairs at the 
mouth of the tube; stamens 4, subequal or equal, 
inserted at the base of the tube, anthers sagittate 
with parallel or subequal thecae; ovary 2-locular, 
ovules many, style short, stigma subcapitate, ex- 
serted. Fruits thin-walled capsules, globose to 
ovoid, dehiscence septicidal; seeds small, reticu- 
late with pitted surface. 

Scoparia is a Neotropical genus of ca. 1 7 species 
with centers of diversity in Mexico and southern 
South America. The plants are native to dry scrub 
and grasslands, and some species have become suc- 
cessful weeds. Scoparia is distinctive within the 
family because of its usually four-lobed corolla with 
very short tube and conspicuous hairs. Unfortunate- 
ly, the corollas are fugaceous, and the characters of 
erect habit, solitary or paired flowers, four calyx 
lobes, and crenatc or pinnatifid leaves must be used 
for identification. One species occurs in Costa Rica, 
and a second is likely to be found here. 



BURGER: FLORA COSTARICENSIS 



57 



Key to the Species of Scoparia 

la. Corolla white or pale lilac; sepals 4, obtuse or rounded at the apex; leaf blades to 35 mm long; 

plants usually becoming >40 cm tall; Mexico to Costa Rica S. dulcis 

Ib. Corolla yellow; sepals 4 or 5, acute at the apex; leaf blades to 20 mm long; plants <30 cm tall; 

Mexico to Nicaragua .5. annua 



Scoparia annua Schldl. & Cham., Linnaea 6: 
375. 1831. 

Small erect annual herbs 10-20(-30) cm tall, 
much-branched from the base or lower half, leafy 
stems 0.4-1.5 mm diam., glabrous, with 4 prom- 
inent longitudinal ridges. Leaves opposite, peti- 
oles not clearly differentiated from the blades, 
glabrous; leaf blades 5-20 mm long (including 
the narrowed base), lower blades ovate to rhombic 
and pinnatifid with rounded lobes to 4 mm long, 
distal blades linear-oblanceolate and entire or with 
a few distal teeth, glandular punctate beneath. In- 
florescences of solitary flowers in leaf axils (21 
node), pedicels 6-18 mm long, filiform, glabrous. 
Flowers with calyx 2 mm long, sepals 4- or 5- 
parted, lanceolate-oblong with acute apices, gla- 
brous; corolla to 3 mm long, rotate, yellow, fu- 
gaceous, lobes 2 mm long, rounded; stamens 
equal, filaments glabrous; ovary ellipsoid, style 
shorter than the ovary, stigma minute. Fruits 2.5- 
3 mm long, 1 .5-2 mm wide, ovoid, subtended by 
sepals ca. 3 mm long; seeds reticulate. 

Scoparia annua ranges from Mexico to central 
Nicaragua in open sunny habitats from sea level 
to 1000 m elevation; it is also found in South 
America. These small weedy plants are likely to 
be overlooked, and it is possible that they will be 
found in northwestern Costa Rica. 



Scoparia dulcis L., Sp. PI. 116. 1753. Figure 4. 

Herbs or subshrubs 0.3-1 m tall, woody at the 
base, often with many lateral branches, leafy 
stems 0.5-4 mm diam., 4- or 6-angled with 4 or 
6 longitudinal ridges, glabrous or puberulent at 
the nodes, nodes with interpetiolar line. Leaves 2 
or 3 (4) at each node, petioles 1-8 mm long (poor- 
ly differentiated), 0.3-0.9 mm wide; leaf blades 
6-35 mm long, 2-15 mm wide, narrowly elliptic 
to oblanceolate (basal leaves ovate to ovate-rhom- 
bic), apex obtuse to acute, with 5-8 teeth/side dis- 
tally, teeth 0.3-3 mm long, base obtuse to cune- 
ate, drying grayish green, glabrous, glandular 
punctate on both surfaces, 2 veins 2 or 3/side. 



Inflorescences of 1-3 flowers axillary to distal 
leaves (leaves often reduced to 6 mm long and 
bract-like), nodes with 2-6 flowers, pedicels 3-9 
mm long, 0. 1-0.2 mm diam., glabrous or minutely 
puberulent. Flowers with 4-parted calyx, 2 mm 
long (-3 mm in fruit), lobes ca. 1.3 mm long, 
elliptic to oblong, apex obtuse or rounded; corolla 
rotate and 3-4 mm wide distally, pale lilac or 
white, white-barbate within the throat; stamens ca. 
1.5 mm long, anthers 0.7 mm long. Fruits 3 mm 
long, 2-3 mm diam., broadly ovoid, smooth, yel- 
lowish gray; seeds 0.3-0.4 mm long, 0.2-0.3 mm 
diam., oblong or irregularly angled, pale brown, 
pits in longitudinal rows. 

Common plants of open sunny sites in decid- 
uous, partly deciduous, and wet evergreen forest 
formations, 0-1300 m elevation. Flowering and 
fruiting throughout the year but collected most of- 
ten in the wet season, June-December. The spe- 
cies ranges from the southeastern United States to 
southern South America and is a pantropical 
weed. 

Scoparia dulcis is recognized by its short, 
much-branched herbaceous habit, nodes with usu- 
ally two or three leaves, dentate leaf blades, small 
flower with white rotate corollas, and thin-walled 
fruits with pitted seeds. This species is a common 
weed in the lowlands of Central America. The 
bushy stems have been used to make small 
brooms and are believed to repel fleas (Standley 
& Williams, 1973). The plants have an aromatic 
odor and are called sweet broom; culantrillo, es- 
coba amarga, escobeta, and escobilla amarga are 
Spanish names. 



Sibthorpia Linnaeus 

REFERENCE O. Hedberg, A taxonomic revision 
of the genus Sibthorpia. Bot. Not. 108: 161-183. 
1955. 

Prostrate or creeping herbs rooting at many 
nodes, perennial, internodes often elongated, 
stems terete, with multicellular hairs, plants dry- 
ing greenish or brown. Leaves alternate or crowd- 



58 



FIELDIANA: BOTANY 



ed, petioles often long, slender, blades orbicular 
to reniform, margin crenate with broad rounded 
or truncated lobes, base cordate, villous to gla- 
brous, venation palmate. Inflorescences of 1-3 
flowers in axils of leaves, bracts and bracteoles 
absent, pedicels long and slender. Flowers very 
small, almost radially symmetric, calyx campan- 
ulate, united in the basal half with 5 (4-8) lobes, 
lobes subequal, acute, persisting in fruit; corolla 
subrotate with very short tube and 5 (4-8) spread- 
ing lobes, white, yellow, pink, or reddish purple, 
lobes rounded; stamens 2-8, as many as the co- 
rolla lobes or 1 or 2 fewer, borne near the apex 
of the tube, filaments slender, subequal, anthers 
slightly sagittate, thecae parallel or slightly diver- 
gent, confluent near the apex, staminodes absent; 
ovary pubescent, ovules few, style short, stigma 
simple or capitate. Fruits thin-walled capsules, 
dehiscence loculicidal; seeds few, oblong to 
ovate, convex on 1 surface, reticulate or smooth. 
Sibthorpia is a genus of six species found in 
Europe, the high mountains of Africa, and higher 
elevations in the Neotropics. The creeping growth 
form, small alternate cordate reniform or orbicular 
leaves with crenate margins, very small axillary 
flowers, almost radially symmetric corolla, short 
corolla tube, and few-seeded capsules distinguish 
this genus. Seed morphology indicates that this 
genus is related to Veronica. 

Sibthorpia repens (Mutis ex L.) Kuntze, Rev. 
Gen. PI. 3; 239. 1898. Willichia repens Mutis 
ex L., Mant. PI. 2: 558. 1771. 5. pichinchensis 
Kunth in H.B.K., Nov. Gen. Sp. 2: 390, tab. 
176. 1817. 5. triandra Suesseng., Repert. Sp. 
Nov. Regni Veg. 39: 18. 1935. 

Creeping herbs, usually with long internodes 
and few lateral branches, rooting at most nodes, 
leafy stems 0.5-1 mm diam., with slender trans- 
lucent multicellular hairs 0.2-0.8 mm long, older 
stems usually glabrescent. Leaves alternate, peti- 
oles 2-22(-30) mm long, 0.3-0.8 mm diam., with 
translucent hairs to 0.8 mm long; leaf blades 6- 
22 mm long, 6.5-25 mm wide, ovate-suborbicular 
to rounded-reniform, apex a rounded lobe, margin 
crenate with 5-9 rounded or truncated lobes/side, 
lobes 0.5-4.5 mm wide, base cordate with sinus 
1-7 mm deep, drying membranaceous, translu- 
cent hairs sparse to dense on both surfaces, major 
veins 3 or 5. Inflorescences of 1 (2) flowers in 
axils of leaves, pedicels 3-12 cm long (to 4 cm 
long in Mexico), filiform, puberulent. Flowers 
with calyx 1.7-2 mm long, united and campanu- 



late, lobes 4 or 5, 0.5-1 mm long, acute; corolla 
2-3 mm long, 3.5-5 mm wide, subrotate, pale li- 
lac to dull purple to brown-purple or dark wine- 
red (white in Ecuador), tube very short, lobes 0.7- 
1 .3 mm wide; stamens 2-4; style 2 mm long, stig- 
ma capitate. Fruits ca. 2 mm long, subglobose, 
lower part enclosed within the calyx cup, surface 
with straight ascending hairs 0.1-0.2 mm long, 
persisting style ca. 0.8 mm long; seeds ca. 0.7 mm 
long, oblong, dark with a whitish reticulum. 

Rarely collected herbs of partly shaded to deep- 
ly shaded sites in evergreen montane forest for- 
mations, 1600-3100 m elevation. Collected with 
flowers in April-June (flowering primarily in De- 
cember-March in northern Central America). 
These plants are locally common in open high- 
elevation forests of Guatemala, but we have seen 
only five collections from Costa Rica. This spe- 
cies ranges from central Mexico to Costa Rica and 
is found in the Andes. 

Sibthorpia repens is recognized by the creeping 
habit, slender stems with many adventitous roots, 
small suborbicular leaves with lobed crenate mar- 
gins, small axillary flowers, subrotate corollas 
with two epipetalous stamens, and few-seeded 
capsules enclosed at their base by the campanu- 
late calyx. This species is quite variable in a num- 
ber of significant taxonomic features. Despite this 
variability, it seems probable that, as Hedberg 
(1955, cited above) and Williams (1972) have 
suggested, the Neotropical material represents a 
single species. These plants are easily mistaken 
for species of Hydrocoryle (Apiaceae) or Dichon- 
dra (Convolvulaceae) with similar growth form. 



Stemodia Linnaeus 
Nomen conservandum 

RKPKRKNCKS B. Turner & C. Cowan, Taxo- 
nomic overview of Stemodia (Scrophulariaceae) 
for North America and the West Indies. Phytolo- 
gia 74: 61-103. 1993. Taxonomic overview of 
Stemodia (Scrophulariaceae) for South America. 
Phytologia 75: 281-324. 1993. 

Erect or procumbent herbs or subshrubs to 3 
m tall, annual or perennial, stems terete or angu- 
lar, often much-branched, usually puberulent, of- 
ten with gland-tipped hairs, drying greenish or 
brown. Leaves opposite, subopposite, or verticil- 
late, simple, sessile or petiolate, blades serrulate 
to pinnately lobed (rarely subentire), pinnately 
veined or subpalmate, often glandular-punctate. 
Inflorescences of 1-3 flowers in axils of leaves 
or bracts (1-6/node), sometimes forming distal ra- 
cemes, thyrses, or panicles, bracts absent when 



BURGER: FLORA COSTARICENSIS 



59 



flowers are subtended by leaves, pedicels slender, 
bracteoles present or absent at the base of the ca- 
lyx. Flowers with deeply 5-parted calyx, sepals 
narrow and acute, equal or more often slightly 
unequal (adaxial sepal often larger), valvate in 
bud, persisting and enlarging slightly in fruit; co- 
rolla tubular to narrowly campanulate, 2-lipped, 
white to blue, lilac, or purple (yellowish when 
faded), 4- or 5-lobed, bilabiate, upper (adaxial) lip 
2-lobed or entire, lower lip 3-lobed, often bearded 
at the base of the lobes; stamens 4, equal or of 2 
unequal pairs, included, borne on the tube, fila- 
ments glabrous or puberulent, anthers glabrous, 
with an enlarged connective separating the 2 par- 
allel or divergent thecae, a staminode sometimes 
present; ovary ovoid, style terete, stigma ligulate, 
often minutely 2-lobed or capitate and reflexed. 
Fruits dry capsules, often bisulcate, dehiscing lo- 
culicidally and partly septicidally from the apex 
(2- or 4-valved), placenta drying to form a peg 
that is free from the apex of the locule; seeds 
many, small, oblong to pyriform or irregular, often 
longitudinally sulcate or ridged, usually with a 
short stipe at one end. 



According to Turner and Cowan (1993) Ste- 
modia is a genus of 29 Neotropical and 20 Old 
World species, including several widespread 
weeds. They are placed in subfamily Antirrhino- 
ideae, tribe Gratioleae, subtribe Stemodiinae. Be- 
cause of the variability of important taxonomic 
characteristics within the genus, it is likely that 
Stemodia may be broken up into smaller genera 
(see Darcyd). Mined (1918) split the genus into a 
number of genera that have generally not been 
accepted. His characters were often variable and 
his nomenclature was flawed, which, as much as 
his splitting, accounts for the lack of interest in 
his classification. We follow Turner and Cowan's 
treatment and their annotations. These generally 
short, weedy plants are recognized by their op- 
posite or ternate leaves, one- to six-flowered 
nodes, calyx with narrow sepals united only near 
the base, tubular two-lipped bluish to purple or 
white corollas, and style often curved at the ter- 
minal stigma. Other important characters are the 
anthers with thecae slightly separated by an ex- 
panded connective, and the seeds often with lon- 
gitudinal ridges or sulci. 



Key to the Species of Stemodia 

la. Flowers and fruits borne on slender pedicels 6-19 mm long; corollas white to pink or yellowish (if 
flowers have a prominent calyx tube, see Torenia) 2 

Ib. Flowers and fruits subsessile or on pedicels < 4 mm long; corollas blue to purple 3 

2a. Flowers 7-9 mm long; fruits 4-5 mm long; plants to 20 cm tall 5. angulata 

2b. Flowers 12-15 mm long; fruits 5-7 mm long; plants to 80 cm tall 5. peduncularis 

3a. Fruits at least 3 mm long; calyx 4-6 mm long; flowers on spike-like axes; leaves subsessile, larger 
leaves to 6 cm long and oblanceolate 5. durantifolia 

3b. Fruits 1.8-2.5 mm long; calyx 2-4 mm long; flowers in leaf axils; leaves petiolate, larger leaves to 
2 cm long and ovate-triangular S. verticillata 



Stemodia angulata Oerst., Vidensk. Meddel. 
Dansk Naturhist. Foren. Kjobenhavn 1853: 22. 
1854. Stemodiacra angulata (Oerst.) Kuntze, 
Rev. Gen. PI. 2: 466. 1891. Stemodia jorullensis 
subsp. reptans Minod, Bull. Soc. Bot. Geneve, 
ser 2, 10: 190. 1918. Figure 2. 

Small herbs 5-20 cm tall, stems to 40 cm long, 
often much-branched, leafy stems 0.3-1.7 mm 
diam., pubescence of thin multicellular translucent 
hairs 0.2-1.2 mm long, slightly viscid. Leaves op- 
posite, petioles 2-9 mm long, 0.2-0.5 mm wide, 
puberulent with thin whitish hairs; leaf blades 6- 
20 mm long, 3-18 mm wide, ovate to ovate-tri- 
angular or ovate-rhombic, apex obtuse or round- 



ed, margin with 7-1 1 teeth/side, base broadly ob- 
tuse or truncate, glandular punctate and sparsely 
puberulent above and below, 2 veins 3-5/side, 
ascending. Inflorescences of 1 (2 or 3) flowers in 
leaf axils, pedicels 4-12 mm long (-19 mm in 
fruit), 0.1-0.2 mm diam., puberulent, bracteoles 
absent. Flowers with calyx 4-6 mm long, sepals 
3-5 mm long, ca. 0.8 mm wide at the base, pu- 
berulent; corolla 7-9 mm long, white with purple 
lines within (yellowish in age), tube to 7 mm long, 
1.4-2 mm diam., lobes to 2 mm long; stamen 
pairs very unequal, 2 borne at the base and 2 in 
the upper part of the tube. Fruits 4-5 mm long, 
ca. 2 mm diam., oblong, pale yellowish brown, 
glabrous; seeds 0.3-0.4 mm long, 0.2-0.3 mm 



60 



FIELDIANA: BOTANY 



diam., variously shaped, brown to black, longi- 
tudinally sulcate with 6-8 ridges. 

Plants of open sunny sites in both seasonally 
dry and wet evergreen forest areas, 10-1400 m 
elevation. Probably flowering and fruiting 
throughout the year, but collected most often in 
August-March. The species ranges from Mexico 
and the West Indies to northwestern South Amer- 
ica. 

Stemodia angulata is recognized by its short 
stature, opposite ovate-triangular leaves, solitary 
axillary flowers on slender pedicels, nearly sepa- 
rate sepals, and small white to rose corollas. 
Standley (1938) included this species under S. pe- 
duncularis, but that species, although similar, is 
larger in almost all its dimensions. In the Flora 
of Panama, specimens of this species were placed 
under 5. jorullensis Kunth in H.B.K., a species of 
western Mexico. 

Stemodia durantifolia (L.) Swartz, Obs. Bot. 
240. 1791. Capraria durantifolia L., Syst. Nat. 
ed. 10: 1116. 1759. Figures 4 and 6. 

Herbs 0.2-0.9(-1.5) m tall, annual or becom- 
ing woody at the base, usually with many branch- 
es, leafy stems 0.5-4.5 mm diam., pubescent with 
crooked translucent multicellular viscid glandular 
hairs 0.1-0.5 mm long, stems 4- angled with 4 lon- 
gitudinal ridges. Leaves 2-3/node, diminishing in 
size distally and intergrading with the bracts, ses- 
sile and clasping the stem; leaf blades 1 .4-7 cm 
long, 4-18 mm wide, smaller leaves narrowly el- 
liptic-oblong to narrowly ovate, larger leaves ob- 
lanceolate, apex acute, margin with 3-12 teeth/ 
side, base auriculate in larger leaves, both surfac- 
es with minute (0. 1 mm) glandular hairs and lon- 
ger thin hairs, 2 veins 2-5/side. Inflorescences 
4-17 cm long, spicate or paniculate to 50 cm long 
with opposite spicate branches and internodes 5- 
15 mm long, puberulent like the stems, bracts (re- 
duced leaves) 5-16 mm long, 2-5 mm wide, ped- 
icels 0-3 mm long, bracteoles 1 or 2, subtending 
the calyx, linear, 2-5 mm long. Flowers with ca- 
lyx 4-6 mm long, ca. 1 mm diam., lobes 2-4 mm 
long, linear-lanceolate, densely glandular-puberu- 
lent; corolla 5-7 mm long, blue to purple, glan- 
dular-puberulent, tube ca. 1 mm diam., yellow 
within the throat, lower lobes ca. 1.5 mm long; 
stamens inserted at the same level near the mouth 
of the tube, connective globose; style 3-4 mm 
long. Fruits 3-4 mm long, 2 mm diam., narrowly 
ovoid, smooth, yellowish brown; seeds 0.2-0.3 
mm long, 0.15 mm diam., narrowly oblong or nar- 



rowed at one end, dark brown, with longitudinal 
rows of rounded tubercles (50X). 

Weedy plants of open sunny moist sites or part- 
ly shaded woodland in seasonally dry vegetation 
of the Pacific slope (rarely collected on the Carib- 
bean slope), 0-900 m elevation. Flowering and 
fruiting throughout the year, but collected most 
often in November-March. The species ranges 
from the southwestern United States, Mexico, and 
the West Indies to Chile and Brazil. 

Stemodia durantifolia is recognized by the 
erect, much-branched, weedy habit, viscid pubes- 
cence, sessile leaves with basal blades oblanceo- 
late, long distal spikes with short internodes, small 
flowers, and blue or purple corollas. The flowers 
are usually two per node but may be as many as 
six per node. Distal bracts may be alternate or 
tightly congested on flowering stems. The brac- 
teoles are difficult to distinguish from the sepals 
but are an unusual character. These plants resem- 
ble species of Hyptis (Lamiaceae). 

Stemodia peduncularis Benth. in DC., Prodr. 10: 
382. 1846. Stemodiacra peduncularis (Benth.) 
Kuntze, Rev. Gen. PI. 2: 466. 1891. Figure 3. 

Erect or decumbent herbs 10-80 cm tall, usu- 
ally with few or no lateral branches on distal 
stems, leafy stems 0.5-2 mm diam., pubescence 
of thin white or viscid hairs 0.2-0.5 mm long, 
glabrescent. Leaves opposite, petioles 4-14 mm 
long. 0.3-0.9 mm wide, puberulent with thin 
whitish hairs, leaf base decurrent on the petiole; 
leaf blades 8-40(-50) mm long, 6-25(-35) mm 
wide, ovate to ovate-triangular or ovate-oblong, 
apex acute, margin with 8-12 teeth/side, base ob- 
tuse to truncate, punctate and minutely (0.1-0.2 
mm) puberulent on both sides, 2 veins 3 or 4/ 
side, strongly ascending. Inflorescences of 1 or 2 
flowers in leaf axils (1-4/node), pedicels 6-18 
mm long (-50 mm in fruit), 0.1-0.2 mm diam., 
with thin whitish hairs 0.1-0.2 mm long. Flowers 
with sepals 4-7 mm long. 0.7-1 mm wide at the 
base, linear to linear-triangular, minutely puberu- 
lent (sometimes glandular punctate); corolla 1 1- 
15 mm long, white or red-veined (yellowish), tube 
8-10 mm long, 2-3 mm diam.. externally gla- 
brous, lobes 2-3 mm long; stamens borne near the 
base of the tube; style 4-7 m long. Fruits 5-7 
mm long, narrowly ovoid; seeds 0.5-1 mm long, 
irregular in shape, black. 

Infrequently encountered plants of shaded sites 
in moist evergreen montane forest formations 
along the Continental Divide and Pacific slope in 



BURGER: FLORA COSTARICENSIS 



61 



the central volcanic highlands and Cordillera de 
Talamanca. 1300-2400 m elevation. Flowering 
and fruiting throughout the year, but collected 
mostly in December-March. The species ranges 
from western Mexico to western Panama. 

Stemodia peduncularis is recognized by its 
short, erect, few-branched habit, opposite serrate 
leaves, mostly solitary flowers on long slender 
pedicels, separate linear sepals, and white tubular 
corollas. Standley (1938) incorrectly included 5. 
angulata under this species. This species resem- 
bles our species of Darcya in general appearance. 

Stemodia verticillata (Miller) Hassler, Trab. 
Mus. Farmacol. 21: 110. 1909. Erinus verticil- 
lotus Miller, Card. Diet. ed. 8. 1768. S. parvi- 
flora W. T. Aiton, Hortus Kew., ed. 2, 4: 52. 
1812. S. arenaria Kunth in H.B.K., Nov. Gen. 
Sp. 2: 357. 1817. Lindernia verticillata (Miller) 
Britton in Britton & Wilson, Bot. Porto Rico 6: 
184. 1925. Figure 2. 

Small herbs 5-15(-25) cm tall, prostrate to as- 
cending, rooting only near the base, internodes 
mostly 4-14 mm long, leafy stems 0.5-1 mm 
diam., densely puberulent with straight or curved 
(often glandular) hairs 0.2-0.8 mm long. Leaves 
usually 2 or 3 (4) per node, petioles 2-1 1 mm 
long, 0.2-0.5 mm wide, puberulent, leaf base de- 
current on the petiole; leaf blades 5-12(-18) mm 
long, 3-10(-14) mm wide, ovate to ovate-trian- 
gular or ovate-oblong, apex obtuse, margin with 
5-9 teeth/side, teeth ca. 0.5 mm long, base obtuse 
or truncated, both surfaces with hairs 0.1-0.5 mm 
long, 2 veins 3-5/side. Inflorescences of usually 
2 subsessile axillary flowers (2-6/node), pedicels 
0.5-2 mm long, minutely puberulent, bracteoles 
absent. Flowers with calyx 2-4 mm long, sepals 
equal or subequal, 0.5 mm wide near the base, 
with thin whitish and glandular hairs; corolla 3- 
5 mm long, blue-violet or purple with white 
throat, tube 1 mm diam., lobes 1-2 mm long; sta- 
mens borne in the lower half of the tube, anther 
thecae 0.3 mm long; style 1-1.5 mm long with 
recurved stigma. Fruits 1.8-2.5 mm long, 1.4-2 
mm wide, ovoid-subglobose, glabrous, pale yel- 
low-brown; seeds ca. 0.5 mm long, 0.2-0.3 mm 
diam., narrowly ovoid, yellowish, with dark tip 
0.05 mm long, surface with longitudinal ridges 
and sulci. 

Uncommon weeds of open sunny or shaded 
sites (often on sand or wet gravel) in evergreen 
or deciduous vegetation, 0-1600 m elevation. 
Probably flowering and fruiting throughout the 



year, but collected in Costa Rica primarily in 
July-August. The species ranges from Mexico 
and the West Indies to Argentina. 

Stemodia verticillata is recognized by its small, 
often prostrate habit, mixture of both glandular 
and eglandular hairs, small petiolate leaves sub- 
tending solitary subsessile flowers, narrow sepals, 
and bluish or purple corollas. Plants with three or 
four leaves per node are distinctive. The glandular 
punctations of the leaf surfaces are often obscure. 
These plants have been called hierba santa (Costa 
Rica) and corrimiento (El Salvador). 



Tetranema Bentham ex Lindley 

REFERENCES M. Grayum & B. Hammel, The 
genus Tetranema (Scrophulariaceae) in Costa 
Rica, with two new species. Phytologia 79: 269- 
280. 1995 (1996). T. Mendez-Larios & J. Villa- 
senor. Revision taxonomica del genero Tetranema 
(Scrophulariaceae). Acta Bot. Mexico 32: 53-68. 
1995. 

Herbs, erect or decumbent, perennial, some- 
times woody at the base but the stems usually 
short, glabrous or pubescent. Leaves opposite and 
often crowded near the base, equal or unequal at 
a node, subsessile and clasping the stem or peti- 
olate, blades dentate to subentire, venation pin- 
nate. Inflorescences axillary, cymose to subum- 
bellate or congested, long-pedunculate, bracteate, 
pedicels short but often elongating in fruit. Flow- 
ers with the calyx deeply 5-parted, campanulate, 
lobes narrow and attenuate-acuminate, with prom- 
inent longitudinal ribs; corolla tubular to campan- 
ulate, 2-lipped, glabrous, white to rose, purple or 
less often red, tube longer than the lobes, upper 
lip emarginate ( 1 -lobed), lower lip with 3 spread- 
ing lobes, imbricate in bud; stamens 4 (rarely 3), 
filaments of 2 longer and 2 shorter pairs, anthers 
2-thecous, a staminode very small (in ours) or ab- 
sent; ovary 2-locular, style slender or thick, stig- 
ma capitate or bilobed. Fruits capsules, ovoid to 
globose, glabrous, dehiscence loculicidal; seeds 
many, angled or tetrahedral, surface reticulate or 
foveolate. 

Tetranema is a genus of six species from Mex- 
ico and Central America. It is distinguished by its 
opposite leaves, axillary and cymose inflorescenc- 
es, separate sepals, four-lobed and two-lipped co- 
rolla, and loculicidal capsules. Our recently dis- 
covered species are unusual in having red (prob- 
ably hummingbird-pollinated) flowers. These dis- 
tinctive plants (Fig. 27) may resemble Razisea 



62 



FffiLDIANA: BOTANY 



spicata Oerst. and Odontonema tubaeforme (Ber- 
tol.) Kuntze of the Acanthaceae. The Costa Rican 
species are similar to T. megaphyllum (Brande- 
gee) L. O. Williams, but that species has smaller 
(2-4 mm) corolla lobes and larger (6-10 mm) flo- 
ral bracts and is endemic to Mexico. Species of 
Tetranema are sometimes cultivated as ornamen- 



tals, especially T. roseum (M. Martens & Galeoti) 
Standl. & Steyerm. The new species come from 
the Pacific slope of central Costa Rica and both 
slopes of southern Costa Rica. The dichotomy in 
Flora of Guatemala separating Tetranema from 
Penstemon and Uroskinnera is inverted (Standley 
& Williams, 1973, p. 321). 



Key to the Species of Tetranema 

la. Inflorescences with 14-30 flowers, peduncle purple; corollas 26-35 mm long, pubescent internally 
with a band of flat yellow hairs along the entire ventral (abaxial) surface and onto the lower lobe; 
leaf apex rounded to short-acuminate; 1200-1600 m on Cerro Turrubares T. floribundum 

Ib. Inflorescences with 2-12 flowers, peduncle green; corollas 44-55 mm long, glabrous throughout or 
rarely pubescent on the lower lobe and mouth; leaf apex long-acuminate; 500-1200 m elevation on 
both Caribbean and Pacific slopes T. gamboanum 



Tetranema floribundum B. Hammel & M. Gra- 
yum, Phytologia 79: 269-280. 1995 (1996). 
Figure 27. 

Erect herbs (0.3-)0.8-2 m tall, decumbent near 
the base and often rooting at the lower nodes, in- 
ternodes to 5 cm long, arachnoid or wooly-pu- 
bescent when young, leafy stems 1 .5-5 mm thick, 
glabrescent. Leaves opposite, subsessile with pet- 
ioles 0-6 mm long, ca. 2 mm thick; leaf blades 
14-24 cm long, 7-13 cm wide, broadly elliptic to 
oblanceolate or spatulate, apex abruptly acute or 
short-acuminate, margins coarsely serrate or un- 
dulate-toothed with teeth ca. 1 mm high, base 
gradually cuneate, mostly glabrous above, mi- 
nutely puberulent on the major veins beneath, lus- 
trous beneath when dry, 2 veins 8-1 I/side, as- 
cending. Inflorescences 15-30 cm long, 14-30- 
flowered, peduncles 13-23 cm long, purple, with 
4 longitudinal ridges, bracts 1-5 mm long, nar- 
rowly triangular, margins ciliate (often only at the 
base), pedicels 10-20 mm long, glabrous. Flow- 
ers with calyx ca. 3 mm long (-5 mm in fruit), 
divided nearly to base, striate and glabrous (ex- 
cept along margins), with a short apical projec- 
tion; corolla 26-35 mm long, 5-6 mm diam., tu- 
bular and slightly expanded distally and curved 
upward, red-orange, lobes 4, ca. 1 3 mm long, 2.5- 
5.5 mm wide, the 3 lower rounded apically and 
reflexed-spreading, puberulent internally abaxial- 
ly; stamens slightly exserted, anthers 0.8-0.9 mm 
long, staminode ca. 0.5 mm long; ovary ca. 3.5 
mm long. Fruits not seen at maturity, ca. 8 mm 
long, ovoid, borne on pedicels ca. 20 mm long. 

Plants of evergreen lower montane forest for- 



mations at 1200-1600 m elevation on Cerro Tur- 
rubares. Flowering in December-January and 
March. This species is known only from an iso- 
lated peak on the Pacific slope of the southeastern 
part of the central plateau in the province of San 
Jose". 

Tetranema floribundum is recognized by its 
larger subsessile elliptic to obovate leaves with 
many ascending secondary veins, inflorescences 
with long purple peduncles, five separate acute 
sepals, curved-tubular red corollas, and isolated 
habitat. Cerro Turrubares is also the location of a 
recently discovered and unusual species of Psy- 
chotria (Burger & Jime'nez, 1994). 

Tetranema gamboanum M. Gray urn & B. Ham- 
mel, Phytologia 79: 269-280. 1995 (1996). Fig- 
ure 27. 

Erect herbs 1-2 m tall, internodes to 1 1 cm (or 
more) long, at first minutely strigulose, 1 .7-5 mm 
thick, glabrous but with a puberulent interpetiolar 
line. Leaves opposite, sessile to subsessile or with 
petioles to 1 cm long, canaliculate above, margins 
ciliate at the base; leaf blades 14-31 cm long, 5- 
1 1 cm wide, elliptic to elliptic-obovate, apex long- 
acuminate, margin coarsely serrate with teeth 1- 
2 mm high, base cuneate and abruptly narrowed 
at the base, glabrous above or with few hairs 
along midvein and major veins, minutely puber- 
ulent along the veins beneath, 2 veins 9-13/side. 
Inflorescences axillary cymes with 2-13 flowers, 
peduncles 9-24 cm long, green, quadrangular 
with 4-winged ribs, bracts 0.5-2 mm long, subu- 
late to narrowly triangular, ciliate along the mar- 
gins, pedicels 9-12 mm long (-20 mm in fruit), 



BURGER: FLORA COSTARICENSIS 



63 



glabrous. Flowers with 5-parted calyx, sepals 3- 
5 mm long (-6 mm in fruit), inner 2 sepals slight- 
ly smaller than the outer, narrowly to broadly 
ovate, imbricate in bud, minutely ciliolate along 
the margin, apex acute, venation parallel; corolla 
44-55 mm long, ca. 5 mm diam., narrowly tu- 
bular, gradually expanded distally and slightly 
curved, bright red, glabrous throughout or with 
hairs at throat and lower lobes, lobes 8-14 mm 
long, 3-4 mm wide, lanceolate or oblong, the 3 
lower lobes obtuse to rounded at the apex and 
spreading-reflexed; stamens slightly exserted, fil- 
aments dilated near base, glabrous, anthers 0.8-1 
mm long, with divergent thecae confluent at the 
base, staminode 1.5-2 mm long; ovary 3-4 mm 
long, narrowly ovoid, style exserted, stigma cap- 
itate. Fruits capsules, 6-9 mm long (not including 
the 1-2 mm style base), subglobose-apiculate, 
glabrous; seeds 0.5-0.7 mm long, 0.4-0.5 mm 
diam., rectangular or oblong-rounded, surface 
with prominent pits (foveolate), dark yellow to 
brown or blackish. 

Newly discovered plants of evergreen rain for- 
est formations on the Caribbean slope of the Cor- 
dillera de Talamanca and on the southern Pacific 
slope, 500-1200 m elevation. Flowering and fruit- 
ing in December-May. This species has been col- 
lected only from between 83W and 84W in the 
southern half of Costa Rica. 

Tetranema gamboanum is distinguished by its 
erect stems with larger ellipsoid leaves with many 
lateral veins, axillary cymes on long peduncles, 
five-parted calyx, and bright red, slightly curved 
corolla tubes. 



Torenia Linnaeus 

Annual or perennial herbs, erect or procum- 
bent, branching from both basal and distal nodes, 



stems 4-angled with longitudinal ridges, puberu- 
lent to hirsute or glabrous. Leaves opposite, pet- 
iolate, blade margins entire to serrate or crenulate, 
venation pinnate. Inflorescences of 1-3 flowers 
in axils of distal leaves or in short terminal or 
axillary few-flowered racemes, linear bracts 
sometimes present, pedicels usually held at a 45 
angle to the stem. Flowers showy or small, calyx 
usually somewhat shorter than the corolla tube, 
united to form a long tube with 3-5 short lobes 
(or 2-lipped), with 3-5 longitudinal ribs or broad 
wings, persisting and enlarging in fruit; corolla 
campanulate to tubular or salverform, bilabiate 
with the upper lip erect and 2-lobed, lower lip 
with 3 lobes, tube usually widened distally, beard- 
ed in the throat; stamens 4, in 2 unequal pairs, 
filaments inserted near the top of the tube and 
usually with a tooth-like appendage at the base, 
anterior 2 arching upward over the stigma (with 
anthers connivent or reduced), thecae oblong to 
linear, divaricate by the enlarged connective; disc 
prominent and saucer-shaped or cupulate; ovary 
oblong, style straight, stigma slightly 2-lobed. 
Fruits capsules, usually enclosed within the per- 
sisting perianth tube, oblong-ellipsoid, dehiscing 
septicidally to the base, placenta linear with sep- 
tum forming wings; seeds globose to variously 
angled, tuberculate or reticulate. 

Torenia is a genus of 40 to 80 species of trop- 
ical Africa and Asia. A few species are cultivated 
ornamentals that have become naturalized in the 
Americas. Torenia is closely related to Lindernia, 
sharing the curved appendaged filaments often 
found in that genus. Species of Torenia in the 
New World can be distinguished from Lindernia 
by their larger flowers, tubular calyx, elongate 
fruits, and globose pitted seeds. 



Key to the Species of Torenia 

la. Corolla 30-40 mm long, lower lobes dark purple or blue-violet; calyx broadly winged and 5-10 
mm wide; leaves 1 6-50 mm long, often truncate at the base T. fournieri 

Ib. Corolla 7-12 mm long, lower lobes pale purple or pale blue- violet; calyx not winged, 2-3 mm 
diam.; leaf blades 8-22 mm long, obtuse to rounded at the base T. thouarsii 



Torenia fournieri Linden in Fourn., Illustr. Hor- 
tic. 23: 129, tab. 249. 1876. Figure 8. 

Weak-stemmed erect herbs 15-50 cm tall, of- 
ten reddish near the base, main stems with 2-5 
branching nodes, leafy stems 1-3 mm diam., 4- 



angled with prominent longitudinal ridges, nodes 
with thin whitish hairs 0.2-0.5 mm long (inter- 
nodes often glabrous). Leaves opposite through- 
out, petioles 3-17 mm long, 0.7-1.4 mm wide, 
with thin whitish hairs along the adaxial margins; 
leaf blades 16-50 mm long, 6-25 mm wide, tri- 



64 



FIELDIANA: BOTANY 



angular to ovate-triangular or ovate-elliptic, apex 
acute, margin with prominent teeth, base truncate 
to obtuse, upper surface with few thin straight 
hairs 0.2-0.5 mm long, 2 veins 3-6/side, strongly 
ascending. Inflorescences of solitary flowers in 
distal leaf axils (2/node) or more often in short 
terminal racemes with 4-8 flowers, bracts 2-5(-12) 
mm long, linear, pedicels 4-18 mm long, 0.6-1 
mm wide, glabrous or very sparsely puberulent. 
Flowers with calyx 13-22 mm long, 5-10 mm 
wide, tube more than half the calyx length, broad- 
ly winged to produce an ovate form with acute or 
acuminate apices, ciliolate; corolla 3-4 cm long, 
2-3 cm wide distally, tube 7-9 mm diam. in the 
center, whitish near the base, upper lip often pale 
blue with the lower lobes very dark blue-violet, a 
yellow patch present within at the base of the cen- 
tral lobe. Fruits 1-2 cm long, included within the 
slightly enlarged calyx. 

Torenia fournieri, a native of southeastern 
Asia, is widely planted as an ornamental. This 
species may become naturalized in lowland and 
mid-elevation areas of Central America, but it 
seems doubtful that the populations persist for 
long. These plants are easily distinguished by 
their small size, opposite (often triangular) leaves, 
few-flowered racemes, broadly winged calyx, and 
relatively large corollas with dramatic dark pur- 
ple-violet coloring. This species is unusual within 
the genus in having broadly winged sepals and in 
lacking basal appendages on the lower filaments. 
Torenia asiatica L. is similar but has prostrate 
stems. The large colorful corollas make these 
plants especially attractive. They can be mistaken 
for members of the Gesneriaceae. 

Torenia thouarsii (Cham. & Schldl.) Kuntze, 
Rev. Gen. PI. 2: 468. 1891. Nonenia thouarsii 
Cham. & Schldl., Linnaea 3: 18. 1828. Linder- 
nia thouarsii (Cham. & Schldl.) Edwin, Phy- 
tologia 19: 361. 1970. Figure 3. 

Procumbent or erect herbs 10-30 cm tall, leafy 
stems 0.6-1.3 mm diam., quadrangular, internodes 
glabrous or with thin whitish hairs 0.2-0.5 mm 
long, nodes usually with thin hairs. Leaves op- 
posite throughout, petioles 1-7 mm long, ca. 1 
mm wide with thin lateral margins, sparsely pu- 
berulent; leaf blades 8-22 mm long, 5-12 mm 
wide, ovate-elliptic to ovate-triangular, apex 
acute, margins with 4 or 5 prominent teeth/cm, 
base obtuse to rounded, surfaces glabrous or 
sparsely puberulent, 2 veins 3-5/side, arcuate as- 
cending. Inflorescences of 1 or 2 flowers axillary 



to foliage leaves (l-r4/node), bracts ca. 2 mm 
long, linear, pedicels 2-15 mm long, ca. 0.5 mm 
diam., glabrous or sparsely puberulent. Flowers 
with calyx 4-12 mm long, 1.3-2 mm diam. (to 
18X3 mm in fruit), narrowly tubular to tubular- 
ellipsoid, glabrous, lobes 2-3 mm long, acute; co- 
rolla 7-12 mm long, bluish, purple, or white, 
lobes crenate; filaments with a linear appendage 
at base, staminode with a rounded tip. Fruits 8- 
10 mm long. 2-3 mm wide, narrowly oblong-el- 
lipsoid, included within the enlarged calyx tube 
or the tube splitting. 

Torenia thouarsii, a native of India, is natural- 
ized in parts of Central and South America. In 
Costa Rica it is occasionally found in wet sites at 
10-2000 m elevation, from the Caribbean low- 
lands to the central highlands. These plants resem- 
ble species of Lindernia and Stemodia but differ 
in the tubular calyx and narrowly oblong fruits. 



Veronica Linnaeus 

Annual or perennial herbs (rarely shrubs), 
prostrate or ascending to erect, often branching 
from the base, stems terete or with longitudinal 
ridges, glabrous or puberulent. Leaves opposite 
near the base and usually alternate distally, some- 
times intergrading with smaller floral bracts, ses- 
sile or petiolate. blades subentire to dentate, cre- 
nate or divided, pubescence of simple multicel- 
lular or glandular hairs, venation usually palmate 
or subpalmate. Inflorescences of solitary flowers 
in leaf axils or of elongate racemes (or spike-like) 
with flowers solitary in the axils of bracts, usually 
terminal, pedicels very short to long and slender. 
Flowers mostly small, calyx deeply 4-lobed (rare- 
ly 5-lobed), the lobes (sepals) subcquai to un- 
equal, slightly overlapping in bud, usually per- 
sisting and slightly enlarged in fruit, glabrous or 
puberulent; corolla rotate (rarely campanulate), 
tube very short, corolla lobes 4 (5). unequal, with 
the lower lobe the smallest, lateral lobes exterior 
in bud, blue to purple or white (rarely reddish); 
stamens 2, borne at either side of the upper lobe, 
exsertcd, filaments attached at the base of the tube 
and free, anthers 2-thecous. thecae confluent at 
the apex; ovary rounded, 2-locular, ovules few to 
many, style simple, persisting in fruit but not en- 
larging, stigma capitate. Fruits capsules, flattened 
at right angles to the plane of the septum, obovoid 
or 2-lobed (obcordate) with a depressed or emar- 
ginate apex, dehiscence loculicidal (also some- 
times septicidal); seeds few to many, ovate to or- 



BURGER: FLORA COSTARICENSIS 



65 



bicular, surfaces smooth to rugulose, often with 
the inner face concave, embryo often U-shaped. 
Veronica is a genus of 150 to 250 species con- 
fined to cool climates. The great majority of spe- 
cies are found in Europe and Asia, but a number 
have become widespread weeds; it seems likely 
that none are native to Central America. The ge- 
nus is distinguished (in our area) by its small her- 
baceous growth form, small leaves becoming al- 
ternate distally, small flowers with prominent ca- 
lyx lobes (sepals), bluish to lilac or white corollas 
with the four lobes usually held in a single plane 
(rotate), two stamens, and unusual fruits. The cap- 



sules are flattened perpendicular to the plane of 
the septum and often have an emarginate or deep- 
ly notched apex. The rounded lateral lobes of the 
fruits (carpels) can be lenticular or rounded. The 
genus is placed in the tribe Veroniceae and is 
closely related to Sibthorpia and the Australasian 
Hebe. Unfortunately, some species of Veronica 
are very similar, and a number of Central Amer- 
ican collections have been misidentified in the 
past. In Costa Rica the genus has not been found 
below 1200 m elevation, and only two species (V. 
polita and V. serpyllifolia) are commonly collect- 
ed. 



Key to the Species of Veronica 

la. Rowers borne on conspicuous pedicels, the pedicels much longer than the calyx lobes, flowers 
arising from the axils of leaves or from short axillary racemes; leaves with prominent teeth along 

the margins; styles 1.3-3 mm long 2 

Ib. Flowers borne on short pedicels, the pedicels much shorter to slightly longer than the calyx lobes, 
flowers arising from the axils of leaves or from the axils of floral bracts differing greatly in size 
and form from the basal leaves; leaves with small or inconspicuous teeth along the margin (except 

in V. arvensis); styles 0.5-2.5 mm long 3 

2a. Corolla < 8 mm wide; fruits about as broad as long, ca. 4 mm wide, venation obscure; fruiting 
sepals ca. 5 mm long; leaf blades often triangular-ovate; styles 1.3-1.6 mm long; occasional at 

1400-3300 m elevation V. polita 

2b. Corolla 8-13 mm wide; fruits distinctly broader than long, 5-9 mm wide, venation prominent; 
fruiting sepals ca. 6 mm long; leaf blades usually ovate-oblong; styles 1.8-3 mm long; rarely 

collected in Costa Rica V. persica 

3a. Lower leaf blades 8-28 mm long, narrowly oblanceolate, leaves gradually intergrading into the 

smaller distal floral bracts; erect stems to 45 cm tall; corolla white with purple lines [pedicels ca. 

1 mm long; styles ca. 0.5 mm long; rarely collected in southern Central America] . . V. peregrina 

3b. Lower leaf blades up to 12 mm long, usually broadly ovate; leaves usually clearly differentiated 

from the smaller floral bracts; erect stems rarely exceeding 20 cm in height; corolla usually bluish 

4 

4a. Pedicels becoming 6 mm long; styles 1.8-2.5 mm long; margins of leaf blades subentire to slightly 

crenate; frequent in Costa Rica V. serpyllifolia 

4b. Pedicels 0.5-1 .2 mm long; styles ca. 0.6 mm long; margins of leaf blades minutely to conspicuously 
dentate-crenate; rarely collected in Costa Rica V. arvensis 



Veronica arvensis L., Sp. PI. 13. 1753. 

Annual herbs, 5-20(-40) cm tall, stems usually 
branching only near the base, leafy stems 0.4-1.3 
mm diam., puberulent with thin curved whitish 
hairs 0.2-0.7 mm long. Leaves opposite (but 
bracts of the inflorescences alternate), sessile or 
the basal leaves with petioles 1-4 mm long; leaf 
blades 4-12 mm long, 3-10 mm wide, broadly 
ovate to ovate-oblong, apex obtuse or rounded, 
margin crenate-dentate with 2-6 teeth 0.3-0.8 
mm high, base obtuse or truncate, with thin hairs 



ca. 0.5 mm long on both surfaces, venation pal- 
mate with 3 or 5 major veins. Inflorescences erect 
spike-like racemes, 2-15 cm long, bracts 4-7 mm 
long, 1-2 mm wide, narrowly oblong to narrowly 
ovate, narrowed at the base, rachis 0.5-1.2 mm 
diam., puberulent, pedicels 0.5-1.2 mm long. 
Flowers with unequal calyx lobes 2.5-4 mm long 
(becoming 5 mm long in fruit), ca. 1 mm wide, 
narrowly oblong to narrowly elliptic-oblong, pu- 
berulent with short thin hairs; corolla blue, upper 
lobe ca. 2 mm long; anthers ca. 0.4 mm long; 
styles ca. 0.6 mm long. Fruits 3.5-4 mm long, 



66 



FffiLDIANA: BOTANY 



ca. 3.5 mm wide, lenticular-obovoid, distally bi- 
lobed with an apical notch 0.5-1 mm deep, pu- 
berulent along the margin. 

Veronica arvensis is found on the slopes of 
Volcan Turrialba (Khan et al. 977), but it is rarely 
collected in Central America. This species is rec- 
ognized by its short stature, leaves with subentire 
margins, very short pedicels, and blue corollas. 
The closely clustered bracts and calyx lobes give 
the inflorescences of this species a spike-like ap- 
pearance. Also, there are few intermediate leaves 
along the stem between the lower foliage leaves 
and the much narrower floral bracts in this spe- 
cies. 

Veronica peregrina L., Sp. PI. 14. 1753. V. xal- 
apensis Kunth in H.B.K., Nov. Gen. Sp. 2: 389. 
1817. V. peregrina var. xalapensis (Kunth in 
H.B.K.) Pennell, Torreya 19: 167. 1919. 

Erect or spreading herbs 5-45 cm tall, branch- 
ing mostly near the base, leafy stems 0.3-3 mm 
diam., glabrous (in var. peregrina) or with minute 
(0.2 mm) gland-tipped hairs (in var. xalapensis). 
Leaves opposite near the base and alternate dis- 
tally, gradually becoming smaller and bract-like 
distally, narrowed at the base but a petiole not 
clearly differentiated; leaf blades 8-28 mm long, 
1-8 mm wide, narrowly oblong to narrowly ellip- 
tic-oblong or oblanceolate (spatulate), apex blunt- 
ly acute or rounded, margin entire or with few 
small teeth, base gradually narrowed and cuneate, 
surfaces glabrous, venation subpalmate with a 
prominent midvein. Inflorescences of solitary 
flowers in leaf axils or flowers in the axils of re- 
duced bract-like distal leaves, spike-like with ped- 
icels ca. 1 mm long, rachis sparsely puberulent. 
Flowers with subequal or strongly unequal calyx 
lobes ca. 4 X 1 mm (-6 mm long in fruit), nar- 
rowly oblong, sparsely puberulent; corolla 2-3 
mm wide, white marked with purple lines; sta- 
mens 2-3 mm long; style ca. 0.5 mm long. Fruits 
2.8-4 mm long, 4-5 mm wide, obovoid-triangular 
with a slight indentation at the apex, compressed- 
lenticular, glabrous or puberulent. 

Plants of open moist sites, often along river- 
beds, 1000-3000 m elevation. Probably flowering 
throughout the year in northern Central America, 
but rarely collected in southern Central America. 
This species is native to southern Europe and is 
now widely naturalized in temperate climates. 

Veronica peregrina is recognized by its white 
flowers, longer erect spicate stems, and narrowly 
oblong subentire leaves. Although glabrous in Eu- 



rope, plants in the Americas range from glabrous 
to glandular puberulent (var. xalapensis). The typ- 
ical glabrous variety is more common in the east- 
ern United States, whereas var. xalapensis is most 
common in the western United States, Mexico, 
and Central America, but they intergrade, and 
both varieties can be found in a broad band across 
the central United States. The longer distal stems 
arc unusual in that there is a gradual change from 
flowers subtended by narrow leaves (to 3 cm 
long) to flowers subtended by linear bracts only 6 
mm long. 

Veronica persica Poiret in Lam., Encycl. 8: 542. 
1808. V. rotundifolia Sesse & Mocifto, Fl. Mex. 
5. 1892. 

Creeping herbs, stems 5-30(-50) cm long 
(-15 cm tall), leafy stems 0.5-2 mm diam., pu- 
berulent with thin whitish hairs 0.1-0.5 mm long, 
often in longitudinal rows. Leaves opposite near 
the base and alternate throughout the distal stems, 
petioles 1-7 mm long, puberulent; leaf blades 
ovate to ovate-orbicular, 6-24 mm long, 4-16 mm 
wide, apex obtuse to rounded, margin with 3-6 
prominent (0.5-2 mm) serrate lobes/side, base ob- 
tuse or truncate, surfaces with whitish hairs to 0.7 
mm long, venation palmate with 3 or 5 major 
veins. Inflorescences of solitary flowers in leaf 
axils (bracts absent), pedicels 10-35 mm long, 
0.2-0.3 mm diam., puberulent. Flowers with 4- 
parted calyx, sepals 4-5 mm long, subequal, to 6 
mm long in fruit, 1-1.5 mm wide at the base, 
lanceolate, surfaces glabrous but the margin cili- 
olate; corolla 8-13 mm wide, upper lobe ca. 4 
mm wide, pale blue or lilac with darker blue lines; 
filaments 2-3 mm long, anthers 0.9 mm long; 
style 1.8-3 mm long. Fruits 3.5-5 mm long, 5- 
9 mm wide, broadly obcordate with apical notch 
and divergent lateral sides, surface glabrous or 
with thin hairs, the veins becoming prominent; 
seeds ovoid, flattened, rugulose. 

Veronica persica, probably a native of the Cau- 
casus and southwestern Asia, is now naturalized 
over much of northern North America and is rare- 
ly found at higher elevations in Mexico and Cen- 
tral America. Standley reported this species as 
growing on the slopes of Volcdn Irazii in Costa 
Rica. This species is recognized by its creeping 
habit, solitary flowers in leaf axils, and larger ob- 
cordate capsules that are distinctly broader than 
long. The broad fruit causes the persisting sepals 
to spread wide as the fruit matures. This species 
has often been confused with V. polita, but that 



BURGER: FLORA COSTARICENSIS 



67 



species has smaller flowers and different fruits. 
Material cited as V. polita in Flora of Guatemala 
and Flora of Panama is actually V. persica. 

Veronica polita Fries, Novit. Fl. Suec. 63. 1819. 
Figure 2. 

Creeping or vining herbs, stems 5-70(-120) 
cm long, branching at both distal and basal nodes, 
leafy stems 0.4-1.5 mm diam., appressed puber- 
ulent with thin whitish hairs 0.1-0.2 mm long. 
Leaves opposite, petioles 4-18 mm long, 0.3-0.6 
mm wide and expanding at the blade, appressed 
puberulent; leaf blades 6-22 mm long, 6-22 m 
wide, triangular to ovate-triangular, apex acute to 
obtuse, margins with 4-8 prominent teeth 0.3-3 
mm high, base truncate to subcordate, surfaces 
with scattered stiff sharp hairs 0.1-0.7 mm long, 
venation palmate with 3 major veins. Inflores- 
cences of solitary flowers in axils of leaves or of 
short (2-4 cm) axillary racemes with narrowly el- 
liptic bracts 2-4 mm long, pedicels 3-15 mm 
long, 0.2-0.3 mm diam., puberulent. Flowers 
with calyx lobes (sepals) 4, subequal or with 1 
distinctly smaller, lobes 3-4 mm long (-5 mm in 
fruit), 1.5-2 mm wide, narrowly obovate or ob- 
ovate, surfaces subglabrous but margins with stiff 
curved hairs; corolla 4-5 mm long, less than 8 
mm wide at anthesis, blue with linear darker 
markings, filaments ca. 1.3 mm long, anthers 0.4 
mm long; styles 1.3-1.6 mm long. Fruits ca. 3 
mm long, 3-4 mm wide, obovate-triangular or 
slightly obcordate, veins usually indistinct; seeds 
obvoid, with one flattened face, rugulose to 
ridged. 

Plants of open moist slopes, damp disturbed 
sites, and partly shaded forest floor, 1400-3000 m 
elevation. Flowering and fruiting throughout the 
year. This species is a native of Europe that is now 
widely naturalized in North America and montane 
areas in Central America. 

Veronica polita is distinguished by its often tri- 
angular leaves with prominent teeth, flowers in 
axils of leaves or on short axillary racemes, well- 
developed pedicels, and blue corollas not extend- 
ing much beyond the obovate sepals. In addition, 
the stems are often long and much-branched, and 
many parts have short sharp hairs. There has been 
a controversy over the correct name for this spe- 
cies. An earlier name, V. didyma Tenore, has been 
used for this species, but many European authors 
consider that name a nomen ambiguum and reject 
it in favor of V. polita. This species is closely 
related to V. persica. 



Veronica serpyllifolia L., Sp. PI. 12. 1753. V. 
tenella All., Fl. Pedemont. 1: 75. 1785. V. hum- 
ifusa Dickson, Trans. Linn. Soc. 2: 228. 1794. 
V. serpyllifolia var. humifusa (Dickson) Vahl, 
Enum. PI. 1: 65. 1805. V. crenulata Sesse & 
Mocino, Fl. Mex. 5. 1892, non Ruiz & Pavon 
1798. Figure 2. 

Small herbs 5-20 cm tall, erect stems un- 
branched, sometimes with repent sterile leafy 
branches forming small mats, leafy stems 0.5-0.9 
mm diam., puberulent with thin ascending whitish 
hairs 0.1-0.2 mm long. Leaves opposite in the 
lower half of the stem, alternate and smaller be- 
low the flowering nodes, petioles 0.5-2 mm long 
or the leaves subsessile; leaf blades 3.5-12 mm 
long, 1.5-9 mm wide, broadly ovate to broadly 
oblong, becoming narrower distally (6X3 mm), 
apex rounded to obtuse, margin subentire to ob- 
scurely crenate with teeth 0.1-0.2 mm high, ob- 
tuse to rounded at the base, glabrous above and 
below, venation subpalmate. Inflorescences 4-15 
cm long, erect, racemes or spike-like, rachis ap- 
pressed puberulent, flowers distant or closely 
spaced, bracts 2-6 mm long, 1-3 mm wide, nar- 
rowly elliptic-oblong to narrowly ovate-elliptic, 
sessile, glabrous, pedicels 2-4 mm long (-6 mm 
in fruit), glabrous or puberulent. Flowers usually 
puberulent at the base, calyx lobes 4, subequal, 
2-4 mm long, ca. 0.9 mm wide, lanceolate, gla- 
brous or minutely puberulent; corolla ca. 4 mm 
long, lobes 2-4 mm wide, pale blue to blue-purple 
and with darker lines; styles 1.8-2.5 mm long. 
Fruits 2.3-4 mm long, 3-4.5 mm wide, obovoid 
with slightly or moderately depressed apex (ob- 
cordate), glabrous or minutely puberulent along 
the distal margins; seeds ovoid, flattened on 1 
face, smooth. 

Plants of damp open or partly shaded sites in 
montane forest formations, 2400-3300 m eleva- 
tion. Probably flowering throughout the year; it is 
common along the Carretera Interamericana in the 
Cordillera de Talamanca. This species, a native of 
western Eurasia, is now widely naturalized in 
North America and higher elevations in Mexico 
and Central America. 

Veronica serpyllifolia is recognized by its nar- 
row unbranched erect stems, small subentire 
leaves, short-pedicellate flowers, and subequal ob- 
long calyx lobes. This species is the second most 
commonly collected species of Veronica in Costa 
Rica. 



68 



FIELDIANA: BOTANY 



List of Accepted Species of Scrophulariaceae 

Key: END-CR = endemic to continental Costa 
Rica; END-CR&WP = endemic to Costa Rica 
and western Panama; END-WP = endemic to 
western Panama; INTRO = introduced weed; 
ORNAM = cultivated ornamental; ORNAM & 
NAT = cultivated and naturalized; ?? = not col- 
lected in Costa Rica but known from nearby ar- 
eas. Total number of species covered is 72; the 
number of documented native species is 48. 

Alectra aspera INTRO 
Alonsoa meridionalis 
Angelonia angustifolia ORNAM 
Anisantherina hispidula 
Antirrhinum majus ORNAM 

Bacopa axillaris 
Bacopa bacopoides ?? 
Bacopa egensis 
Bacopa laxiflora 
Bacopa monnieri 
Bacopa monnierioides 
Bacopa repens 
Bacopa salzmannii 
Bacopa sessiliflora 
Benjaminia reflexa 
Buchnera pusilla 
Buchnera weberbaueri 

Calceolaria irazuensis END-CR 
Calceolaria mexicana 
Calceolaria microbe/aria 
Calceolaria perfoliata 
Calceolaria tripartita 
Capraria biflora 
Castilleja an'ensis 
Castilleja irasuensis END-CR 
Castilleja lentil END-CR 
Castilleja quirosii END-CR&WP 
Castilleja talamancensis END-CR 
Castilleja tayloriorum END-CR 
Cymhalaria muralis INTRO 

Darcya costaricensis END-CR 
Darcva reliquiarum END-WP 
Digitalis purpurea ORNAM & NAT 

Escobedia grandiflora 
Hemichaena fruticosa 

Lamarouxia gutierrezii END-CR&WP 

Lamarouxia lanceolata 

Lamarouxia viscosa 

Leucocarpus perfoliatus 

Limosella acaulis 

Linaria canadensis ?? 

Linaria vulgaris ORNAM 

Lindernia Crustacea 

Lindernia diffusa 

Lindernia dubia 

Lophospernnim eruhescens ORNAM & NAT 

Maurandya barclaiana ORNAM 
Maurandya scandens ORNAM 



Mazus pumila INTRO 
Mecardonia prociunbens 
Micranthemum umhroxum ?? 
Mimulus glabratus '.''.' 

Penstemon gentianoitles ORNAM 

Rus.\t'lia equisctiformis ORNAM & NAT 
Russelia sa rmt-n to MI 

Schistophragma mt-.\icana 
Scoparia an nun 
Scoptiria tiulci\ 
Sibthorpia rcpena 
Stemodia angulata 
Stemodid (Inniiitifolia 
Stemodia peduncularis 
Stemodia verticilltita 

Tetranema florilntnditm END-CR 
Tetnnit-nui gamboantun END-CR 
Torcnia fournieri ORNAM 
Ton-nia thouarsii INTRO 

Veronica arvenis INTRO 
Veronica peregrina INTRO 
Veronica persica INTRO 
Veronica polita INTRO 
Veronica serpyllifolia INTRO 



SCHLEGELIACEAE Reveal 

By William Burger and Kerry Barringer 

REFERENCES W. D'Arcy, Scrophulariaceae, in 
Flora of Panama. Ann. Missouri Bot. Gard. 66: 
173-272, 1979. R. Olmstead & R Reeves, Evi- 
dence for the polyphyly of the Scrophulariaceae 
based on chloroplast rbcL and ndhF sequences. 
Ann. Missouri Bot. Gard. 82: 176-191, 1995. J. 
Reveal, Newly required suprageneric names in 
vascular plants. Phytologia 79: 68-76, 1996. 

Shrubs, subshrubs or lianas, erect or climbing, 
terrestrial or epiphytic, autotrophic, bisexual, 
stems without internal phloem, glabrous or with 
simple, eglandular or glandular, unicellular or 
multiccllular hairs, nodes lacking intcrpetiolar 
lines or glandular fields; stipules absent. Leaves 
opposite or subopposite, simple, entire (serrulate 
in Synopsis), usually coriaceous to subcoriaceous, 
glabrous or puberulent, pinnately veined. Inflo- 
rescences racemes, cymes, or of solitary flowers 
in leaf axils, sometimes in dense fascicles, brac- 
teoles present, pedicels well developed. Flowers 
bisexual, small to large, often showy, calyx irreg- 
ularly lobed, 5-lobed or split, persistent and usu- 
ally accrescent in fruit; corolla tubular to funnel- 
form, bilaterally symmetric or almost radially 



BURGER: FLORA COSTARICENSIS 



69 



symmetric, 5-Iobed, the tube straight (not saccate 
or spurred), stamens 4, borne on the corolla tube, 
filaments tree, alternating with the corolla lobes, 
anthers 2-thecous, thecae equal, distinct, a stami- 
node present, disc absent; pistil solitary, ovary su- 
perior. 2-locular, ovules many, unitegmic, borne 
on 2 large axile placentas, the style 1, terminal, 
stigmas simple. Fruits baccate, indehiscent; seeds 
many, with reticulate exotesta, the embryo small, 
straight. 

Schlegeliaceae is a family of four Neotropical 
genera and about 30 species. The genera Gibson- 
iothamnus and Schlegelia are found in Costa Rica, 
and they are closely related. Until recently the 
genera of this family were included in either the 
Scrophulariaceae or Bignoniaceae but were atyp- 



ical for either family. Evidence from chloroplast 
DNA sequences (Olmstead & Reeves, 1995, cited 
above) indicates that the family is not closely re- 
lated to either of these families but is a distinct 
group, probably most closely related to the basal 
Verbenaceae, such as Callicarpa. In recognition 
of their distinctiveness, Reveal (1996, cited 
above) established the family Schlegeliaceae for 
this group. Our representatives are distinguished 
from most Bignoniaceae by their simple entire 
leaves and baccate fruits. The family also differs 
from many bignons in lacking tendrils, interpetio- 
lar lines, and interpetiolar gland fields on the 
stems. The Schlegeliaceae are distinguished from 
most Scrophulariaceae by their woody habit and 
baccate-indehiscent fruits. 



Key to the Genera of Schlegeliaceae in Costa Rica 

la. Epiphytic shrubs; corollas usually reddish to purple (less often white), tubular and almost radially 
symmetric; staminodes < 5 mm long; fruits with fleshy covering; leaves usually 4-8 cm long . . . 
Gibsoniothamnus 

Ib. Lianas or rarely small trees or epiphytic shrubs; corollas white to pinkish (in Costa Rica), tubular- 
campanulate and usually clearly bilaterally symmetric; staminodes usually > 5 mm long; fruits with 

a hard outer covering; larger leaves usually > 10 cm long (except in 5. brachyantha) 

Schlegelia 



Gibsoniothamnus L. O. Williams 

REFERENCES W. D'Arcy, Gibsoniothamnus 
(Scrophulariaceae) in Flora of Panama, Part 9. 
Ann. Missouri Bot. Card. 66: 220-227. 1979. A. 
Gentry, Note on Gibsoniothamnus. Fieldiana, Bot. 
34: 55. 1971. A. Gentry, Gibsoniothamnus (Scro- 
phulariaceae) in Panama. Ann. Missouri Bot. 
Gard. 61: 533-537. 1974. A. Gentry, New or 
noteworthy species of middle American Bigno- 
niaceae. Wrightia 7: 83-89. 1982. P. C. Standley 
& L. O. Williams, Gibsoniothamnus (Scrophular- 
iaceae) in Flora of Guatemala. Fieldiana, Bot. 24: 
356-359. 1973. L. O. Williams, An overlooked 
genus of the Scrophulariaceae. Fieldiana, Bot. 32: 
211-214. 1970. 

Shrubs or small trees, epiphytic or rarely ter- 
restrial, to 6 m tall, young stems terete or angular, 
glabrous or pubescent, leaf base persistent and be- 
coming conspicuously raised, older stems with a 
light-colored bark. Leaves opposite or suboppos- 
ite, equal or unequal (anisophyllous) at a node, 
clearly articulated at the base, petioles usually 
short, merging with the base of the blade, blades 
simple and entire, often elliptic, coriaceous or 



subcoriaceous (rarely chartaceous), glabrous or 
puberulent, usually with glandular pits scattered 
on the underside, domatia often present in the ax- 
ils of veins and midvein below. Inflorescences of 
solitary axillary flowers or flowers in terminal 
cymes, panicles, racemes, or condensed and fas- 
ciculate, peduncles present or absent, pedicels 
elongate, merging with the calyx and often the 
same color as the calyx, usually bracteate. Flow- 
ers usually colorful throughout, calyx cupulate or 
campanulate, glabrous or puberulent, lobes usu- 
ally 5, narrow and tooth-like to elongated and lat- 
erally compressed or winged, persistent and often 
accrescent in fruit; corolla tubular to narrowly 
funnelform, often brightly colored in purples and 
reds, radially symmetric or slightly 2-lipped, gla- 
brous externally or ciliolate along the distal edges, 
lobes 5, rounded and subequal, much shorter than 
the tube, tube interior with hairs near the stamen 
insertion; stamens 4, included, equal or in 2 sub- 
equal pairs, inserted above the base of the tube, 
filaments filiform, bent near the apex, anthers with 
a broad connective and 2 separate parallel or 
slightly divaricate thecae, staminode 1, usually 
shorter than the stamens; disc lacking; ovary 2- 



70 



FIELDIANA: BOTANY 



Gibsoniothamnus 
pterocalyx 



Schlegelia 
parvi flora 



Schlegelia 
fasti giata 




FIG. 9. Schlcgcliaceae: epiphytic shrubs, vines, or trees with berry-like fruits; species of Gibsoniothamnus and 
Schlegelia. 



BURGER: FLORA COSTARICENSIS 



71 



locular, ovules many, style slender, longer than the 
stamens, stigma capitate. Fruits fleshy berries, 
globose to subglobose, juicy or mucilaginous; 
seeds many, seed coat reticulate, the margins or 
the reticulae bearing elongate filaments that are 
sticky or mucilaginous; endosperm absent in ma- 
ture seeds, embryo straight. 

Gibsoniothamnus is a genus of 12 species na- 
tive to southern Mexico, Central America, and ad- 
jacent Colombia. The paucity of earlier collec- 
tions resulted in D'Arcy's broad specific concepts 



(1979), but additional collecting appears to justify 
the recognition of a larger number of geographi- 
cally isolated species. The domatia vary from 
tufts of hairs in vein axils to well-defined pits or 
pockets in the lower leaf surface; they may be 
lacking in some species. Our species are less pu- 
bescent and have more prominent calyx lobes 
than does G. cornutus (J. D. Smith) A. Gentry, 
which is restricted to Mexico and Guatemala and 
probably includes G. pithecobius (Standl. & Stey- 
erm.) L. O. Williams. 



Key to the Species of Gibsoniothamnus 

la. Flowers usually borne in distal fascicles of 5-25 flowers; leaf blades elliptic-oblong to obovate, to 
5 cm wide 2 

Ib. Flowers 1-3 in the axils of leaves or terminal; leaf blades narrowly elliptic to ovate or oblanceolate, 

to 3 cm wide 3 

2a. Leaf blades to 8 cm long and to 3 cm wide, the major veins deeply impressed above when 

dried; young stems usually with stiff straight hairs; flowers in fascicles of 5-1 1 

G. epiphyticus 

2b. Leaf blades to 15 cm long and to 6 cm wide, the major veins not deeply impressed when dried; 
young stems glabrous or with few hairs; flower in fascicles of 10-25 G. parvifolius 

3a. Corolla red or purple; sepal lobes and veins not expanded laterally to form winged margins, calyx 
drying blackish or yellowish; 1000-1500 m elevation in evergreen (rarely, deciduous) forests of 
the Pacific slope G. parvifolius 

3b. Corolla lavender, rose, or white, sepal lobes expanded laterally from the veins and somewhat 
winged, often drying yellowish; 300-1400 m elevation in evergreen forests on the Caribbean slope 
and along the Continental Divide 4 

4a. Corolla lilac or rose; calyx lobes only slightly expanded along the veins, leaves to 70 mm long; 
400-900 m elevation on the Caribbean slope G. pterocalyx 

4b. Corolla white, calyx lobes expanded laterally along the midveins and conspicuously winged, leaves 
to 25 mm long; 1200-1400 m elevation on Cerro Colorado, Panama G. stellatus 



Gibsoniothamnus epiphyticus (Standl.) L. O. 
Williams, Fieldiana, Bot. 34: 120. 1972. Cler- 
odendrum epiphyticum Standl., Publ. Field 
Mus. Nat. Hist. Bot. Ser. 22: 168. 1940. Figure 9. 

Epiphytic shrubs to ca. 1 m tall, leafy branches 
2-4 mm thick, terete to slightly angulate below 
the raised leaf bases, with stiff straight hairs 0.3- 
1 mm long but glabrescent. Leaves opposite or 
subopposite, usually of 2 different sizes at each 
node (anisophyllous), petioles 4-9 mm long, 0.8- 
1 .7 mm thick, puberulent or glabrous; leaf blades 
2-8 cm long, 1.2-3 cm wide, elliptic to elliptic- 
oblong or ovate-elliptic, apex acuminate to acute 
or rounded, margin entire and reflexed, base cu- 
neate to acute, subcoriaceous, lustrous above, gla- 
brous or with few straight hairs to 1 mm long, 
midvein impressed above, 2 veins 2 or 3/side, 



domatia in vein axils beneath. Inflorescences of 
5-1 1 flowers in short axillary panicles or from a 
fasciculate base near the apex of a branchlet, 1 
peduncles to 7 mm long, bracts 2-4 mm long, 
narrowly triangular, pedicels 15-20 mm long, 
merging gradually with the calyx, glabrous or 
with straight hairs to 0.8 mm long. Flowers with 
calyx 8-1 1 mm long, 3-4 mm diam., glabrous or 
slightly puberulent on the angles, lobes 4-6 mm 
long, 1-1.5 mm wide at the base; corolla 14-21 
mm long, 3-4 mm diam., tubular, glabrous, red- 
violet or purple, lobes 1.5-2.5 mm long; stamens 
attached 3-4 mm from base of tube, filaments 18- 
20 mm long; staminode 4-5 mm long; pistil gla- 
brous, style 19-22 mm long. Mature fruits to 1 
cm diam., becoming white. 

Rarely collected plants of evergreen montane 
cloud forest formations of the Caribbean slope, 



72 



FIELDIANA: BOTANY 



600-1500 m elevation. Flowering in April-July. 
This species is known only from central Costa 
Rica. 

Gibsoniothamnus epiphyticus is recognized by 
its epiphytic habit, stiff leaves with deeply im- 
pressed venation, fasciculate flowers, prominent 
winged calyx lobes, and red or purple tubular co- 
rollas. The stiff multicellular hairs are distinctive 
when present, but the pubescence is often decid- 
uous. This species is similar to G. pterocalyx but 
differs from that species in having narrower, sub- 
coriaceous leaves with veins impressed above, 
red-violet corollas, and shorter, more triangular 
calyx lobes. G. mirificus A. Gentry of central Pan- 
ama is also similar but that species has solitary 
flowers, longer calyx lobes, and thinner leaves. 

Gibsoniothamnus parvifolius Barringer (Novon 
9: 476. 1999). Figure 9. 

Epiphytic shrubs to 2 m tall, leafy stems 1-7 
mm thick, glabrous to sparsely pubescent with 
hairs 0.2-0.6 mm long, winged or angled in early 
stages but soon terete and glabrescent. Leaves 
equal or sometimes strongly unequal at a node 
(with the smaller leaves more rounded), petioles 
2-15 mm long, 0.4-2 mm thick, glabrous; leaf 
blades 1.6-12 cm long, 0.6-5 cm wide, blades 
lanceolate to narrowly elliptic or obovate-elliptic, 
apex acuminate, margin entire and reflexed on 
drying, base acute or cuneate and decurrent on the 
petiole, drying subcoriaceous, lustrous and dark 
olive-green above, glabrous above, glabrous or 
with thin hairs beneath, 2 veins usually 2-5/side, 
impressed above, pit domatia often present near 
the vein axils beneath. Inflorescences of solitary 
flowers in distal leaf axils (2/node), bracteoles ca. 
1 mm long, linear, pedicels 4-30 mm long, 0.3- 
1.5 mm thick, glabrous or with few glandular 
hairs in early stages, bracteolate at base. Flowers 
glabrous externally, calyx campanulate, tube ca. 4 
mm long with truncated distal margin, lobes 4- 
10 mm long, 0.5-1 mm wide, narrowly triangular, 
glabrous, spreading, drying dark; corolla 18-38 
mm long, 1.5-5 mm diam., tubular, reddish violet 
to wine-red or red, lobes 2-4 mm long, upper 
lobes broadly ovate, lateral and median lobes 
ovate, usually more darkly colored than the tube; 
filaments 15-26 mm long, staminode 1.7-5 mm 
long; pistil glabrous, style elongate, ca. 18 mm 
long. Fruits becoming 10 mm in diam., globose 
or oblate, fleshy, white at maturity. 

Plants of evergreen cloud forest formations in 
the Chiriqui highlands and in the Cordillera de 



Guanacaste and Cordillera de Tilaran, at 500- 
1500 m elevation. Flowering in February and Au- 
gust; fruiting in August. This species is known 
only from Costa Rica and adjacent Panama. 

Gibsoniothamnus pan'ifolius is distinguished 
by its often small glabrous leaves, solitary axil- 
lary flowers, narrow calyx lobes, slender tubular 
purplish or reddish corollas, and white fleshy 
fruits. The specimens from northern Costa Rica 
tend to have more angled stems and shorter sta- 
minodia than the Panamanian specimens. This 
species can be distinguished from G. epiphyticus 
of central Costa Rica by its small obovate or lan- 
ceolate leaves, its calyx lobes more than 5 mm 
long, and its one- or two-flowered inflorescences. 
The larger-leaved collections resemble G. gran- 
diflorus A. Gentry & Barringer of central Panama 
(see Fig. 9). Two Costa Rican collections were 
provisionally placed under G. grandiflorus. but 
they are now placed under an expanded circum- 
scription of G. parvifolius; they are G. Herrera 
3473 (MO) and J. D. Smith 6730 (us). 

Gibsoniothamnus pterocalyx A. Gentry; Ann. 
Missouri Bot. Gard. 61: 535. 1974. Figure 9. 

Epiphytic shrubs to 2 m tall, leafy stems 0.7- 
3 mm thick, glabrous, at first slightly angled but 
soon terete and pale grayish. Leaves opposite or 
subopposite, unequal or subequal at a node, artic- 
ulated at the base, petioles 2-13 mm long, 0.4- 
0.8 mm thick, glabrous; leaf blades 1.7-7 cm 
long, 1.5-3.4 cm wide, narrowly elliptic to nar- 
rowly obovate or oblanceolate, apex bluntly acute 
or rounded and slightly emarginate, margin entire 
and slightly revolute, base acute to cuneate, dry- 
ing subcoricaceous to chartaceous, dark olive 
green and lustrous above, glabrous, 2 veins 2 or 
3/side, strongly ascending; pit domatia often pre- 
sent near vein axils beneath. Inflorescences of 
solitary axillary flowers or few flowers in con- 
densed axillary panicles, 1 peduncles to 1 cm 
long, bracts 2-3 mm long, narrowly triangular, 
sparsely villous, pedicels 7-25 mm long, 0.4-0.8 
mm thick, glabrous, merging gradually with the 
base of the calyx. Flowers pale lilac or rose, gla- 
brous externally, calyx 8-12 mm long, 2-4 mm 
diam., calyx lobes 5-9 mm long, 0.4-1.3 mm 
wide, linear and slightly winged at the base; co- 
rolla 18-25 mm long, lilac or pink, glabrous, tube 
3-5 mm diam., lobes 2-3 mm long, rounded; fil- 
aments 1 5-20 mm long, staminode 5-6 mm long, 
filiform; ovary glabrous, style 16-19 mm long. 



BURGER: FLORA COSTARICENSIS 



73 



Fruits becoming up to 10 mm diam., obovoid to 
oblong with truncated apex, white. 

Infrequently collected plants of evergreen low- 
er montane rain forest formations on the Carib- 
bean slope, 400-900 m elevation (intensive col- 
lecting over many years at Monteverde indicates 
that the upper altitudinal limit of 900 m is real, 
and not an artifact of insufficient collecting). 
Flowering in April-July and October. This species 
ranges along the Caribbean escarpment from 
northern Costa Rica to western Panama. 

Gibsoniothamnus pterocalyx is recognized by 
its epiphytic habit, glabrous parts, small narrow 
lustrous leaves, usually only one to three pink or 
lilac (white) flowers at a node, and prominent se- 
pal lobes. The leaf undersides are often purplish, 
especially along the veins. This species is super- 
ficially similar to G. parvifolius (q.v.). 

Gibsoniothamnus stellatus A. Gentry & Bun m- 
ger, Novon 5: 121-122, 1995. 

Epiphytic shrubs, stems angled and with raised 
leaf bases, leafy stems ca. 2 mm thick, glabrous, 
becoming grayish. Leaves of the same node un- 
equal or subequal, petioles 3-5 mm long, gla- 
brous, without lateral wings; leaf blades 5-25 
mm long, 4-16 mm wide (larger blades 15-25 X 
9-16 mm), ovate-lanceolate to elliptic, apex 
acute, base cuneate, drying coriaceous, glabrous, 
punctate; 2 veins 2 or 3/side, domatia sometimes 
present near the base beneath. Inflorescences of 
solitary flowers axillary to distal leaves, bracteate 
at the base, pedicels 10-12 mm long, expanded 
distally, glabrous. Flowers with campanulate pur- 
plish calyx, glabrous, tube 6-8 mm long, lobes 8- 
9 mm long, triangular with wings 3-4 mm wide; 
corolla 20-25 mm long, 4-5 mm diam., tubular, 
white, upper and lateral lobes 3-4 mm long and 
wide, ciliolate along the edge; filaments 17-18 
mm long, staminode 5-6 mm long; pistil gla- 
brous. Fruits not seen at maturity. 

Gibsoniothamnus stellatus is distinguished by 
its shrubby epiphytic habit, small leaves, flowers 
with winged calyx lobes (giving a star-like form), 
and white corollas. This species is known only 
from two collections from 1200-1400 m elevation 
on Cerro Colorado in the provinces of Bocas del 
Toro and Chiriqui in western Panama. They were 
flowering in April and July. This species is closely 



related to G. alatus of eastern Panama and adja- 
cent Choc6, which also has white corollas and 
winged calyx lobes but has much larger leaves. 



Schlegelia Miquel 

REFERENCES A. Gentry, Schlegelia (Bignoni- 
aceae) in Flora of Panama. Ann. Missouri Bot. 
Card. 60: 923-930, 1973. P. Standley & L. O. 
Williams, Schlegelia (Scrophulariaceae) in Flora 
of Guatemala. Fieldiana Bot. 24, pt. 9: 396-400, 
1973. 

Shrubs, woody vines, lianas or small trees, ter- 
restrial or epiphytic, wood without anomalous 
vasculature, stems usually terete; pseudostipules 
small, usually appressed against the branchlets. 
Leaves opposite, simple, with thick terete peti- 
oles, usually becoming articulated at the base, 
blades often coriaceous, margins entire, venation 
pinnate. Inflorescences terminal or axillary pan- 
icles, fascicles or axillary racemes, peduncles usu- 
ally woody, often with conspicuous small bracts 
beneath the pedicels. Flowers with a subcoria- 
ceous tubular or cupulate calyx, entire or splitting 
irregularly into small lobes, persisting and split- 
ting in fruit; corolla campanulate to narrowly tu- 
bular, white to pink or red (yellow), glabrous ex- 
ternally but glandular lepidote on the lobes within, 
lobes 5, rounded; stamens 4, in 2 unequal pairs, 
borne on the lower half of the corolla tube, in- 
cluded, anthers glabrous, thecae divaricate, a 
small staminode usually present; ovary 2-locular 
(sometimes 1- locular near the apex), placenta 
central on the septum in each locule, ovules many, 
style simple, stigma bifid (trifid). Fruits berries 
with hard crustaceous indehiscent pericarp, usu- 
ally globose; seeds narrow and angular, wingless, 
small, embedded in pulp. 

Schlegelia is a genus of ca. 15 species, ranging 
from eastern Mexico and the West Indies to Bra- 
zil. The simple, opposite, usually stiff leaves, 
shrubby, often epiphytic or vining habit, usual 
lack of conspicuous pubescence, woody inflores 
cence branches, calyx tubes with poorly devel- 
oped lobes, and round berry-like fruits with stiff 
rind help distinguish this genus. Dr. William 
D'Arcy (MO) is currently studying this genus. 



74 



FIELDIANA: BOTANY 



Key to the Species of Schlegelia 

la. Corollas usually < 20 mm long; calyx tube 4-6 mm long at anthesis; fruits up to 12 mm diam.; 
leaves usually drying stiffly coriaceous and with the minor venation usually not evident on the 

upper surface; plants often epiphytic; 2-2100 m elevation 2 

Ib. Corollas 20-43 mm long; calyx 5-12 mm long; fruits 12-40 mm diam.; leaves drying stiffly 
chartaceous to subcoriaceous, the minor veins often elevated on the upper surface; plants usually 

vines or lianas; 10-900(-1500) m elevation 3 

2a. Corollas mostly 8-1 1 mm long; leaf blade acute to obtuse or rounded at apex; commonly collected 

S. parviflora 

2b. Corollas usually 15-20 mm long; leaf blade obtuse to rounded at apex; rarely collected 

S. brachyantha 

3a. Corolla 20-28 mm long, narrowly tubular with short (1-3 mm) lobes; fruits 12-16 mm diam.; 
inflorescences of many (> 15) flowers in dense woody fascicles or panicles, peduncles 1-20 cm 

long S. fastigiata 

3b. Corolla 30-43 mm long, tubular-campanulate with prominent (to 10 mm) lobes; fruits 30-40 
mm diam.; inflorescences of few (1-5) flowers in racemose arrangements, peduncles usually < 
1 cm long S. nicaraguensis 



Schlegelia brachyantha Griseb., Cat. PI. Cub. 
191. 1866. 

Lianas or vine-like shrubs to 6 m high, leafy 
stems 1-8 mm thick, glabrous or sparsely puber- 
ulent near the apex and on the new growth, drying 
grayish or dark, terete or slightly quadrangular. 
Leaves with petioles 3-9 mm long, 1-2.5 mm 
thick, becoming articulated and thicker at the 
base, glabrous or with few minute (0.1-0.2 mm) 
hairs; leaf blades 4-12 cm long, 1.5-8 cm wide, 
broadly elliptic to broadly oblong or oblong-or- 
bicular, apex bluntly obtuse to rounded or short- 
acuminate, base obtuse to rounded and subtrun- 
cate, drying subcoriaceous and slightly lustrous 
above with margin often recurved, glabrous above 
and below, punctate beneath, major 2 veins 3-57 
side, loop-connected distally. Inflorescences ax- 
illary to leaves or leafless nodes, apparently ses- 
sile fascicles with few (2-5) flowers, peduncles 
very short with bracts ca. 2 mm long on congested 
nodes, pedicels 2-8 mm long, glabrous and dry- 
ing black. Flowers not seen at anthesis, flower 
buds 3-5 mm long, with few minute hairs, drying 
dark; corolla 15-20 mm long, white or pink, 5- 
6 mm wide at throat, lobes ca. 5 mm long. Fruits 
6-10 mm diam., green turning red. 

Rarely collected plants of evergreen forest for- 
mations, 20-1800 m elevation. In Costa Rica it is 
only known from the high plateau southeast of 
Chatham Bay, Cocos Island, flowering and fruit- 
ing in April (R. Foster 4125 F). Also known from 
the Chiriqui highlands in Panama, Venezuela, and 
the West Indies. 



Schlegelia brachyantha is distinguished by its 
smaller, often rounded leaves, few-flowered, ses- 
sile inflorescences, and few collections from 
widely separated geographical areas. 

Schlegelia costaricensis Stand!, described in 
the Flora of Costa Rica, part 3 (Publ. Field Mus. 
Nat. Hist. Bot. Ser. 18: 1128, 1938) has been 
transferred to the Boraginaceae as Bourreria cos- 
taricensis (Standl.) A. Gentry (see Phytologia 26: 
67-68, 1973). 

Schlegelia fastigiata Schery, Ann. Missouri Bot. 
Gard. 29: 367. 1942. Figure 9. 

Lianas or vines, climbing stems ca. 15 mm 
diam., leafy stems 1.8-6 mm thick, terete or 
somewhat flattened, glabrous (minutely puberu- 
lent). Leaves with petioles 3-8 mm long, 1 .5-3.5 
mm thick, glabrous, usually drying dark and ar- 
ticulated at the base; leaf blades 10-26(-33) cm 
long, 3-9(-13) cm wide, narrowly elliptic-oblong 
to narrowly ovate-oblong, apex acuminate or 
acute, base obtuse, drying stiffly chartaceous to 
subcoriaceous, glabrous above and below, the mi- 
nor venation slightly elevated on the upper sur- 
face (dried), 2 veins 8-16/side. Inflorescences 
congested woody panicles borne on short (1-2 
cm) peduncles or longer (8-14 cm) leafless stems, 
usually with more than 15 flowers, woody pani- 
cles (fascicles) 2-8 cm long, 2-10 cm wide, 
woody branches short and grayish, pedicels 4-8 
mm long, ca. 0.4 mm thick, minutely puberulent, 
drying black, subtended by a pair of small (1 mm) 
bracts on a slender 2 peduncle 2-4 mm long. 



BURGER: FLORA COSTARICENSIS 



75 



Flowers with truncated calyx tube 5-8 mm long, 
cupular with entire or slightly lobed distal margin, 
5-7 mm diam. at apex, purple to red-violet or 
pink, usually glabrous, splitting in fruit and to 14 
mm wide; corolla 20-28 mm long, tubular, gla- 
brous externally, 4-6 mm diam., 4-8 mm wide at 
the spreading lobes, white, lobes 2-5 mm long, 
rounded, pink to pale violet; filaments 8-12 mm 
long, anthers 1-1.5 mm long, divaricate, ca. 2 mm 
wide; ovary lepidote (Gentry, 1973b). Fruits 12- 
1 6 mm long, subglobose, yellow to orange or pur- 
ple, surface somewhat muriculate, glabrous; seeds 
ca. 2 X 0.5 mm, narrowly triangular or curved. 

Plants of the evergreen rain forest formations 
of the Caribbean lowlands and slopes, 10-900 m 
elevation. Probably flowering and fruiting 
throughout the year, but our collections were col- 
lected between July and March. The species rang- 
es from Guatemala to Ecuador. 

Schlegelia fasti giata is recognized by its vining 
habit, thinner leaves with more 2 veins than our 
other species of Schlegelia, flowers borne in 
densely branched, woody fascicles on slender 
bracteate pedicels, calyx cup with entire margin, 
and narrowly tubular white corolla with pink 
lobes. The minor venation is usually clearly ele- 
vated on the upper leaf surface of the dried leaves, 
a helpful characteristic in making specific deter- 
minations. This species was incorrectly placed un- 
der S. sulphurea Diels in Flora of Panama, mis- 
spelled as 5. sulfurea (Gentry, 1973b); that spe- 
cies has yellow corollas and is endemic to eastern 
Ecuador. 

Schlegelia nicaraguensis Standl., Trop. Woods 
16: 44. 1928. 5. silvicola L. O. Williams, Fiel- 
diana, Bot. 34: 126, tab. 3. 1972. 

Lianas or woody vines (rarely trees or epi- 
phytic shrubs), climbing stems ca. 15 mm diam., 
leafy stems 2-6 mm thick, terete or 4-angled, mi- 
nutely puberulent in early stages, glabrescent, 
grayish and with prominent lenticels; pseudosti- 
pules to 3 mm long. Leaves with petioles 3-14 
mm long, 1.5-4.2 mm thick, rounded but with 2 
adaxial ridges, glabrous, drying grayish or yel- 
lowish; leaf blades (4-)8-22 cm long, (2-)3.5- 
10 cm wide, elliptic to elliptic-oblong, elliptic-ob- 
ovate or elliptic-ovate, apex acuminate to acute or 
rounded, base obtuse to acute, drying subcoria- 
ceous and grayish, glabrous above and below or 
with a few hairs along the midvein beneath, minor 
venation flat or slightly elevated above, 2 veins 
5-8/side. Inflorescences with 1-5 flowers, ter- 



minal, axillary, or borne on older stems, race- 
mose, peduncles 6-10 mm long from a short (1- 
6 mm) woody base, rachis with small (1-2 mm) 
bracts, pedicels 8-20 mm long, ca. 0.8 mm thick, 
minutely (0.1-0.2 mm) puberulent. Flowers with 
cupulate or campanulate calyx 9-12 mm long, 7- 
10 mm diam., margin entire or with short (1-2 
mm) rounded lobes, glabrous or with few minute 
hairs near the base, purple or violet with white 
spots in life; corolla 33-42 mm long, tubular- 
campanulate and bilabiate, yellowish white or 
white, with rose or purplish lobes, tube 3-4 mm 
diam. at base and widening to the mouth, lobes 
6-10 mm long, with rounded apices; stamens 16- 
23 mm and 14-18 mm long, anthers divaricate, 
thecae ca. 3 mm long, staminode 3-5 mm long; 
ovary unilocular near the apex. Fruits 4-5 cm 
long, 3-4 cm diam., obovoid to globose with nar- 
rowed base, surface lustrous and slightly muri- 
culate, drying brown; seeds ca. 3 mm long, 0.8- 
1 mm wide in the center. 

Uncommon vines of evergreen rain forest for- 
mations along the Caribbean slope, 100-600 m 
elevation (to 1500 m in Nicaragua). Probably 
fruiting throughout the year (Costa Rican flower- 
ing collections have been made only in October 
and December-February). This species ranges 
from Veracruz, Mexico, to Panama. 

Schlegelia nicaraguensis is recognized by its 
woody vining habit, subcoriaceous leaves, few- 
flowered inflorescences, large bilabiate corollas, 
and large fruits. This species is similar to 5. par- 
asitica Sw. of Jamaica (Gentry, 1973b). 

Schlegelia parviflora (Oerst.) Monachino, Phy- 
tologia 3: 103. 1949. Dermatocalyx parviflorus 
Oerst., Vidensk. Meddel. Dansk Naturhis. For- 
en. Kjobenhavn 1855: 29. 1856. S. fuscata A. 
Gentry, Ann. Missouri Bot. Gard. 60: 925. 1973 
(1974). Figure 9. 

Epiphytic shrubs or less often (in Costa Rica) 
woody vines, lianas, or small trees, stems slightly 
quadrangular or terete, leafy stems 2-7 mm thick, 
glabrous or sparsely puberulent at the shoot apex; 
pseudostipules 2-3 mm long. Leaves with peti- 
oles 6-30 mm long, 0.7-3 mm thick, becoming 
articulated at the base and thickened, glabrous; 
leaf blades (4-)6-18 cm long, (2-)3-13 cm wide, 
broadly elliptic to ovate-elliptic or ovate-oblong, 
apex bluntly acute to obtuse or rounded, base ob- 
tuse to cuneate and often slightly decurrent on the 
petiole, drying coriaceous and yellowish gray, 
glabrous above and below, minutely punctate be- 



76 



FIELDIANA: BOTANY 



neath, 2 veins 4-8/side, minor venation usually 
obscure above. Inflorescences axillary to leaves 
or from older leafless nodes, fascicles of 5-40 
flowers in racemose or paniculate arrangements, 
2-5 cm long (to 8 cm in fruit), 1 peduncles 1- 
10 mm long, 1-2 mm thick, glabrous, usually dry- 
ing yellowish, bracts 1-2 mm long, subulate, ped- 
icels 1-4 mm long (measured from the subtending 
bracteoles), becoming 1.5 mm thick in fruit. 
Flowers with calyx 4-6 mm long, 3-4 mm wide, 
cupulate to campanulate, glabrous, lobes 0.5-1.5 
mm long, rounded distally; corolla 8-1 1 mm 
long, to 10 mm wide across the lobes, white with 
pink-tinged lobes, glabrous externally, lobes 2-4 
mm long, with purple lines within; filaments ca. 
4 mm long, thecae 1.3 mm long, lilac; style ca. 6 
mm long. Fruits 8-12 mm diam., globose or 
somewhat oblate, becoming purple or wine-red, 
surface lustrous and slightly muriculate, subtend- 
ing calyx to 10 mm wide, pale green; seeds 2-3 
mm long, 0.5-1 mm thick, usually wider at one 
end, drying black. 

Common woody plants in wet evergreen forest 
formations on both the Caribbean and Pacific 
slopes, 4-2200 m elevation. Probably flowering 
and fruiting in all months, but with lowland plants 
usually flowering in December-February and 
highland plants with peak flowering in June-Au- 
gust. This species ranges from Belize to Brazil. 

Schlegelia parviflora is recognized by its usu- 
ally epiphytic habit (also vines or trees), stiffly 
coriaceous leaves, lack of pubescence, fascicles of 
flowers on few-branched axes, cupulate or cam- 
panulate calyx, white corolla with pink and purple 
coloring, and subglobose purplish fruits. The low- 
er elevation collections tend to have larger leaves, 
but there is great variation in leaf size, and a cline 
is not evident. The flowers and fruits of this spe- 
cies are distinctly smaller than those of its sym- 
patric congeners. A very few Costa Rican collec- 
tions fit the description of S. fuscata, with smaller 
flowers drying black on consistently racemose 
axes. However, such collections seem to be no 
more than extreme variants (perhaps plants of ex- 
posed windy sites) in a species that exhibits a 
broad range of variation in leaf and inflorescence 
morphology. 



BIGNONIACEAE 

By William Burger and Alwyn Gentry (t) 

REFERENCES A. Gentry, Bignoniaceae in Flora 
of Panama. Ann. Missouri Bot. Gard. 60: 781- 



9771. 1973 (1974). A. Gentry, Co-evolutionary 
patterns in Central American Bignoniaceae. Ann. 
Missouri Bot. Gard. 61: 728-759. 1974. A. Gen- 
try, Bignoniaceae of southern Central America; 
distribution and ecological specificity. Biotropica 
8: 117-131. 1976. A. Gentry, Bignoniaceae, Part 
I (Crescentieae and Tourrettieae). Fl. Neotrop. 
Monogr. 25(1): 1-130. 1980. A. Gentry, Evolu- 
tionary patterns in Neotropical Bignoniaceae. 
Mem. New York Bot. Gard. 55: 118-129. 1990. 
A. Gentry, Bignoniaceae Part II (tribe Teco- 
meae). Fl. Neotrop. Monogr. 25(11): 1-370. 1992. 
P. C. Standley & L. O. Williams, Bignoniaceae in 
Flora of Guatemala. Fieldiana, Bot. 24(10): 153- 
232. 1974. 

Trees, shrubs, or more often lianas (rarely 
herbs or twining herbs), often climbing with the 
aid of twisting stems or tendrils, the tendrils sim- 
ple or distally trifid (with sticky pads in Mansoa 
parvifolia), often becoming woody, stems terete 
or angulate, glabrous or pubescent, often with in- 
terpetiolar ridges or gland fields at the nodes; stip- 
ules absent, pseudostipules present or absent. 
Leaves opposite or whorled (alternate in a few 
genera), simple, 2-foliolate, 3-foliolate, pinnately 
compound or palmately compound (occasionally 
twice compound as in Jacaranda, Pleonoioma, 
and Tourrettia), petiolate. the distal leaflet often 
replaced by a tendril in climbing species, blades 
usually entire, sometimes serrate (not lobed), gla- 
brous to densely puberulent, venation palmate or 
pinnate, domatia rarely present. Inflorescences 
terminal, axillary, or cauliflorus, cymose, panic- 
ulate, racemose, fasciculate, or of solitary flowers 
in leaf axils, bracts and bracteoles often present, 
small and caducous, pedicels usually well devel- 
oped. Flowers bisexual, nearly always large and 
showy, calyx united and tubular to campanulate, 
usually with 5 lobes or teeth (rarely entire, bila- 
biate, calyptrate, or spathe-like); corolla united 
and funnelform to campanulate with a basal tube, 
usually 2-lipped and bilaterally symmetrical (rare- 
ly radially symmetric), the 5 lobes imbricate in 
bud (rarely valvate) and usually rounded distally; 
stamens usually 4 (rarely 5 or 2), alternate with 
corolla lobes, filaments usually of 2 unequal pairs, 
borne on the lower half of the tube and free, an- 
thers 2-thecous (1-thecous), free, a staminode usu- 
ally present, an annular disc usually present; ova- 
ry superior, 2-locular with 2 axile placentas or 
unilocular with 2-4 intruded parietal placentas 
(rarely 4-locular as in Tourrettia), style simple, 
stigma 2-lobed, rounded. Fruits mostly 2-valved 



BURGER: FLORA COSTARICENSIS 



77 



capsules with septicidal or loculicidal dehiscence 
(fruits fleshy or hard and indehiscent in a few gen- 
era); seeds mostly flat and winged in capsular 
fruits, often embedded in pulp or mucilage in in- 
dehiscent fruits, lacking endosperm, cotyledons 
leaf-like. 

The Bignoniaceae are a family of about 112 
genera with over 800 species in tropical and tem- 
perate areas of the world but with the greatest 
concentration of genera and species (ca. 600) in 
the Neotropics. The family is often easy to iden- 
tify because of its woody stems, large showy five- 
lobed and two-lipped corollas, androecium of usu- 
ally four functional stamens with filaments of two 
lengths, superior ovary with many ovules, and 
usually capsular fruits with thin-winged seeds. 
Gland fields are sometimes present at the nodes, 
on the petioles, or at the base of the leaf blade. 
The stems are woody (except in Tourettia) and 
often climbing; many species have opposite com- 
pound leaves with entire margins. Gibsoniotham- 
nus and Schlegelia, often assigned to Bignoni- 
aceae, have now been placed in the family Schle- 
geliaceae (q.v.). On the basis of floral morphology 
the Bignoniaceae were thought to be closely re- 
lated to the Gesneriaceae, Pedaliaceae, and Scro- 
phulariaceae. However, recent DNA studies also 
suggest relationships with Buddleiaceae and Ver- 
benaceae (Olmstead & Reeves, 1995). 

This is publication No. 2 in the Gentry Invita- 
tion Series, based on data and annotations left by 
the late Dr. Alwyn H. Gentry, who died in an 
airplane crash in Ecuador on 3 August 1993. The 
series acknowledges the many contributions made 
by Dr. Gentry to our understanding of the Big- 
noniaceae. A summary of his life and a complete 
bibliography were published by James Miller 
(Ann. Missouri Bot. Card. 83 pt. 4). For infor- 
mation on the computerized databases left by 
Gentry, contact the Missouri Botanical Garden. It 
has been a priviledge to work on Costa Rica's 
Bignoniaceae using the many annotations and 
publications that Alwyn Gentry left us. The Big- 
noniaceae had been a very difficult family on 
which to work. Many genera and species were 
described from material that included only flowers 
or only fruits. The resulting number of monotypic 
genera and genera with disimilar species created 
considerable confusion. An additional problem 
was that tropical trees, and especially lianas, had 
been poorly sampled (Standley & Williams, 
1974). Al Gentry stepped into this situation and 



attacked the problem with intelligence and vigor, 
both in the field and in herbaria. He made order 
where there had been considerable chaos, and we 
are deeply indebted to him for this work. His ef- 
forts in understanding tropical forests and their 
diversity, together with his work on behalf of con- 
servation, inspired many of his students and col- 
leagues. Al Gentry's untimely death cut short 
what was already a rich and productive botanical 
career. 

Thanks to Gentry's work, most of the species 
of New World Bignoniaceae are well character- 
ized and their taxonomy has been put into good 
order. The percentage of Costa Rican bignon spe- 
cies that range from Mexico far into South Amer- 
ica greatly exceeds that for most plant families in 
our flora. There appear to be two reasons for this. 
The first is that we now have sound biological 
species concepts recognizing considerable varia- 
tion within each binomial, thanks to Al Gentry's 
extensive field work. The second and more fun- 
damental reason is related to the thin membra- 
nous-winged seeds, which have proved to be very 
successful dispersal agents in many of the genera. 
As Al Gentry pointed out (1983), genera and spe- 
cies lacking such seeds usually have much more 
limited geographical ranges (cf. Amphitecna and 
Parmentiera). Interestingly, although a few gen- 
era of Bignoniaceae can tolerate the cold winters 
of the northeastern United States, no indigenous 
member of the family is found above 2300 m el- 
evation in Costa Rica. This treatment has also 
benefited from comparison with a draft of the 
family for the Manual Flora of Costa Rica project 
by Quirico Jimenez and J. F. Morales. Our inde- 
pendent work on these plants resulted in almost 
identical treatments, thanks to the earlier annota- 
tions by Al Gentry. 

The Bignoniaceae appear to be a modern line- 
age in which recent diversification has produced 
a great number of species differing in a variety of 
minor morphological traits. Some of these minor 
variations have been the basis for erecting what 
appeared to be an excessive number of genera. 
Gentry (1973a, 1979) addressed this issue and 
made important advances in creating better ge- 
neric concepts. Although further "lumping" 
would seem to be desirable, current generic con- 
cepts have provided a useful information retrieval 
system that should not be radically altered until 
the evidence for doing so becomes substantial. 



78 



FIELDIANA: BOTANY 



Key to the Genera and Unusual Species of Bignoniaceae 

la. Trees and shrubs, stems woody and erect or spreading, tendrils absent [native and ornamental 

species] 2 

Ib. Lianas or vines, stems woody or herbaceous, stems climbing or clambering, tendrils often present 

16 

2a. Leaves pinnately compound, the central rachis with > 1 pair of lateral leaflets or bipinnately 

compound 3 

2b. Leaves simple, trifoliolate or palmately compound (with all leaflets arising from the apex of 

the petiole) 8 

3a. Leaves bipinnate (twice compound), the central rachis with opposing lateral 2 rachises on 
which the leaflets are borne; flowers lavender to bluish purple, staminode larger than the 

stamens and resembling a style [both native and planted ornamental species] 

Jacaranda 

3b. Leaves pinnate, the central rachis bearing the usually opposite leaflets; flowers not lavender 

or bluish, staminode much smaller than the stamens, not resembling a style 4 

4a. Corollas 3.5-6 cm long, yellow or orange to red-orange; native and introduced trees . . 5 
4b. Corollas 6--12 cm long, yellow, red or dark purple to maroon; introduced ornamental trees 

6 

5a. Stamens included; flowers usually yellow; leaflets 2-15 cm long; native small trees 

often planted for ornament Tecoma stans 

5b. Stamens exserted; flowers usually orange-red; leaflets 1-3.5 cm long; introduced or- 
namental shrubs with clambering branches Tecoma capensis 

6a. Corollas 8-12 cm long, red or red-orange, curved and opening upward, glabrous externally; 
calyx split down 1 side and spathe-like; commonly planted in lower elevation evergreen 

areas Spathodea campanulata 

6b. Corollas 6-9 cm long, yellow or dark purple to maroon, not usually opening upward, 
glabrous or puberulent externally; calyx not split down 1 side (not spathe-like); rarely 

planted in Central America 7 

7a. Corolla dark purple or maroon, glabrous externally; leaves 7-9rfoliolate; fruits pendulous, 

ellipsoid-oblong Kigelia pinnata 

7b. Corolla yellow, puberulent externally; leaves 5-7-foliolate; fruits not ellipsoid [plants not 

recorded from Costa Rica and not included in text] Haplophragma adenophyllum 

8a. (from 2b) Epiphytic shrubs; leaves simple, opposite, usually coriaceous; corollas radially sym- 
metric or only slightly 2-lipped (note that these plants have now been transferred to Schlege- 

liaceae) 9 

8b. Terrestrial trees (not epiphytic); corollas usually clearly bilaterally symmetric and 2-lipped [nei- 
ther red nor narrowly tubular] 10 

9a. Corollas usually red or purple, narrowly tubular and radially symmetric; fruit with fleshy 

covering; leaves usually < 10 cm long Gibsoniothamnus 

9b. Flowers white or pinkish, tubular-campanulate; fruit with hard outer covering; leaves to 18 

cm long Schlegelia 

lOa. Leaves opposite, usually 2-9-foliolate; corolla thin and without a transverse fold on the lower 
(abaxial) side, white to yellow, rose, or magenta; flowers usually borne in racemose panicles 

on distal stems; fruits dehiscent capsules with thin flat overlapping seeds 11 

lOb. Leaves fasciculate, alternate or opposite, simple or 3-foliolate; corolla thick and stiff with a 
transverse fold on the lower side, white to greenish white; flowers usually 1 to few and borne 
on thicker branches or trunk; fruits indehiscent with fibrous-fleshy or hard outer shells, seed 

angular and embedded in white fleshy pulp 14 

1 la. Flowers < 18 mm long; anthers pilose; fruits spirally twisted; leaflets 7-9 [often cuneate 

at base] Godmannia 

lib. Flowers > 20 mm long; anthers glabrous; fruits usually straight; leaflets (l-)2-7 . . 12 

12a. Corolla deeply split down the sides and strongly 2-lipped; rarely collected small trees of 

the Caribbean slope Tynanthus macranthus 

BURGER: FLORA COSTARICENSIS 79 



12h. Corolla not deeply split down the side, slightly 2-lipped; rare and common species of 

many habitats 13 

13a. Fruits usually held erect and 6-18 cm long; inflorescences with < 11 purple flowers; 
leaves simple and 3-foliolate on the same branches; small trees and shrubs of seasonally 

very dry Guanacaste Arrabidaea costaricensis 

13b. Fruits usually pendant and > 15 cm long; inflorescences usually with > 10 pink flowers 
or the corollas yellow or white; leaves rarely simple, 3-7-foliolate; small to large trees 

found in many habitats and often planted as ornamentals Tabebuia 

14a. (from lOb) Leaves opposite; 3-foliolate (often cruciform); calyx split on 1 side; fruits with 

fibrous exocarp (not a hard shell) Parmentiera 

14b. Leaves alternate or fasciculate; simple or 3-foliolate; calyx not split on 1 side; fruits with a 

hard exocarp 15 

15a. Leaves alternate; ovules on 2 parietal placentas or axile near base; seeds large (> 13 mm) 

Amphitecna 

15b. Leaves fasciculate; ovules on 4 parietal placentas; seeds small (< 8 mm) Crescentia 

16a. (from Ib) Leaves simple; tendrils absent; fruits spherical and indehiscent, seeds angular and wing- 
less; stems lacking 4-8 phloem areas in cross-section; with 1 axile placenta in each locule (now 

transferred to Schlegeliaceae) Schlegelia 

16b. Leaves usually compound (simple leaves sometimes present on young growth or the base of new 

shoots); tendrils often present; fruits usually elongate or flattened, dehiscing to release flat, usually 

winged seeds; stems with 4-8 phloem areas; with usually 2 axile placentas in each locule . . 17 

17a. Leaflets 3-15 mm long on young climbing stems, becoming 30 mm long on distal stems [4 leaflets/ 

node]; tendrils ending with small disc-like adhesive pads Mansoa parvifolia 

17b. Leaflets not so small on young climbing stems, becoming more than 3 cm long on distal stems; 

tendrils not ending in disc-like pads 18 

18a. Some leaves twice compound, some petioles bearing 2 or 3 petiolules with 3 or 5 leaflets each 

19 

1 8b. Leaves never twice compound, simple or with petioles bearing 2 or 3 leaflets [plants with woody 

stems; inflorescences never with sterile and fertile dimorphic flowers] 21 

19a. Herbaceous vines; inflorescence subspicate with 2 different kinds of flowers (distal flowers 
sterile); corolla greenish to purple; fruits ellipsoid and covered with prominent hooked spines 

Tourrettia lappacea 

19b. Woody vines and lianas; inflorescence racemose or paniculate, with 1 kind of flower; fruits 

linear, without spines 20 

20a. Corollas white or yellowish white, glabrous externally; tendrils tri-fid at the apex 

Pleonotoma variabilis 

20b. Corollas magenta or purple, puberulent externally; tendrils simple [plants not known from 

between Belize and Colombia and not included in the text] Arrabidaea inaequalis 

2 la. Each tendril terminating with usually 3 sharp stiff claw-like tips or hooks [corolla glabrous exter- 
nally] 22 

21b. Each tendril terminating with 1-3 slender tips, not hard and claw-like or hooked 24 

22a. Corolla deep purple or magenta; calyx 4- or 5-lobed; Golfo Dulce area in Central America 

[capsules usually linear] Parabignonia steyermarkii 

22b. Corolla yellow; calyx subtruncate to spathe-like; widely ranging in Central America ... 23 
23a. Capsules linear, 1.5-2 cm wide, > 15 cm long, valves not splitting in half (2/fruit); thin 

flexible Macfadyena 

23b. Capsules oblong, 2.5-4 cm wide, to 15 cm long, valves splitting longitudinally in half at 

maturity (4/fruit), thick woody Melloa quadrivalvis 

24a. Corolla orange, long-tubular and slightly curved (6-9 cm long, 3-12 mm diam.), lobes valvate in 

bud; planted for ornament in parks and gardens Pyroslegia venusta 

24b. Corolla not orange, without a long narrow slightly curved tube, lobes imbricate in bud; planted 

for ornament and/or native wild species 25 

25a. Branchlets 6-sided, hexagonal in cross-section, with 6 prominent longitudinal ribs along the young 
stems [fruits ellipsoid to oblong, 2.5-7 cm wide, surface smooth to tuberculate] 26 

80 FIELDIANA: BOTANY 



25b. Branchlets 2- or 4-sided to terete or subterete in cross-section, with 0-4 longitudinal ribs along 

the young stems 28 

26a. Calyx double, with an inner and outer whorl of distal lobes; leaves with dendroid (stellate 
or branched) hairs; tendrils usually with 3 distal tips; corolla strongly 2-lipped with the upper 

and lower lobes connate at anthesis Amphilophium 

26b. Calyx simple, with 1 distal whorl of lobes; leaves with simple hairs; tendrils with 3-15 distal 

divisions; corolla weakly 2-lipped, lobes free and reflexed at anthesis 27 

27a. Corolla white, tube strongly curved near the base; fruit oblong with a covering of short 

spines; widely ranging Pithecoctenium crucigerum 

27b. Corolla rose-purple, tube not curved; fruits without spines; not known from between central 

Nicaragua and Colombia and not included in the descriptions Distictis 

28a. Leaves with minute (0. 1 mm) pellucid flat glands on both surfaces; branchlets with a hollow center; 
capsules < 9 mm wide, linear and 24-45 cm long, surface minutely puberulent [corolla puberulent 

externally, white or greenish yellow with purple lobes) Stizophyllum 

28b. Leaf surfaces without pellucid-punctate glands; branchlets without a hollow center (sometimes 
hollow in Paragonia); capsule > 10 mm wide, linear to oblong or rounded, surfaces glabrous to 

puberulent, smooth to spiny or muricate 29 

29a. Leaves with branched or irregular dendroid hairs (at least in the vein axils beneath); capsules 
oblong (3-10 cm wide, ca. 5-10 mm thick), valves woody and flattened, surface smooth [seeds 

thin with broad lateral wings] 30 

29b. Leaves with slender simple or flat rounded-peltate hairs or glabrous; capsules without the above 

combination of characters 31 

30a. Interpetiolar glandular fields absent at nodes; calyx > 20 mm long; corolla yellow; seeds 6- 

12 cm wide, wings brown and opaque Callichlamys latifolia 

30b. Interpetiolar gland fields present at the nodes; calyx < 9 mm long; corolla lavender or rose; 

seeds 3-5 cm wide, wings translucent Xylophragma seemannianum 

3 la. Corolla 15-20 cm long with long narrow tube and distal rotate lobes (tubular-slaverform). white 
and puberulent externally; fruits oblong-cylindric, 9-22 cm long, 6-1 1 cm wide and 5-8 cm thick 

Tanaecium jaroba 

31b. Corolla < 15 cm long (mostly tubular-campanulate) and without the combination of characters 

listed above 32 

32a. Lower surface of leaf with gland fields in axils of basal veins or with gland field at apex of the 

petiole [corolla pink to magenta or white] 33 

32b. Lower surface of leaf lacking gland fields, apex of petiole lacking a gland field 35 

33a. Leaf axils with a conical structure made up of 3 imbricate series of scale-like pseudostipules 
or vegetative parts with the strong odor of onion or garlic [fruits with verrucose or smooth 

surface] Mansoa 

33b. Leaf axils lacking a conical structure of imbricate scale-like pseudostipules and vegetative 

parts lacking an onion-like odor 34 

34a. Native wild plants; young stems lacking conspicuous pseudostipules (rounded leaf-like pseu- 
dostipules present in C. diversifolia); corolla pink or lavender to purple, glabrous or puber- 
ulent externally; fruits oblong or linear Cydista 

34b. Vines planted for ornament (rarely collected in Costa Rica); young stems with broadly ovate 

leaf-like pseudostipules; corolla magenta to violet, glabrous externally; fruits linear 

Saritea magnified 

35a. Corolla strongly bilabiate, split down the sides beyond the middle, usually white; rarely collected 

Tynanthus 

35b. Corolla not strongly bilabiate, not split down the sides to the middle, colors various; common to 

rare species 36 

36a. Flowers yellow or white (sometimes with reddish markings in Mussatia); disc present beneath the 

ovary; valves of the fruits usually woody, > 25 mm wide, never cchinate 37 

36b. Flowers rose to magenta or lavender (white in species of Arrabidaea, Cydista, and Lundia); disc 
present or absent; valves of the fruits not woody unless echinate or tuberculate, usually < 25 mm 
wide -41 

BURGER: FLORA COSTARICENSIS 81 



37a. Corolla < 20 mm long, calyx < 2 mm long, widely campanulate; young stems strongly 
tetragonal in cross-section, with 4 prominent longitudinal ridges Mussatia 

37h. Corolla > 25 mm long, calyx > 4 mm long, cupular to tubular-campanulate; young stems 
terete or weakly tetragonal in cross-section, without 4 prominent ridges 38 

38a. Young stems with interpetiolar gland fields at the nodes; fruits tetragonal (square) in cross- 
section, ca. 2 cm thick, narrowly oblong [corolla glabrous externally] 

Ceratophytum tetragonolobum 

38b. Young stems lacking interpetiolar gland fields; fruits flattened and broadly oblong to elliptic 
or suborbicular 39 

39a. Corolla glabrous externally or with minute flat peltate hairs; capsules narrowed at the base 
and somewhat stipitate, flattened and elliptic to oblong or suborbicular . . . Anemopaegma 

39b. Corolla minutely puberulent externally; capsules not stipitate, flattened and elliptic to oblong 
40 

40a. Inflorescences 10-20 cm long; calyx entire; tendrils trifid at the apex 

Distictella magnoliifolia 

40b. Inflorescence 2-6 cm long; calyx usually split 1-2 mm along 2 sides; tendrils simple 

Adenocalymma inundatum 

4 la. (from 36b) Pseudostipules (cataphylls) forming small bromeliad-like clusters in distal leaf axils 

(especially below the inflorescence); fruits ellipsoid to suborbicular and echinate, 5-9 cm long 

with spines to 8 mm long [disc absent, venation pinnate] Clytostoma binatum 

41b. Pseudostipules not forming small bromeliad-like clusters in leaf axils; fruits mostly linear and 

smooth to verrucose 42 

42a. Calyx split down 1 side and spathe-like, apex thickened and curved; fruit surface with minute 

hairs that often give a grayish metallic luster [disc absent; fruits linear-oblong or ellipsoid; gland 

fields absent at the nodes] Phyrganocydia 

42b. Calyx not split down 1 side, not spathe-like or with curved thickened tip; fruit surface not grayish 

lustrous 43 

43a. Anthers pubescent; flower buds with narrow conical tip, the apex splitting off to produce an entire 

margin on the calyx cup at anthesis (also splitting as growth progresses) [corollas white to rose 

or magenta, minutely puberulent externally; disc absent; fruits 20-60 cm long, 1.5-2 cm wide 

margin and midvein prominent, surface densely puberulent] Lundia 

43b. Anthers glabrous; flower buds and calyx not calyptrate (sometimes coming off as a conical cap in 

Paragonia} 44 

44a. Tendrils ending in 3 short tips; corolla dark purple to deep maroon, glabrous or subglabrous 

externally; fruits (30-)50-110 cm long, linear, surfaces flat and glabrous [disc present] 

Martinella obovata 

44b. Tendrils simple or bifid at apex; corolla whitish or rose to purple, usually minutely puberulent 

externally; fruits < 60 cm long, surface with raised edges or raised midrib, glabrous or puberulent 

45 

45a. Calyx > 15 mm long, 2-lipped or lobed; fruits becoming black, with tuberculate or verrucose 

surface [disc present] Arrabidaea verrucosa 

45b. Calyx > 10 mm long, usually truncate at the apex; fruits not becoming dark with tuberculate or 

verrucose surface 46 

46a. Disc absent beneath the ovary; young stems terete or tetragonal, lacking interpetiolar gland fields; 

stems with 8-16 phloem areas in cross-section Cydista 

46b. Disc present beneath the ovary; young stems terete, often with interpetiolar gland fields; stems 

with 4 phloem areas in cross-section 47 

47a. Capsule valves flat and smooth (except in A. verrucosa), seeds usually with translucent wings 

differentiated from the brown center; tendrils simple at the apex; interpetiolar gland fields often 

present; leaflets lacking a faint sweet odor when crushed; deciduous to evergreen wet forests . . . 

Arrabidaea 

47b. Capsule valves convex and slightly rough-surfaced, seeds uniformly brownish; tendrils simple or 

minutely bifid at the apex; interpetiolar gland fields rarely present; leaflets with faint sweet odor 

when crushed; evergreen lowland rain forests Paragonia pyramidata 

82 FIELDIANA: BOTANY 



Amphitecna 
isthmica 




A. sessilifolia 

FIG. 10. Bignoniaccae: trees with simple leaves, fruit a calabash or pepo; species of Amphitecna. 



BURGER: FLORA COSTARICENSIS 



83 



Parmentiera macrophylla, 




FIG. 1 1. Bignoniaccac: trees with simple or compound leaves, fruit a calabash or pepo: species of Crescentia and 
Parmentiera. 



84 



FIELDIANA: BOTANY 



'A Tabebuia 

palustris 



Tynnanthus 
macranthus 



Godmania 
aesculi folia 




Arrabidea costaricensis 

FIG. 12. Bignoniaccac: trees and shrubs with 2- to 7-foliolatc leaves, fruits long narrow capsules. 



BURGER: FLORA COSTARICENSIS 



85 



Tabebuia 
chrysantha 




Tabebuia impetiginosa 

FIG. 13. Bignoniaceac: trees with palmately compound leaves and showy flowers; species of Tabebuia. 



86 



FIELDIANA: BOTANY 



Jacaranda 
i mimosifolia 



isK Jacaranda copaia 




FIG. 14. Bignoniaceae: trees with bipinnately compound leaves and lavender or bluish flowers; species of Jaca- 
randa. 



BURGER: FLORA COSTARICENSIS 



87 




FIG. 15. Bignoniaccac: ornamental trees and shrubs and an ornamental vine. 



88 



FIELDIANA: BOTANY 



Mansoa 
parvifolia "713 



Pleonotoma l^variabilis 




Tanaecium jaroba 



FIG. 16. Bignoniaccac: unusual vines with twice -compound leaves, with very small leaves, or with a very long 
narrow corolla tube. 



BURGER: FLORA COSTARICENSIS 



89 



Parabignonia 
steyermarkl i 




Melloa quadrivalvis 

FIG. 17. Bignoniaccac: vines climbing with tendrils ending in 3 sharp hard claws. 



90 



FIELDIANA: BOTANY 



&&OT Pi thecoctem um 
crucigerum 




FIG. 18. Bignoniaccac: vines with cchinatc or prominently tubcrculatc fruits. 



BURGER: FLORA COSTARICENSIS 



91 



Ceratophytum 
tetragonolobum 



Cydlsta potosina 



Callichlamys 
1 at i folia 




hyacinthina f 

FIG. 19. Bignoniaccae: vines with broad or thick elongate fruits. 



92 



FIELDIANA: BOTANY 



Anemopaegma 
chrysoleucum 




A. orbiculatum 

FIG. 20. Bignoniaccac: vines with ellipsoid or rounded fruits; species of Anemopae^ma. 



BURGER: FLORA COSTARICENSIS 



93 



Adenocalymma 
inundatum 



Xylophragma 
seemanniana 




Amphilophium 
paniculatum 



magnoTri fol i a 



FIG. 21. Bignoniaceae: vines with flattened ellipsoid or oblong fruits. 



94 



FIELDIANA: BOTANY 




Phryganocydia corymbosa 



Mansoa standleyi 30 
cm 



FIG. 22. Bignoniaccae: vines with flattened elongate fruits more than 2 cm wide. 



BURGER: FLORA COSTARICENSIS 



95 




Paragonla ^pyrami data <^ Stizophyllum riparium 

Fto. 23. Bignoniaceae: vines with very long linear flattened fruits. 



96 



FIELDIANA: BOTANY 



Cydista ^ttaequinoctialis 



Cydista diversi folia 



Cydista heterophy 1 1 a~*y 




lilacina 



FIG. 24. Bignoniaccae: vines with elongate narrow fruits; species of Cydista. 



BURGER: FLORA COSTARICENSIS 



97 



Arrabidaea candicans 




A. chica ^<^| /^^^^m \'^ K. conjugata 

FIG. 25. Bignoniaceac: vines with elongate flattened narrow fruits; species of Arrabidea. 



98 



FIELDIANA: BOTANY 



Arrabldaea patellifera Stizophyllum 

inaequilaterum 




Arrabidaea mollisslma 

FIG. 26. Bignoniaceae: vines with elongate narrow fruits and usually puberulcnt leaves 



Amphilophium 
pannosum 



BURGER: FLORA COSTARICENSIS 



99 



Adenocalymma Martius ex Meissner 

Lianas, climbing with tendrils, stem terete, 
with 4 phloem areas in cross-section, glandular 
fields absent at the nodes; pseudostipules small, 
subulate. Leaves opposite, 3-foliolate or 2-folio- 
late with terminal tendril, petiolate, leaflets with 
petiolules, margins entire, venation pinnate. Inflo- 
rescences terminal or axillary, narrow racemes, 
flower buds subtended by caducous bracts. Flow- 
ers with cupular calyx, 5-lobed and 2-lipped or 
truncated, usually with peltate glands near the 
rim; corolla tubular-funnelform to tubular-cam- 
panulate, yellow, usually puberulent on the exte- 
rior; stamens 4, filaments of 2 lengths, included, 
anthers glabrous, thecae straight and divaricate to 
slightly divergent, a staminode present; disc pul- 
vinate, ovary puberulent or with peltate hairs, 2- 
locular, ovules in 2 series on each placenta. Fruits 
oblong woody capsules, valves parallel to the sep- 
tum, rounded or slightly compressed, median vein 
not elevated; seeds almost wingless or with 2 lat- 
eral wings poorly differentiated from the central 
body, brownish or translucent distally. 

Adenocalymma is a Neotropical genus of about 
36 species ranging from Mexico to Argentina, 
with most of the species in eastern Brazil. One 
species is endemic to mangroves in eastern Mex- 
ico, two species are found in eastern Panama- 
Choco, and the following wide-ranging species 
occurs in Central America. 

Adenocalymma inundatum Mart, ex DC., Prodi. 
9: 201. 1845. Tabebuia calderonii Standl., J. 
Washington Acad. Sci. 14: 244. 1924. A. hin- 
tonii Sandw. Kew Bull. 1936: 10. 1936. A. 
calderonii (Standl.) Seibert, Carnegie Inst. 
Washington Publ. 522: 428. 1940. Figure 21. 

Lianas, stems 1-15 cm diam., tendrils to 20 cm 
long, leafy stems 1 .7-8 mm diam., essentially gla- 
brous, terete, drying dark with whitish lenticels 
but becoming grayish in age, interpetiolar line of- 
ten present; pseudostipules 2-3 mm long. Leaves 
with petioles 2-8 cm long, 0.8-2 mm diam., gla- 
brous or minutely puberulent at the apex, petio- 
lules of lateral leaflets 7-27 mm long, often thick- 
ened below the blade; leaflet blades 3.5-16 cm 
long, 2.5-7 cm wide, ovate to broadly elliptic- 
ovate or elliptic-oblong, apex acuminate, tips to 
1 5 mm long, base obtuse or slightly rounded and 
truncated, drying stiffly chartaceous and lustrous 
above, glabrous except for few minute (0.05-0.1 
mm) peltate hairs beneath, 2 veins 5-8/side, 3 



veins often elevated on the dried upper surface. 
Inflorescences 2-8 cm long, peduncles 6-22 mm 
long, minutely papillate-puberulent, bracts 6-12 
mm long, 4-7 mm wide, early caducous, pedicels 
2-7 mm long. Flowers with calyx 4-8 mm long, 
4-6 mm diam., minutely papillate-puberulent, 
lobes 1-2 mm or margin irregularly split; corolla 
3.5-6.5 cm long, funnelform-campanulate, yel- 
low, tube 5-18 mm diam., minutely puberulent 
with crooked hairs externally, lobes 10-20 mm 
long, narrowed below and rounded distally, be- 
coming reflexed; filaments 15-20 and 10-16 mm 
long, thecae 2-3 mm long. Fruits 9-18(-27) cm 
long, 2-3.5 cm wide, 13-22 mm thick, oblong, 
rounded at both ends, valves convex or flattened, 
pale grayish and rugulose-lenticellate; seeds 16- 
19(-21) mm long, 44-64(-76) mm wide, central 
area 17-20 mm wide, lateral wings membrana- 
ceous. 

Evergreen lianas in deciduous and partly decid- 
uous forests of the Pacific lowlands (in Costa 
Rica) and often found in riverine gallery forests, 
2-400 m elevation (to 900 m in Chiapas). Flow- 
ering primarily March-August in Central Ameri- 
ca, fruiting December-February. This species 
ranges from Mexico to Brazil and occurs on Gre- 
nada. 

Adenocalymma inundatum is recognized by its 
woody climbing habit and simple tendrils, oppo- 
site two- or three-foliolate leaves, short inflores- 
ences, papillate-puberulent yellow corollas, nar- 
rowly oblong woody fruit, and thick-bodied, 
winged seeds. The generally glabrous parts and 
large floral bracts that leave conspicuous scars are 
also distinctive characteristics. The floral bracts 
have the same texture as the calyx but are present 
for only a short time. Gentry used the name A. 
apurense (Kunth in H.B.K.) Sandw. for this ma- 
terial but later decided that A. apurense was re- 
stricted to South America. 



Amphilophium Kunth 

Lianas, climbing with trifid tendrils, stems te- 
rete, with 4 (5) phloem areas in cross-section, 
nodes with interpetiolar ridges, lacking gland 
fields; pseudostipules leaf-like, early deciduous. 
Leaves opposite, 3-foliolate or 2-foliolate and of- 
ten with a tendril, petiolate; leaflets with entire 
margins and pinnate venation. Inflorescences ter- 
minal, often on short lateral branches, racemose 
panicles with short lateral branches, bracts sub- 
tending the pedicels. Flowers with a cupulate- 



100 



FIELDIANA: BOTANY 



campanulate calyx with thick inner 2-3-lobed ca- 
lyx and- thinner 5-lobed spreading exterior; corol- 
la tubular and bilabiate, thick, whitish at first but 
becoming purple at anthesis, glabrous or puberu- 
lent externally, upper lip with 2 united lobes, low- 
er lip with 3 united lobes, lobes triangular; sta- 
mens 4, filaments of 2 lengths, anthers glabrous, 
thecae thick and divaricate, staminode present; 
disc annular; ovary 2-locular, ovules multiseriate 
in each locule. Fruits woody capsules, valves 
somewhat compressed, parallel to the septum, 
smooth to tuberculate; seeds thin, with 2 lateral 



membranous wings not clearly differentiated from 
the body of the seed. 

Amphilophium is a genus of about eight South 
American species, with two wider ranging species 
present in Costa Rica. Unusual features are a thick 
two-layered calyx and the stiff, closed corolla, 
split almost halfway into two lobes that must be 
forced open by pollinators. The rough-surfaced 
fruits and corollas that are white but turn purple 
at anthesis are also unusual. Distictis laxiflora 
(DC.) Greeenm. of Mexico and central Nicaragua 
also has six-angled stems, but that species has 
larger corollas and smooth elliptic fruits. 



Key to the Species of Amphilophium 

la. Fruit surface muriculate; pseudostipules 3-10 mm long and caducous; stems and petioles sparsely 

to densely puberulent with simple to dendroid hairs 0.05-0.4 mm long and usually drying grayish 

to yellowish gray; common plants in deciduous, partly deciduous, and evergreen forest formations 

A. paniculatum 

Ib. Fruit surface verrucose; pseudostipules 5-18 mm long and often persisting; stems and petioles 
villous with simple or branched hairs 1-3 mm long and drying yellowish to yellowish brown; 
uncommon plants of evergreen rain forest formations A. pannosum 



Amphilophium paniculatum (L.) Kunth in 
H.B.K., Nov. Gen. Sp. 3: 116. 1819. Bignonia 
paniculata L., Sp. PI. 623. 1753. A. molle 
(Schldl. & Cham.) Linnaea 5: 120. 1830. A. 
paniculatum var. molle (Schldl. & Cham.) 
Standl., Publ. Field Mus. Nat. Hist. Bot. Ser. 18: 
1114. 1938. Figure 21. 

Lianas to ca. 15m high, stems to 10 cm diam., 
tendrils to 17 cm long, leafy stems 1.8-5 mm 
diam., with 6 prominent longitudinal ridges (hex- 
agonal in cross-section), sparsely to densely pu- 
berulent with short (0.1-0.4 mm) scurfy-stellate 
and minute (0.05 mm) peltate hairs; pseudosti- 
pules 3-10 mm long, usually rounded, caducous. 
Leaves usually 2-foliolate, petioles 3-7 cm long, 
puberulent in longitudinal lines, petiolules 1-4.5 
cm long; leaflet blades 3-16 cm long, 2-11 cm 
wide, broadly ovate to ovate-oblong or ovate-el- 
liptic, apex acuminate to caudate-acuminate, base 
rounded and cordate to subcordate or truncate, of- 
ten minutely scurfy puberulent beneath, larger 
(0.2 mm) peltate rounded hairs beneath near the 
base, venation subpalmate, 2 veins 5-7/side, dis- 
tal veins strongly ascending. Inflorescences ter- 
minal, often on short lateral shoots and apparently 
axillary, 7-30 cm long, racemiform panicles with 
short lateral branches, 1 peduncle ca. 3 cm long. 



puberulent like the stems, bracts caducous, pedi- 
cels 1-9 mm long. Flowers with calyx 7-12 mm 
long, 5-7 mm diam., cupulate. minutely puberu- 
lent, with usually reflexed outer lobes 3-4 mm 
long, inner calyx 2- or 3-lobed; corolla 20-35 
mm long, 5-14 mm diam., white in early stages 
and becoming purple at anthesis, glabrous or with 
few hairs externally, lobes 11-20 mm long, nar- 
rowly triangular and not opening; filaments ca. 17 
and 14 mm long, thecae ca. 2 mm long; ovary 
puberulent, style ca. 24 mm long. Fruits 5-15 cm 
long, 2.8-5 cm wide, 13-30 mm thick, oblong or 
oblong-ellipsoid, surface muriculate and with mi- 
nute peltate hairs; seeds 11-15 mm long, 27-42 
mm wide, central area ca. 10 mm wide, brownish 
or transparent distally. 

Common and widespread plants of both sea- 
sonally deciduous forests and evergreen rain for- 
est formations, 5-1800 m elevation. Flowering in 
the wet season and early dry season (Gentry, 
1973b), but Costa Rican collections were made 
primarily in May-July and October-January. 
Probably fruiting throughout the year. This spe- 
cies ranges from Veracruz, Mexico, to northern 
Argentina. 

Amphilophium paniculatum is recognized by its 
unusual calyx with inner and outer layers, corollas 
that are thick and held shut at anthesis, and the 



BURGER: FLORA COSTARICENSIS 



101 



woody oblong fruits with muriculate surface. 
Stems with six prominent longitudinal ridges, 
dendritic hairs (when present), and ovate blades 
with palmate venation are useful vegetative char- 
acteristics. The thick corolla lobes are united and 
the two lips remain connivent until forced open 
by large bees. The flowers are full size and white 
or yellowish a day before anthesis; they then be- 
come purple. The thick two-layered calyx may be 
protection against nectar-robbing animals. A great 
range of variation makes separation of this species 
into two varieties based on pubescence impracti- 
cal. Because of its foliage and hexagonal stems, 
this species can be confused with Pithecoctenium 
echinatum, but that species has simple hairs and 
tendrils with as many as 15 distal segments. 

Amphilophium pannosum (DC.) Bur. & K. 
Schumann in Mart., Fl. Bras. 8 (2): 209. 1896. 
Bignonia pannosa DC, Prodr. 9: 148. 1845. A. 
oxylophium J. D. Smith, Bot. Gaz. 31: 119: 
1901. Figure 26. 

Lianas or slender vines to 3 cm diam., tendrils 
to 18 cm long, leafy stems 1.5-5 mm diam., with 
6 longitudinal ridges, densely villous with hairs 
0.5-3.7 mm long yellowish brown when dried; 
pseudostipules 5-18 mm long, equally broad, sub- 
sessile. Leaves usually 2-foliolate, petioles 2-7 
cm long, 1.3-2 mm diam., densely villous, peti- 
olules 1-4 cm long; leaflet blades 6-14 cm long, 
3.5-10 cm wide, ovate to ovate-elliptic, apex acu- 
minate, base rounded and truncate to cordate, up- 
per surface with simple hairs 0.5-2 mm long, low- 
er surface densely villous with branched hairs to 
2 mm long, venation subpalmate, 2 veins 4-6/ 
side. Inflorescences 5-18 cm long, 1 peduncles 
24 cm long, densely villous, 2 peduncles 5-12 
mm long, bracts 1-1.5 mm, linear lanceolate, ca- 
ducous, 2 bracteoles present below the calyx. 
Flowers with outer calyx 8-14 mm long, densely 
villous, lobes to 6 mm long, rounded, recurved or 
rotate, to 24 mm wide; corolla 22-30 mm long, 
tubular, 6-9 mm diam., at first yellowish white but 
turning purple, subglabrous externally, lip 1 1-14 
mm long; filaments ca. 15 and 12 mm long, the- 
cae ca. 2 mm long; style ca. 2 cm long. Fruits 5- 
10 cm long, 3-5.5 cm wide, elliptic-oblong, com- 
pressed, surfaces prominently verrucose with tu- 
bercles ca. 2 mm long, a central vein prominent 
or not, yellowish pubescent; seeds ca. 12 X 48 
mm. 

Uncommon plants of wet evergreen forest for- 
mations on both the Caribbean and southern Pa- 



cific slopes, 5-1400 m elevation (but note that 
nearly all our collections come from the southern 
Pacific slope). Flowering in January, May, and 
July-September. The species ranges from central 
Costa Rica to northern Argentina. 

Amphilophium pannosum is recognized by its 
climbing habit, six-angled stems, tendrils with 
usually trifid tips, densely villous yellow-brown 
pubescence, rounded pseudostipules, usually two- 
foliolate leaves, double-margined calyx, and ver- 
rucose fruit. Specimens of A. pilosum Standl. 
(1000-1400 m, in central Honduras) will key here 
because of similar pubescence and often retaining 
their pseudostipules, but that species may be an 
unusual variant of A. paniculatum. 



Amphitecna Miers 
Nomen conservandum 

REFERENCE A. Gentry, Amphitecna in Bigno- 
niaceae Part I (Crescentieae and Tourrettieae). 
Flora Neotropica Monogr. 25(1): 1-130. 1980. 

Small to medium-sized trees, stems without 
distinct phloem fields in cross-section, glabrous, 
becoming terete; pseudostipules and glandular 
fields absent at the nodes. Leaves alternate (sub- 
opposite), simple, sessile to short-petiolate, blades 
entire, subglabrous or glabrous (domatia absent), 
venation pinnate. Inflorescences terminal, axil- 
lary or on short-shoots borne on older branchlets 
or trunks (cauliflorus), flowers solitary or fascic- 
ulate, subtended by small bracts, pedicels well de- 
veloped and drying black, with or without brac- 
teoles, glabrous. Flowers drying black, calyx 
large, cupular or tubular, splitting along 1 or 2 
sides or opening irregularly, usually glabrous; co- 
rolla tubular-campanulate to funnelform, usually 
bent upward above a contracted region below the 
middle of the tube, greenish white to yellowish 
white, thick-textured, 5-(4-) lobed and slightly 2- 
lipped; stamens 4, slightly exserted, filaments 
equal or of 2 lengths, anthers glabrous, thecae 
thick and somewhat divergent; a staminode pres- 
ent; ovary 1- or 2-locular, ovules many on axile 
placentas (when 2-locular) or on 2 parietal pla- 
centas (where unilocular). Fruits pendulous, an 
indehiscent hard-surfaced and thin-walled pepo 
(calabash), globose to ellipsoid, usually subtended 
by a circular perianth scar, the outer shell easily 
broken; seeds more than 10 mm diam., embedded 
in a soft or fleshy white pulp, surfaces smooth and 
without wings. 

Amphitecna (formerly called Dendrosicus and 



102 



FffiLDIANA: BOTANY 



including Enallagma) is a genus of 20 species 
ranging -from Mexico, Central America, and the 
West Indies to northern South America. There are 
two centers of species diversity: southern Mexico 
to Belize and Guatemala, and Costa Rica to Pan- 
ama. The tree habit, lack of conspicuous hairs, 
simple large alternate leaves, large flowers with 
thick tissues that dry black, and pepo-like fruit 
with rounded seeds are unusual traits among Big- 
noniaceae and are characteristic of the tribe Cres- 
centieae. The perianth usually forms a circum- 
scissle abscission scar around the base of the fruit. 
The corolla often has an abaxial transverse inden- 
tation just distal to the constricted section of the 
corolla tube, where the distal portion of the wid- 
ening corolla tube bends upward. The large size, 



strong odor (usually unpleasant), large amount of 
nectar, and thick texture of the flowers are char- 
acteristic of flowers pollinated by bats. The flow- 
ers often vary significantly in size within a single 
species; leaf sizes and shapes also vary greatly 
within species. More important, herbarium mate- 
rial suggests that there may be populations inter- 
mediate between generally recognized species in 
this genus. Whether this variation is due to hy- 
bridization, overly narrow species concepts, or in- 
herent variability is difficult to determine at this 
time because many species are represented by 
only a few collections. Gentry (1990) interpreted 
the greater number of localized endemic species 
in this genus as being a result of mammal-dis- 
persed seeds, in contrast to genera with wind-dis- 
persed seeds. 



Key to the Species of Amphitecna 

la. Plants of ocean shores and low-elevation swamp forests; fruits subglobose and rounded distally; 
leaf blades often rounded distally, coriaceous A. latifolia 

Ib. Plants of well-drained soils, 5-2200 m elevation; fruits usually oblong to ellipsoid with a narrowed 
apex (sometimes globose in A. gentryi); leaf blades usually acute to acuminate at the apex, char- 
taceous to subcoriaceous 2 

2a. Flowers borne on older leafless stems and trunks; leaf blades to 50 cm long [calyx 20-34 mm long, 
corolla 38-50 mm long; 10-600 m elevation] A. kennedyi 

2b. Flowers terminal on leafy shoots or on short-shoots near the leaves; leaf blades to 30 cm long . . . 
3 

3a. Flowers usually in clusters or 2-8, pedicels often with linear bracteoles 1-4 mm long 2-5 mm 
above the pedicel base; leaves often drying dark; ( 1 200-) 1 500-2200 m elevation [calyx 22-32 mm 
long, corolla 33-53 mm long; fruits 13-17 cm long] A. sessilifolia 

3b. Flowers usually 1 or 2, pedicels lacking linear bracteoles near the base; leaves often drying brown 
or grayish; 5-1400 m elevation 4 

4a. Calyx 22-28 mm long, corolla 44-60 mm long; fruits 10-14 cm long; 5-800 m elevation. Pacific 
slope A. isthmica 

4b. Calyx 15-20 mm long, corolla 25-40 mm long; fruits 5-8 cm long; 600-1400 m elevation, Carib- 
bean slope A. gentryi 



Amphitecna gentryi W. Burger, sp. nov. Figure 
10. 

Arbor parva. Folia alternata, glabra, anguste 
obovata-elliptica vel elliptica-oblonga, acuminata. 
Inflorescencia terminalia, floribus 1-2, pedicelis 
glabris. Calyx 15-20 mm longus, bipartitus, lobis 
rotundatis, glabris. Corolla 22-28 mm longa, 
campanulata, alba. Fructus globosus vel ellipso- 
ideus, seminibus in pulpa inclusis. TYPUS: Costa 
Rica, Puntarenas, Monteverde, in lower commu- 
nity, alt. 1350 m. March 1981. Haber 478 (holo- 
typus MO 3162341, isotypus CR). 

Small trees 5-10 m tall, trunks to 18 cm diam.. 



leafy stems 2-6 mm diam., terete, glabrous, yel- 
lowish or pale gray, expanded below the leaf base. 
Leaves alternate, petioles 0-8 mm long, 0.7-1.8 
mm diam., glabrous, often with lustrous hard yel- 
lowish tissue at the base; leaf blades 6-16.5 cm 
long, 2-6.5 cm wide, elliptic-obovate, obovate or 
narrowly elliptic-oblong, apex acute to caudate- 
acuminate, gradually narrowed to the cuneate or 
acute base, drying chartaceous and grayish or 
grayish brown, glabrous above and below, 2 
veins 7-12/side. Inflorescences terminal, of 1 
flower (rarely 2) on leafy shoots or subterminal 
on short lateral shoots, subtended by hard yellow- 
ish subulate bracts 1-2 mm long, pedicels 30-42 



BURGER: FLORA COSTARICENSIS 



103 



mm long, glabrous, sometimes with very small (1 
mm) yellowish bracteoles 1-5 mm above the 
base. Flowers with calyx 15-20 mm long, ca. 10 
mm diam., glabrous, splitting near the base into 2 
broad lobes, rounded distally; corolla 25-40 mm 
long, white or cream-green, tube 6-1 1 mm diam., 
papillate-puberulent in the distal half, lobes ca. 3 
X 10 mm; filaments 20-23 mm long, thecae ca. 
5 mm long; ovary not examined. Fruits 5-8 cm 
long, 4-5 cm diam., oblong to subglobose, with 
a slightly narrowed apex, surface smooth, green 
drying brown; seeds 11-14 mm long, 14-17 mm 
wide, 6-9 mm thick. 

Plants of lower montane rain forest formations 
of the Caribbean slope, 600-1400 m elevation. 
Flowering in February-March and July; fruiting 
in March, June-July, and September. As presently 
known, this species ranges from Volcan Cacao 
eastward to Braulio Carillo National Park in 
northern and central Costa Rica. 

Amphitecna gentry! is recognized by its small 
tree habit, smaller subsessile leaves turning gray- 
ish when dried, usually solitary terminal flower, 
smaller corollas, and globose or oblong fruit. 
Specimens placed here include Gomez et al. 
21603, Hammel & Chavarria 17533 A, and the 
following collections by William Haber and as- 
sociates: 478, 5841, 6673, 7473, 8235, and 9272. 

Gentry recognized these plants as distinct and 
designated these plants as Amphitecna sp. aff. A. 
donnell-smithii (Sprague) L. O. Williams, a low- 
land species of Belize and Guatemala with oblan- 
ceolate leaves of similar length. This species also 
resembles two highland species of northern Cen- 
tral America described by L. O. Williams: A. mol- 
inae, of Honduras, and A. silvicola, of the border 
between Guatemala and Mexico. More important, 
it should be noted that there are collections that 
appear to be intermediate between this species 
and A. isthmica at elevations of 300-600 m (Her- 
rera 1966, 21 18), implying that the plants placed 
here could also be considered a subspecific high- 
land element of A. isthmica. Nevertheless, the 
characteristics used in the key seem to separate 
the two taxa effectively, and only further collect- 
ing can determine whether the intermediates are 
unusual individuals or are parts of a cline. 

Amphitecna isthmica (A. Gentry) A. Gentry, 
Taxon 25: 108. 1976. Dendrosicos isthmicus A. 
Gentry, Phytologia 26: 442. 1973. Figure 10. 

Small trees 5-10 m tall, trunks to 15 cm diam., 
leafy stems 1.7-5 mm diam., glabrous, pale yel- 



low or grayish, terete. Leaves subsessile or with 
petioles to 10 mm long, 1-2.5 mm diam., gla- 
brous, often with hard lustrous yellowish tissue at 
the base; leaf blades 6-22(-32) cm long, 2-8(-10) 
cm wide, narrowly elliptic-obovate to narrowly el- 
liptic-oblong, apex acuminate, gradually narrowed 
to the cuneate base, drying stiffly chartaceous and 
grayish, glabrous, 2 veins 8-17/side. Inflores- 
cences terminal (axillary), usually 1 (2) flower on 
leafy stems, subtended by few hard subulate yel- 
lowish bracts 1-2 mm long, pedicel 30-48 mm 
long, 0.8-1.7 mm diam., glabrous, without brac- 
teoles. Flowers drying black or dark brown, calyx 
22-30 mm long, 11-18 mm wide, splitting to near 
the base into 2 broad obtuse lobes, glabrous; co- 
rolla 44-60 mm long, 30-40 mm wide at the 
lobes, greenish white, papillate puberulent in the 
distal half externally, tube 5-14 mm diam., lobes 
3-8 mm long, with reflexed rim; filaments 20-30 
mm long, thecae 4-5 mm long; ovary bilocular to 
above the middle. Fruits 9-14 cm long, 3-8 cm 
diam., ellipsoid-cylindric, narrowed at the ends, 
surfaces smooth and punctate, perianth scar thick- 
ened and disc-like at base; seeds ca. 10-14 mm 
long, 4 mm thick. 

Plants of wet evergreen rain forest formations 
on the Pacific slopes, 5-800 m elevation. Flow- 
ering material has been collected in February, 
May, August, and October-December. The spe- 
cies ranges from northern Costa Rica to Colom- 
bia. 

Amphitecna isthmica is recognized by its low- 
land rain forest habitat, small tree habit, narrow 
alternate leaves that usually dry grayish, usually 
solitary terminal flowers on long pedicels, calyx 
splitting into two rounded lobes, and ellipsoid-cy- 
lindrica fruits. This species lacks the bracteoles on 
the pedicels seen in A. gentryi and A. sessilifolia 
and is separated from A. kennedyi by having only 
terminal flowers. This species appears to be re- 
stricted to the evergreen Pacific slope in Costa 
Rica, which helps distinguish it from the closely 
related A. gentryi. 

Amphitecna kennedyi (A. Gentry) A. Gentry, 
Taxon 25: 108. 1973. Demdrosicos kennedyi A. 
Gentry, Phytologia 26: 441. 1973. Figure 10. 

Small trees and shrubs 3-15 m tall, leafy stems 
1.5-1 1 mm diam., glabrous, ridged below the leaf 
base but becoming terete and pale grayish. Leaves 
with petioles 2-12 mm long, 1-4 mm diam., gla- 
brous, often with lustrous hard tissue developed 
at the base; leaf blades 14-35(-50) cm long, 5- 



104 



FIELDIANA: BOTANY 



13(-21) cm wide, elliptic-obovate to narrowly ob- 
ovate or oblanceolate, apex usually short-acumi- 
nate, gradually narrowed to the cuneate base, dry- 
ing chartaceous and brownish or grayish, glabrous 
above and below, 2 veins 8-17(-22)/side, often 
weakly loop-connected near the margin. Inflores- 
cences cauliflorous, 1-3 flowers on short-shoots 
borne below leafy nodes on branchlets or on the 
surface of larger stems, subtended by stiff grayish 
subulate bracts 2-3 mm long, pedicels 25-45 mm 
long, 0.6-1.7 mm diam., glabrous. Flowers dry- 
ing black, calyx 20-34 mm long, 9-20 mm diam., 
split to near the base to form 2 lobes, glabrous; 
corolla 38-56 mm long, tubular-campanulate, 1 3- 
19 mm wide at the mouth, greenish white to yel- 
lowish green, with peltate or papillate hairs on the 
distal half externally, lobes forming a reflexed bi- 
labiate rim; filaments 24-28 mm long, thecae 4- 
5 mm long; ovary lepidote, 1-locular with 2 pa- 
rietal placentae. Fruits 11-16 cm long, 5-7 cm 
diam., ellipsoid, apiculate at base and apex; seeds 
13-15 mm long, embedded in pulp. 

Plants of evergreen rain forest formations of 
both Caribbean and southern Pacific slopes (in 
Costa Rica), 10-600 m elevation. Flowering col- 
lections were made in January-April and July- 
September. This species ranges from the east coast 
of Honduras to central Panama. 

Amphitecna kennedyi is distinguished by its 
small tree habit, flowers borne at leafless nodes or 
on the older stems, corolla with poorly developed 
lobes forming a reflexed rim, ellipsoid fruits, and 
lowland rain forest habitat. The lower half of the 
corolla often has a smoother texture demarcated 
distally by a transverse ridge. 

Amphitecna latifolia (Miller) A. Gentry, Taxon 
25: 108. 1976. Crescentia latifolia Miller, Card. 
Diet. ed. 8: 306. 1768. C. obovata Benth., Bot. 
Voy. Sulphur 130. 1844. Enallagma latifolia 
(Miller) Small, Fl. Miami 171. 1913. -A. obovata 
(Benth.) L. O. Williams, Fieldiana, Bot. 36: 25. 
1973. Dendrosicos latifolius (Miller) A. Gentry, 
Taxon 22: 644. 1973. Figure 10. 

Small trees (2-)4-8(-12) m tall, trunks to 20 
cm diam., branches often crooked, leafy stems 
2.5-12 cm diam., terete, glabrous, often becoming 
pale gray, lenticellate. Leaves alternate or subop- 
posite, petioles 2-14 mm long but poorly differ- 
entiated from the leaf, glabrous; leaf blades 10- 
28 cm long, 4-14 cm wide, narrowly or broadly 
obovate to obovate-oblong or oblong, apex short- 
acuminate to rounded, base obtuse to cuneate and 



slightly decurrent on the petiole, drying subcori- 
aceous to coriaceous, glabrous or with minute 
(0.05 mm) peltate hairs beneath, 2 veins 7-13/ 
side, weakly loop-connected near the margin. In- 
florescences terminal or below leafless nodes, of 
1-3 flowers, subtended by stiff subulate bracts 2- 
3 mm long, pedicels (22-)40-63 mm long, gla- 
brous. Flowers glabrous externally, drying black, 
calyx 28-38 mm long, tube 7-15 mm diam., split- 
ting along the lateral sides nearly to the base to 
form 2 divergent lobes; corolla 45-65 mm long, 
tube 1 2-24 mm diam., 3-4 cm wide at the apex, 
tubular-campanulate, greenish white or pale yel- 
lowish green, lobes 5-15 mm long; filaments ca. 
25 mm long, anthers 5-6 mm long; style 5-6 cm 
long. Fruits 7-10 cm long, 6-10 cm wide, glo- 
bose or subglobose, green, surface smooth and 
hard, with thickened disc-like perianth scar at 
base; seeds 13-16 mm long, 14-17 mm wide, 5- 
8 mm thick, surface smooth. 

Small trees growing just inland from coastal 
mangroves and in low-elevation swamps in ev- 
ergreen and deciduous areas of both the Pacific 
and Caribbean coasts, 0-20(-200) m elevation. 
Flowering and fruiting throughout the year (more 
abundantly in the wet season). This species ranges 
from southern Florida, the West Indies, Mexico, 
and Central America to Venezuela and Ecuador. 

Amphitecna latifolia is recognized by the tree 
habit, stiff simple alternate leaves, glabrous flow- 
ers drying black and arising directly from distal 
stems, globose indehiscent fruit, and rounded 
seeds embedded in pulp. The flowers have a dis- 
agreeable odor. Common names are calabasillo de 
playa, jicarita, jicaro de playa and swamp cala- 
bash (Belize). 

Amphitecna sessilifolia (J. D. Smith) L. O. Wil- 
liams, Fieldiana, Bot. 36: 25. April 1973. Ta- 
bebuia sessilifolia J. D. Smith, Bot. Gaz. 25: 
156. 1898. Neotuerckheimia gonoclada J. D. 
Smith, Bot. Gaz. 47: 259. 1909. Enallagma ses- 
silifolia (J. D. Smith) Standl., Publ. Field Mus. 
Nat. Hist., Bot. Ser. 18: 1120. 1938. A. sessili- 
folius (J. D. Smith) A. Gentry, Taxon 22: 646. 
Nov. 1973. Figure 10. 

Trees or shrubs to 12 m tall, trunks 7-25 cm 
diam., leafy stems 4-1 1 mm diam., glabrous, with 
ridges beneath the leaf base, pale gray, becoming 
terete. Leaves alternate, petioles 2-10 mm long, 
poorly differentiated from the blade, glabrous; 
leaf blades 10-35 cm long, 2-11 cm wide, nar- 
rowly obovate to oblanceolate or narrowly oblan- 



BURGER: FLORA COSTARICENSIS 



105 



ceolate, apex acute to acuminate, gradually nar- 
rowed to the cuneate base, drying chartaceous, 
subglabrous with minute (0.05 mm) peltate hairs, 
2 veins (8-)l 1-22/side, often arising at 70-90 
from the midvein. Inflorescences terminal fasci- 
cles of 2-8 flowers, basal bracts to 7 mm long, 
pedicels 22-65 mm long, 1-2 mm diam., gla- 
brous, drying black, often with 2 small (1-4 mm) 
linear bracteoles 2-5 mm above the base. Flowers 
with calyx inflated before anthesis, 22-32 mm 
long, 11-19 mm diam., glabrous, splitting later- 
ally into 2 lobes, margins rounded; corolla 33-53 
mm long, 25-35 mm wide distally, white with 
greenish tinge, tube 6-13 mm diam., bent upward 
near the middle, lobes 3-12 mm long, glabrous 
proximally but with minute (0. 1 mm) peltate hairs 
on the lobes externally; filaments ca. 26 and 24 
mm long, thecae ca. 5 mm long. Fruits 12-17 cm 
long, 6-9 cm diam., subglobose to ellipsoid-ob- 
long with narrowed apex to 1 cm long, surface 
smooth, glabrous, drying dark; seeds 15-20 mm 
long, 15-24 mm wide. 

Understory trees within evergreen montane rain 
forests, 1200-2200 m elevation. Flowering 
throughout the year. This species ranges from the 
Cordillera de Tilaran to the Chiriqui highlands of 
western Panama. 

Amphitecna sessilifolia is recognized by its tree 
habit, alternate oblanceolate leaves, large 2-lobed 
calyx, pale green corollas, smooth oblong-ellip- 
soid fruits, and restriction to montane forests. An- 
other important distinction is that the thick calyx 
splits apart, often resulting in a whitish tissue ex- 
posed along the edge. A number of specimens 
from Monteverde have more clearly differentiated 
petioles and corolla lobes lacking peltate hairs ex- 
ternally; these have been annotated A. haberi (an 
unpublished name) by Gentry. Because they are 
otherwise identical to A. sessilifolia and live in 
the same kind of habitat, it seems best to consider 
them no more than a local variant of A. sessili- 



folia. Common names are calabacero, calabash, 
quacalillo, jicarilla, jicaro. 



Anemopaegma Martius ex Meisner 
Nomen conservandum 

Lianas or vines (in Central America), climbing 
with simple or distally trifid tendrils, stem with 8 
(rarely 4) phloem areas in cross-section, terete; 
nodes usually with interpetiolar lines, gland fields 
absent; pseudostipules leaf-like to small or absent. 
Leaves opposite, 2-5-foliolate, petiolate, often 
bearing a tendril coiled near the tip, blades often 
drying yellowish or grayish green, margins entire, 
venation pinnate. Inflorescences axillary or ter- 
minal, few-flowered racemes or 1 or 2 flowers, 
bracts small, pedicels short. Flowers with cupular 
or campanulate calyx, truncated and subentire or 
slightly lobed, often with glands below the margin 
externally; corolla tubular-campanulate or funnel- 
form, 5-lobed and somewhat 2-lipped, bright yel- 
low to pale yellow or white, externally glabrous 
or with minute glandular peltate hairs; stamens 4, 
included, filaments of 2 lengths, anthers glabrous, 
thecae straight and divaricate, staminode present; 
disc pulviniform; ovary usually stipitate, ellipsoid, 
puberulent or with peltate hairs, 2-locular with 
ovules in 2-6 series in each locule. Fruits woody 
or coriaceous capsules, elliptic to orbicular, valves 
strongly flattened or rounded, smooth; seeds flat, 
completely surrounded by a membranaceous sub- 
orbicular wing or wingless and with larger brown 
corky seed area and thinner distal area. 

Anemopaegma is a genus of about 46 species 
ranging from Mexico to Brazil and Argentina; 
most of the species are South American. The great 
variation found within some species has resulted 
in a large number of names, and it is often diffi- 
cult to demarcate species. Many of the proposed 
species probably will have to be brought together 
into broader species concepts. 



Key to the Species of Anemopaegma 

la. Leaves usually with 4 or 5 leaflets A. orbiculatum 

Ib. Leaves with no more than 2 or 3 leaflets 2 

2a. Pseudostipules conspicuous (4-9 mm long); tendrils lacking trifid tips; fruits ellipsoid, seeds brown 

and lacking a translucent outer wing A. chrysoleucum 

2b. Pseudostipules absent or minute; tendrils often trifid near the tip; fruits flattened and oblong, seeds 

pale yellowish with thin translucent circumferential wing (where known) 3 

3a. Leaves usually puberulent beneath, leaflet margins not involute [stems terete, lacking corky ridges; 

corolla tube subglabrous externally] A. puberulum 



106 



FIELDIANA: BOTANY 



3b. Leaves subglabrous beneath, leaflet margins often involute 4 

4a. Corolla tube with minute peltate hairs externally; stems tetragonal and often with corky ridges; 

seeds not known A. santaritensis 

4b. Corolla tube glabrous; stems subterete and longitudinally striate; seeds suborbicular 

. . A. chrysanthum 



Anemopaegma chrysanthum Dugand, Caldasia 
4: 307. 1947. 

Lianas or slender vines, stems to 2 cm diam., 
tendrils to 13 cm long, trifid at tip, leafy stems 
1.2-8 mm diam., glabrous, longitudinally striate, 
grayish and becoming lenticellate; pseudostipules 
minute. Leaves foliolate, petioles 8-50 mm long, 
1-2.5 mm thick, glabrous, petiolules 8-40 mm 
long; leaflet blades 5-17(-20) cm long, 3-9(-12) 
cm wide, broadly ovate to elliptic, apex acute or 
short-acuminate, base acute to obtuse, drying stiff- 
ly chartaceous and grayish green, subglabrous, 2 
veins 3-7/side. Inflorescences short axillary ra- 
cemes with 2-8 flowers, peduncles ca. 10 mm 
long, pedicels 10-15 mm long. Flowers subgla- 
brous, calyx 6-9 mm long, 4-6 mm diam. with 
distal glands, margin subentire; corolla 50-70 
mm long, yellow, mouth 12-22 mm wide, lobes 
10-15 mm long; filaments 24-32 and 16-23 mm 
long, thecae 4-5 mm long. Fruits 8-20 cm long, 
5-8 cm wide, 1 cm thick, oblong-elliptic, flat- 
tened, surface smooth, yellow-brown, stipe 1-2 
cm long; seeds 40-55 mm diam., suborbicular. 

Plants of evergreen forest formations of the 
northern cordilleras in Costa Rica; 350-1000 m 
elevation. Flowering in March-April. This species 
ranges from Mexico to Ecuador. 

Anemopaegma chrysanthum is recognized by 
its three-tipped tendrils, two-foliolate opposite 
leaves, few-flowered axillary racemes, glabrous 
yellow corollas, and broad flat fruits with subor- 
bicular seeds. Rarely collected in Central Ameri- 
ca, this species has only recently been found in 
Costa Rica. 

Anemopaegma chrysoleucum (Kunth in H.B.K.) 
Sandw., Lilloa 3: 459. 1938. Bignonia chryso- 
leucum Kunth in H.B.K., Nov. Gen. Sp. 3: 134. 
1819. A. puncticulatum Pittier & Standl., J. 
Wash. Acad. Sci. 15: 461. 1925. A. macrocarpa 
Standl., Publ. Field Columb. Mus., Bot. Sen 4: 
262. 1929. Figure 20. 

Lianas or slender vines, stems to 2 cm diam., 
tendrils to 17 cm long, simple, leafy stems 1.2-5 
mm diam., minutely puberulent at the nodes, lon- 



gitudinally striate, interpetiolar lines usually well 
developed; pseudostipules 4-9 mm long, broadly 
ovate. Leaves 2-(3-) foliolate, petioles 8-34 mm 
long, 0.8-1.5 mm diam., glabrous or minutely 
(0.1-0.2 mm) puberulent, petiolules 10-18 mm 
long; leaflet blades 5-13(-18) cm long, 2-6(-8) 
cm wide, narrowly ovate-elliptic to elliptic, apex 
acute or acuminate, base acute to obtuse, drying 
chartaceous and yellowish green or gray-green, mi- 
nutely punctate, glabrous except for minute hairs 
on the midvein above, 2 veins 5-8/side. Inflores- 
cences of 1 or 2 axillary flowers (1-4/node) or a 
raceme to 9 cm long with ca. 5 flowers, peduncles 
3-15 mm long, 1 mm diam., minutely puberulent, 
pedicels 6-14 mm long. Flowers with calyx 8-1 1 
mm long, 7-9 mm diam., rounded glands and mi- 
nute peltate hairs or subglabrous, margin entire; co- 
rolla 58-105 mm long, white to yellowish white, 
glabrous externally, narrowed base of tube ca. 2 
cm long, mouth 1 2-20 mm wide, lobes 1 2-25 mm 
long, 17-24 mm wide; filaments 24-34 and 16-25 
mm long, thecae 4-5 mm long. Fruits 7-13 cm 
long, 3-5 cm wide, 2 cm thick, ellipsoid, surface 
smooth and lustrous, yellow-brown, stipe ca. 1 cm 
long; seeds ca. 15 X 22 mm, brown throughout, 
margin thin but not translucent. 

Plants of river edges and swampy areas in low- 
land evergreen rain forest formations on both the 
Caribbean and Pacific slopes, 0-50 m elevation. 
Flowering throughout the year. This species rang- 
es from Mexico to Venezuela and Peru. 

Anemopaegma chrysoleucum is recognized by 
its slender climbing stems with simple tendrils, 
usually bifoliolate opposite leaves, small leaf-like 
pseudostipules, few-flowered axillary inflores- 
cences, large white to yellowish corollas, and el- 
lipsoid fruits with flat seeds that lack thin trans- 
lucent membranous wings. This species differs 
from its congeners by having seeds with stiff 
cardboard-like texture that are dispersed by water. 

Anemopaegma orbiculatum (Jacq.) DC., Prodr. 
9: 190. 1845. Bignonia orbiculata Jacq., Sel. 
Stirp. Am. Hist. 184, tab. 180, fig. 79. 1763. 
Pithecoctenium panamense Benth., Bot. voy. 
Sulphur 129. 1844. Figure 20. 

Slender lianas to 5 m high, stems to 25 mm 
diam., longitudinally striate, hollow, tendrils 8-17 



BURGER. FLORA COSTARICENSIS 



107 



cm long, simple or with trifid arms to 5 mm long, 
leafy stems 1.5-9 mm diam., minutely (0.1-0.2 
mm) puberulent with thin hairs, an interpetiolar 
line present or absent; pseudostipules rarely pres- 
ent. Leaves 4- or 5-foliolate, petioles 3-14 cm 
long, 1.2-2.2 mm diam., minutely puberulent, 
longitudinally striate, tendril scar ca. 2 mm wide, 
petiolules 4-40(-55) mm long; leaflet blades 5- 
15(-22) cm long, 3-8(-9.5) cm wide, ovate-ellip- 
tic to ovate, apex acuminate, base obtuse to 
rounded and subcordate, drying chartaceous and 
greenish, punctate above, minutely and sparsely 
puberulent beneath, 2 veins 5-7/side. Inflores- 
cences axillary. 2-10 cm long, racemes with 1- 
1 flowers (rarely with a cymose lateral branch at 
the first node), peduncles 3-30 mm long, ca. 1.2 
mm diam., minutely puberulent, pedicels 6-12 
mm long. Flowers with cupular-campanulate ca- 
lyx 6-10 mm long, 5-9 mm wide, margin sub- 
entire or with 5 irregular minute teeth, minutely 
(0.1 mm) puberulent; corolla 33-70 mm long, 
tube 14-20 mm wide at the mouth, yellow and 
often with the lobes paler, glandular puberulent 
externally (more densely near the base), lobes 8- 
18 mm long, the lower reflexed; filaments ca. 16 
and 23 mm long, thecae ca. 4 mm long, divaricate 
at 180. Fruits 9-14 cm long, 6-10 cm wide, ca. 
2 cm thick at center, suborbicular to broadly el- 
liptic, with a basal stipe 8-15 mm long, valves 
slightly convex or flattened, surface smooth and 
lustrous, yellowish brown; seeds 30-38 mm long, 
40-47 mm wide, elliptic-orbicular, body ca. 14 
mm wide, wings transparent near the periphery. 

Plants of forest edges in evergreen lowland rain 
forest formations of the Caribbean slope and Ni- 
coya peninsula, 5-900 m. Flowering in April- 
September. Nearly all Costa Rican collections 
come from near La Selva and the Rio Sarapiqui- 
Rio Frio drainage areas; we have no collections 
of this species from southern Costa Rica. This 
species ranges from El Salvador to Colombia and 
Venezuela. 

Anemopaegma orbiculatum is recognized by its 
climbing stems with simple or trifid tendrils, op- 
posite four- or five-foliolate leaves, subentire ca- 
lyx cups, bright yellow corollas, and suborbicular 
fruits with orbicular winged seeds. This is the 
only bignoniaceous vine in Central America with 
consistently five-foliolate or four-foliolate leaves. 

Anemopaegma puberulum (Seibert) Miranda, 
Anal. Inst. Biol. Mexico 24: 93. 1953. Chodan- 
thus puberulum Seibert, Carnegie Inst. Wash- 
ington Publ. 527: 425. 1940. Figure 20. 



Slender lianas to 2 cm diam., tendrils 1 1-27 
cm long and trifid at the tip, leafy stems 2-6 mm 
diam., glabrous or puberulent with thin straight 
hairs 0.1-0.3 mm long, longitudinaly striate when 
dry, interpetiolar ridge usually present; pseudo- 
stipules minute. Leaves 2-foliolate with tendril or 
tendril scar, petioles 18-40 mm long, 1.5-2.3 mm 
diam., glabrous or minutely puberulent, petiolules 
10-45 mm long, 1-2.2 mm diam.; leaflet blades 
8-21 cm long, 4-12 cm wide, ovate to ovate-el- 
liptic, apex acuminate, base obtuse to truncate or 
subcordate, drying stiffly chartaceous, usually gla- 
brous above, punctate, minor venation raised 
above (dried), lower surface glabrous to pubes- 
cent with straight thin hairs to 0.1-0.4 mm long, 
2 veins 4-7/side, 3 veins prominent on both sur- 
faces. Inflorescences axillary, 2-8-flowered ra- 
cemes to 6 cm long, peduncles 14-20 mm long, 
ca. 1.5 mm diam., glabrous or puberulent, pedi- 
cels ca. 7 mm long. Flowers with cupular calyx 
6-9 mm long, 5-8 mm diam., distal margin sub- 
entire or slightly lobed, puberulent, glabrous or 
with few minute peltate hairs; corolla 50-71 mm 
long, tubular campanulate, yellow, glabrous or pa- 
pillate-puberulent externally, tube 12-21 mm 
diam. at mouth, lobes 10-15 mm long, 10-14 mm 
wide, rounded, filaments ca. 14 and 20 mm long, 
thecae 3-4 mm long. Fruits 10-21 cm long (in- 
cluding the 11-20 mm stipe), 6-8 cm wide, ca. 
13 mm thick, oblong-elliptic, valves flattened, 
woody, with a median longitudinal line, surface 
lustrous, yellowish brown; seeds 35-60 mm 
diam., suborbicular with thin translucent periph- 
eral wing, body 14-18 mm long. 

Uncommon plants of evergreen or partly decid- 
uous forest formations on the Caribean and south- 
ern Pacific slopes in Costa Rica, 5-800(-1200) m 
elevation. Probably flowering throughout the year 
(mostly in the dry season, January-May). This 
species ranges from Veracruz, Mexico, to Ecua- 
dor. 

Anemopaegma puberulum is recognized by its 
vining habit with trifid tendrils, opposite two-fo- 
liolate leaves, few-flowered axillary inflorescenc- 
es, subentire calyx cups, large yellow subglabrous 
corollas, flattened oblong-elliptic woody fruit, and 
seeds with thin suborbicular peripheral wings. The 
glabrous lustrous upper leaf surface with elevated 
minor venation, lack of conspicuous pseudostip- 
ules, and fruits with a longitudinal line down the 
middle also help distinguish this uncommon spe- 
cies from its congeners. Specimens with densely 
puberulent stems and leaves do look quite differ- 
ent from those that are essentially glabrous, but 
they seem to differ in no other way, and there are 



108 



FffiLDIANA: BOTANY 



intermediate collections. For this reason, A. chry- 
santhum may prove to be a synonym of A. pub- 
erulum. 

Anemopaegma santaritense A. Gentry, Ann. 
Missouri Bot. Card. 58: 93. 1971. Figure 20. 

Slender lianas climbing with trifid (simple) ten- 
drils 3-12 cm long, leafy stems 1.5-4 mm diam., 
glabrous or minutely papillate-puberulent, longi- 
tudinally ridged or with longitudinal corky ridges, 
tetragonal, interpetiolar ridge present or absent; 
pseudostipules absent. Leaves 2-foliolate, with a 
tendril or tendril scar, petioles 15-34 mm long, 
0.9-1.3 mm diam., glabrous or minutely papillate- 
puberulent along the upper (adaxial) surface, pet- 
iolules 6-17 mm long; leaflet blades 5-11 cm 
long, 2.5-6.5 cm wide, ovate to narrowly ovate- 
elliptic or ovate-orbicular, apex acuminate to cau- 
date-acuminate, base obtuse to somewhat round- 
ed, drying chartaceous and greenish, glabrous or 
with minute papillate hairs along the midvein on 
both surfaces, 2 veins 3-5/side. Inflorescences 
axillary, flowers 1 or 2 or 2-8 on a raceme to 9 
cm long, subglabrous, bracts 0.5-1 mm long, ped- 
icels 5-14 mm long. Flowers with cupular calyx 
6-10 mm long, 5-8 mm diam., glabrous or with 
few minute papillate hairs, margin entire; corolla 
52-75 mm long, tubular-campanulate, 17-20 mm 
diam. at mouth, pale yellow or white, glabrous or 
with minute peltate hairs externally, lobes 8-12 
mm long; filaments ca. 20 and 18 mm long, the- 
cae 3-4 mm long; ovary stipitate, ca. 2 mm long. 
Fruits not known. 

Rarely collected plants of evergreen rain forest 
formations on the Caribbean slope and Golfo Dul- 
ce, 0-1000 m elevation. Flowering collections 
have been made in March-May and September in 
Costa Rica and Panama. This species ranges from 
eastern Costa Rica to Colombia. 

Anemopaegma santaritensis is recognized by 
its climbing habit and usually trifid tendrils, te- 
tragonal stems, opposite two-foliolate leaves, few- 
flowered axillary inflorescences, calyx cups with 
subentire margins, and larger yellow or white co- 
rollas with minute peltate hairs. The upper leaf 
surfaces are dull green and often reflexed along 
the margins when dried. 



Arrabidaea DeCandolle 

Lianas climbing with simple tendrils (rarely 
small trees or shrubs), stems terete, with 4 phloem 
areas in cross-section, interpetiolar ridges and 



gland fields often present at the nodes; pseudo- 
stipules usually small and inconspicuous. Leaves 
opposite, petiolate, 3-foliolate or 2-foliolate with 
tendril or tendril scar (rarely simple), leaflets pe- 
tiolulate, margins entire, puberulent to subgla- 
brous, venation pinnate or palmate, domatia rarely 
present. Inflorescences often terminal pyramidal 
thyrses or panicles, also axillary or from leafless 
nodes, usually many-flowered, bracts minute, ped- 
icels usually puberulent. Flowers small to large, 
calyx cupulate to tubular or campanulate (rarely 
patelliform), distally truncated and entire or mi- 
nutely 5-denticulate, puberulent; corolla tubular- 
campanulate or funnelform, usually somewhat 2- 
lipped, pale pink to purple (white), usually dense- 
ly puberulent externally, lobes 5, unequal, round- 
ed at the apex; stamens 4, included, filaments of 
2 lengths, anthers glabrous, thecae usually divar- 
icate, staminode present; disc annular; ovary nar- 
rowly cylindrical, with peltate hairs, 2-locular, 
ovules 2-seriate. Fruits long-linear capsules, sep- 
ticidally dehiscent, valves flattened parallel to the 
septum, coriaceous and usually with a raised lon- 
gitudinal median vein, smooth (rarely tubercu- 
late); seeds much wider than long with 2 mem- 
branaceous lateral wings (rarely corky), body of 
the seed usually differentiated from the thin 
wings. 

Arrabidaea is a genus of about 70 species rang- 
ing from Mexico and the West Indies to Argen- 
tina; the majority of species are South American. 
Arrabidaea inaequalis (DC. ex Spreng.) K. 
Schum. is disjunct between Belize and Colombia 
but should be searched for in our flora; it is easily 
recognized by its twice-compound leaves. Two 
other species are included in this treatment that 
have not been collected in Costa Rica: A. florida 
and A. pubescens (q.v.). Our species of Arrabi- 
daea are mostly lianas with simple tendrils and 
two- or three-foliolate leaves, usually having pa- 
niculate inflorescences, narrowly cupular calyx 
with entire or minutely dentate margin, purple to 
lilac or rose corollas that are puberulent external- 
ly, long narrow fruits with flattened valves, and 
flat seeds, often with thin lateral wings transparent 
at the tips. A number of our species have very 
short flowering periods, which may account for 
their poor representation in herbaria. There is 
great variation within many of the species, which 
can make determination of specimens lacking 
flowers very difficult. These plants are easily con- 
fused with species of Cydista (lacking a disc at 
the base of the ovary) and Lundia (pubescent an- 
thers). 



BURGER: FLORA COSTARICENSIS 



109 



Key to the Species of Arrabidaea 

la. Shrubs, small trees, or rarely lianas, tendrils usually absent; gland fields absent; simple leaves often 
present beneath 3-foliolate leaves on distal stems; inflorescences short racemes with < 10 flowers; 

fruits often held erect [plants of dry deciduous forests, flowering in May-June] 

A. costaricensis 

Ib. Lianas, tendrils usually present, gland field present or absent; simple leaves only found on young 
plants (except in A. panel lifera); inflorescences usually paniculate with > 10 flowers; fruits pen- 
dulous 2 

2a. Calyx 14-23 mm long, tubular; fruits with prominently tuberculate valves, becoming black or dark 
brown; seeds 16-18 mm long [leaves 2-foliolate; flowering in the wet season in evergreen forest 
formations] A. verrucosa 

2b. Calyx 1-9 mm long, cupulate, short-tubular or flattened and saucer-like; fruits with smooth or 
slightly muricate valves becoming yellowish brown to dark brown, seeds 6-14 mm long 3 

3a. Calyx 1-4 mm long, flattened and saucer-like (patelliform), entire and lacking teeth; corolla tube 
glabrous externally but with the lobes puberulent and whitish [deciduous forests, flowering in July- 
October] A. patellifera 

3b. Calyx 3-9 mm long, cupulate to short-tubular, entire or with minute teeth; corolla tube puberulent 
above the narrowed base, lobes pink to purple 4 

4a. Corolla 8-18 mm long [flowers in dense distal clusters on large inflorescences; leaves usually 
subglabrous and drying grayish; interpetiolar gland fields usually absent; not known to occur be- 
tween Nicaragua and central Panama] A. florida 

4b. Corolla 1 8-55 mm long 5 

5a. Leaves usually drying reddish brown above and below, usually subglabrous and lustrous with minor 
venation raised above and below; plants of wet evergreen forests 5-900 m elevation [flowering 
March-August] A. chica 

5b. Leaves not drying reddish brown and lustrous above and below, usually conspicuously puberulent 
on the veins or on the lower surface; plants of deciduous and partly deciduous forest formations 
(also evergreen forests in A. candicans), 0-400 m elevation 6 

6a. Lower leaf surfaces pale grayish with a dense tomentum of minute hairs; flowering in July-Decem- 
ber [fruits with the midvein and lateral edges usually prominent; central area of seed clearly de- 
marked from the translucent wings] 7 

6b. Lower leaf surface not pale grayish with a dense tomentum, hairs 0.1-1 mm long; flowering in 

January-August 8 

7a. Flowering in November-December; seeds 24-34 mm wide; collected in Costa Rica in partly 

deciduous and evergreen forests of the Pacific slope A. candicans 

7b. Flowering in July-September; seeds 17-24 mm wide; not known from Costa Rica 

A. pubescens 

8a. Surface of fruit puberulent and soft to the touch; calyx 4-6 mm long [margin often with minute 
teeth and whitish]; flowering in December-early March A. mollissima 

8b. Surface of fruit glabrous or minutely puberulent, not soft to the touch; calyx 4-9 mm long; flowering 
in April-August 9 

9a. Calyx cup with small (0.3 mm) distal teeth; stems and leaves with simple and/or branched hairs; 

fruits with the midvein not elevated; mostly flowering in April-June (in Costa Rica) 

A. corallina 

9b. Calyx cup with entire distal margin (or sometimes split); stems and leaves with simple hairs; fruits 
with the midvein usually elevated; mostly flowering in June-August A. conjugate 

Arrabidaea candicans (L. C. Rich.) DC., Prodr. Lianas to 7 cm diam., climbing with simple 

9: 185. 1845. Bignonia candicans L. C. Rich., tendrils 11-21 cm long, leafy stems 1-6 mm 

Act. Soc. Hist. Nat. Paris 1: 110. 1792. A. pa- diam., minutely puberulent, becoming grayish, 

chycalyx Sprague, Bull. Herb. Boiss., ser. 2 6: glandular fields conspicuous, an interpetiolar line 

373. 1906. A. rhodothyrsus Kra'nzl., Fedde Re- usually present. Leaves with petioles 45-97 mm 

pert. 17: 20. 1921. Figure 25. long, 1-2.3 mm diam., minutely puberulent, pet- 

110 FIELDIANA: BOTANY 



iolules 11-55 mm long, 0.6-1.4 mm diam.; leaflet 
blades 4.5-20 cm long, 3-12 cm wide, ovate to 
broadly ovate or ovate-elliptic, apex acute to acu- 
minate, base obtuse to subtruncate or slightly 
rounded, drying chartaceous, densely and minute- 
ly grayish puberulent beneath, 2 veins 4-7/side, 
strongly ascending. Inflorescences terminal or ax- 
illary to distal leaves, 20-40 cm long, to 30 cm 
wide, a pyramidal thyrse, peduncles 2-4 mm 
diam., densely minutely grayish puberulent, lat- 
eral branches 2-7 cm long, pedicels 1-3 mm long. 
Flowers with calyx 3-5 mm long, 2-4 mm diam., 
tubular-conical, densely minutely puberulent, 
margin entire; corolla 20-38 mm long, 6-9 mm 
diam. distally, purple or rose (white within), 
densely puberulent externally, lobes 5-1 1 mm 
long, 7-10 mm wide, rounded; filaments ca. 16 
and 13 mm long, thecae 2-2.5 mm long. Fruits 
12-35 cm long, 7-12 mm wide, linear to linear- 
oblong, valves smooth, midvein and margins 
slightly elevated; seeds 7-9 mm long, 24-34 mm 
wide, central dark area 14-18 mm wide, clearly 
differentiated from the translucent wings. 

Plants of evergreen rain forest and partly de- 
ciduous forest formations, 5-900 m elevation. 
Flowering November-December, fruiting Febru- 
ary-June. Uncommon in Costa Rica, it is known 
only from the Pacific slope, from Tacares south- 
ward to Golfo Dulce. This species ranges dis- 
junctly from Mexico to Brazil. 

Arrabidaea candicans is recognized by the 
climbing habit, simple tendrils, two- or three-fo- 
liolate leaves, large inflorescences, puberulent ca- 
lyx with entire margin, puberulent rose to purple 
corollas, long linear fruits, and transparent-winged 
seeds. The leaflets with grayish lower surface and 
the gland fields conspicuous below the nodes are 
useful vegetative distinctions. 

Arrabidaea chica (Humboldt & Bonpland) Ver- 
lot, Rev. Hortic. 40: 154. 1868. Bignonia chica 
Humboldt & Bonpland, PI. Aequin. 1: 107, pi. 
31. 1808. Figure 25. 

Lianas to 35 m high, stems to 15 cm diam., 
tendrils to 12 cm long, leafy stems 1.5-5 mm 
diam., glabrous or sparsely and minutely (0.1 
mm) puberulent, longitudinally striate, nodal 
glandular fields visible on young stems. Leaves 
with petioles 32-85 mm long, 0.8-1.9 mm diam., 
usually glabrous, petiolules (4-)10-32(-46) mm 
long; leaflet blades (3-)5-16 cm long, 2-8 cm 
wide, narrowly ovate-elliptic to elliptic-oblong or 
elliptic, acuminate at the apex, base obtuse or 



rounded, drying chartaceous and usually reddish 
brown, usually glabrous above and below (in Cen- 
tral America), 2 veins 4-7/side, minor venation 
raised on both surfaces (dried). Inflorescences 
terminal, often 3-parted at a branchlet apex, 8-36 
cm long, peduncles 4-8 cm long, ca. 1.7 mm 
diam., very sparsely puberulent, drying dark, 
proximal lateral branches to 7 cm long, pedicels 
4-7 mm long. Flowers with calyx 3-5 mm long, 
2-3 mm diam., cupulate, minutely papillate pu- 
berulent, grayish green, distally subentire with 
small (0.2 mm) teeth; corolla 18-34 mm -long, 
tube 6-1 1 mm diam., lavender to purple (white 
within), densely papillate puberulent externally, 
lobes 6-12 mm long, to 12 mm wide; filaments 
ca. 12 and 9 mm long, thecae 1.5-2 mm long. 
Fruits 14-28 cm long, 9-14 mm wide, linear- 
oblong, valves flat, smooth and light to dark 
brown, midvein and margins slightly raised; seeds 
8-10 mm long, 23-38 mm wide, wings strongly 
differentiated and transparent. 

Uncommon plants of wet evergreen rain forest 
formations on both Caribbean and Pacific coasts 
(rarely collected in moist sites in seasonally very 
dry deciduous formations), 5-900 m elevation. 
Flowering in late March-August. This species 
ranges from Mexico to Argentina. 

Arrabidaea chica is recognized by its climbing 
habit with simple tendrils, two- or three-foliolate 
leaves, nearly entire cupular calyx, pale purple co- 
rollas densely puberulent externally, and narrow 
glabrous fruits with winged seeds. The leaves are 
usually glabrous and usually become reddish 
brown and lustrous when dried. These plants have 
been widely used as a source of red dyes; such a 
use has been reported around Santa Ana (Eche- 
verria 32 & 37 F). Gentry (1973b) remarked that 
puberulent forms of this species intergraded with 
A. candicans, but in Central America it may be 
better to assign all the puberulent forms to A. can- 
dicans. 

Arrabidaea conjugata (Veil.) Mart., Flora 24 (2), 
Beibl. 46. 1841. Bignonia conjugata Veil., Fl. 
Flum. 245. 1825; 6: tab. 18. 1827. Figure 25. 

Lianas climbing to the tops of medium-size 
trees, to 5 cm diam., leafy stems 1 .5-5 mm diam., 
sparsely very minutely puberulent, glabrescent, 
gland fields often conspicuous; pseudostipules ca. 
2 mm long, conical. Leaves with petioles 2-13 
cm long, 0.8-2 mm diam., sparsely minutely pu- 
berulent, longitudinally striate, petiolules 8-55 
mm long; leaflet blades 5-17(-21) cm long, 3.5- 



BURGER: FLORA COSTARICENSIS 



111 



10(-13) cm wide, ovate-elliptic to ovate or ellip- 
tic-obovate, apex acuminate, base cuneate to 
rounded, drying chartaceous to subcoriaceous, 
glabrous or with minute (0. 1 mm) hairs along the 
veins beneath and usually with longer (0.2-0.5 
mm) hairs in vein axils (domatia), 2 veins 4-77 
side, strongly ascending (subpalmate). Inflores- 
cences terminal, 14-34 cm long, pyramidal pan- 
icles with 3-5 pairs of lateral branches, peduncles 
to 10 cm long, 2.3 mm diam., pedicels 1-4 mm 
long, densely puberulent. Flowers with tubular 
calyx 4-7 mm long, 3-5 mm diam., grayish with 
a dense minute (0. 1 mm) puberulence, margin en- 
tire or 5-denticulate, submarginal glands present; 
corolla 30-42 mm long, 7-12 mm wide at the 
mouth, tubular-campanulate, magenta (white 
within), puberulent with short crooked hairs, lobes 
4-11 mm long, 7-9 mm wide; filaments 12-16 
and 8-1 1 mm long, thecae ca. 2 mm long. Fruits 
12-31 cm long, 8-14 mm wide, linear-oblong, 
midvein often slightly elevated, surface slightly 
muricate; seeds 7-11 mm long, 19-27 mm wide, 
brown throughout except for transparent distal 2- 
3 mm. 

Evergreen lianas of the seasonally dry decidu- 
ous forests of northwestern Costa Rica (and in 
partly deciduous forests elsewhere), 5150 m el- 
evation. Flowering in June-September; fruiting in 
the dry season (Gentry, 1973b). The short flow- 
ering season may account for a paucity of collec- 
tions. This species ranges from northwestern Cos- 
ta Rica to Brazil. 

Arrabidaea conjugata is recognized by its 
climbing habit with simple tendrils, two- or three- 
foliolate glabrate leaves, terminal pyramidal pan- 
icles, densely puberulent grayish calyx with sub- 
marginal glands, puberulent magenta corolla, and 
long linear flattened fruit with two-winged seeds. 
The conspicuous gland fields below the nodes, 
leaves with longer hairs in vein axils beneath 
(when present), and raised minor venation also 
help to distinguish this species. 

Arrabidaea corallina (Jacq.) Sandw., Kew Bull. 
1953: 460. 1954. Bignonia corallina Jacq., 
Fragm. Bot. 37, tab. 42, fig. 1. 1800-1809. B. 
glabrata Kunth in H.B.K., Nov. Gen. Sp. 3: 
137. 1819. B. obliqua Kunth in H.B.K., Nov. 
Gen. Sp. 3: 135. 1819. Figure 25. 

Lianas to 20 m high, stems to 12 cm diam., 
tendrils to 15 cm long, leafy stems 2-4 mm diam., 
sparsely to densely puberulent with simple or 
branched hairs 0.2-0.5 mm long, gland fields pre- 



sent or absent at the nodes. Leaves mostly trifo- 
liolate, petioles 2-9 cm long, 0.7-2 mm diam., 
usually densely puberulent with simple or 
branched hairs, petiolules 3-38 mm long; leaflet 
blades 5-18 cm long, 3-9(-12) cm wide, ovate- 
elliptic to broadly rounded-elliptic or elliptic-ob- 
ovate, apex short-acuminate, base narrowed to 
rounded-truncate, drying thin chartaceous, green 
or brown, minutely puberulent on the veins above, 
sparsely to densely puberulent beneath with sim- 
ple or branched hairs 0.2-0.8 mm long, 2 veins 
4-7/side. Inflorescences usually short-branched 
(raceme-like) panicles from the axils of fallen 
leaves or terminal, 8-30 cm long, peduncles 2-24 
mm long, 1.5-2 mm diam., densely puberulent, 
bracts 1-4 mm long, linear, pedicels 1-12 mm 
long. Flowers with cupulate calyx 5-9 mm long, 
4-7 mm diam., sparsely puberulent with multi- 
cellular hairs 0.1-0.2 mm long, margin subentire 
or with narrow apiculate teeth 0.3-1 mm long 
(sometimes split and appearing lobed); corolla 
27-46 mm long, 8-12 mm wide at the mouth, 
magenta to lavender, puberulent externally, lobes 
5-14 mm long, to 12 mm wide; filaments ca. 15 
and 10 mm long, thecae 2.5-3 mm long. Fruits 
12-47 cm long, 11-21 mm wide, valves flat and 
smooth (minutely pitted), midvein apparent or 
not; seeds 1 1-14 mm long, 32-50 mm wide, cen- 
tral area ca. 14 mm wide, thin transparent edges 
ca. 5 mm wide. 

Rarely collected deciduous lianas of seasonally 
dry deciduous forest formations, 1-300 m eleva- 
tion. Flowering in April-June (July-November in 
Panama; Gentry, 1973b). The species ranges from 
Mexico to Argentina. 

Arrabidaea corallina is recognized by its 
climbing habit, simple tendrils, two- or three-fo- 
liolate leaves with usually dense pubescence of 
branched or simple hairs, puberulent magenta or 
lavender corollas, long narrow fruits, and winged 
seeds. The calyx tube is usually sparsely puberu- 
lent distally and of thin texture with venation dry- 
ing darker, and narrow teeth (0.3 mm) are usually 
present along the irregular distal margin. 

Arrabidaea costaricensis (Kranzl) A. Gentry, 
Brittonia 25: 231. 1973. Saldanhaea costaricen- 
sis Kranzl, Fedde Repert. 17: 124. 1921. Arra- 
bidaea erecta Miranda, Anal. Inst. Biol. Mex. 
24: 91. 1953. Figure 12. 

Shrubs or small trees to 5 m tall (rarely woody 
vines), tendrils absent or simple, leafy stems 2-5 
mm diam., densely puberulent with straight or 



112 



FIELDIANA: BOTANY 



curved hairs 0.1-0.3 mm long, soon glabrescent 
and pale grayish, gland fields absent, interpetiolar 
lines present or absent. Leaves simple to 3-fo- 
liolate, petioles 3-55 mm long, 0.8-1.7 mm 
diam., densely puberulent, petiolules 5-27 mm 
long with central petiolule much longer than lat- 
erals; leaflet blades 3-12 cm long, 2.5-7.5 cm 
wide, ovate-elliptic to broadly ovate, apex acute 
or short-acuminate, base narrowed and obtuse to 
rounded, drying chartaceous and grayish green or 
brown, smooth or slightly scabrous above. With 
straight hairs ca. 0.2 mm long, lower surface 
densely puberulent, the soft straight hairs to 0.3 
mm long, venation subpalmate, 2 veins 4-6/side, 
strongly ascending. Inflorescences terminal or ax- 
illary at leafless nodes, 3-7 cm long, racemes with 
fewer than 10 flowers, peduncle 5-14 mm long, 
densely puberulent, bracts caducous or to 4 mm 
long, pedicels 2-5 mm long. Flowers with calyx 
3-5 mm long, 3-4 mm diam., cupular, surface 
grayish with dense minute puberulence, marginal 
teeth 5, 0.3-0.7 mm long, 0.3-0.5 mm wide at 
base; corolla 24-32 mm long, 5-6 mm wide at 
the mouth, reddish purple (white), densely puber- 
ulent with crooked multicellular hairs externally. 
Fruits 8-21 cm long, 6-14 mm wide, linear-ob- 
long valves flattened, with prominent midrib, 
brown, smooth or with pits or dots; seeds 6-12 
mm long, 18-26 mm wide, central area 13-16 
mm wide. 

Rarely collected shrubs or vines of seasonally 
very dry deciduous forest formations in northern 
Guanacaste, 10-300 m elevation (to 900 m in 
Honduras). Flowering in May-June; fruiting in 
June-January. This species ranges from Mexico 
along the Pacific slope to northwestern Costa 
Rica. 

Arrabidaea costaricensis is distinguished by its 
often shrubby habit and usual lack of tendrils 
(sometimes present), grayish indumentum of 
straight simple hairs, presence of both simple and 
three-foliolate leaves, few-flowered inflorescenc- 
es, and linear-oblong fruits with two-winged 
seeds. The capsules are sometimes held upright, 
and they may open explosively. The simple leaves 
are usually borne near the base of a shoot or at 
the initiation of a new shoot, with the following 
leaves three-foliolate. 

Arrabidaea florida DC., Prodr. 9: 184. 1845. A. 
panamensis Sprague, Bull. Herb. Boiss. se. 2, 
6: 371. 1906. 



Lianas, tendrils to 8 cm long (often lacking on 
specimens), leafy stems 2-5 mm diam., minutely 
(0.05 mm) puberulent and lenticellate, longitudi- 
nally striate, interpetiolar lines usually present, 
gland fields absent. Leaves 2- or 3-foliolate, pet- 
ioles 10-60 mm long, 0.8-1.5 mm diam., minute- 
ly puberulent, petiolules 5-24 mm long; leaflet 
blades 5-14 cm long, 2.5-8 cm wide, ovate-ellip- 
tic to ovate-oblong, apex acute to acuminate, base 
acute to obtuse or rounded, drying stiffly charta- 
ceous and grayish green or brown, lustrous and 
glabrous above, minutely puberulent beneath, 2 
veins 4-7/side. Inflorescences terminal or axil- 
lary to distal leaves, 9-35 cm long, open panicles 
to 30 cm wide, 2 peduncles 1 .5-7 cm long, distal 
branches grayish puberulent, flowers crowded in 
distal clusters, subtended by narrow bracts 1-4 
mm long, pedicels 0-3 mm long. Flower buds 
narrowly ellipsoid, calyx 3-4 mm long, 2-3 mm 
diam., minutely puberulent with grayish hairs, 
margin entire or with minute teeth 0.1-0.3 mm 
long; corolla 12-18 mm long, campanulate with 
spreading lobes, pink or lavender (white), throat 
often white, 5-7 mm wide at the mouth, lobes 3- 
6 mm long; filaments ca. 8 and 6 mm long, thecae 
1.5 mm long. Fruits 10-26 cm long, 8-11 mm 
wide, valves flat, brown with raised midrib, 
smooth; seeds 6-9 mm long, 17-28(-36) mm 
wide, dark central area 11-14 mm wide, wings 
translucent. 

Rarely collected plants (in Costa Rica) of moist 
forest formations, 1-1 100 m elevation. Flowering 
in July-October. The species ranges disjunctly 
from Mexico to Bolivia and Brazil. 

Arrabidaea florida is distinguished by its 
climbing habit with simple tendrils, generally gla- 
brous two- or three-foliolate leaves, large inflo- 
rescences with many distal clusters of small flow- 
ers, pink to lavender puberulent corollas, long 
narrow fruit, and two-winged seeds. Sterile plants 
of this species resemble Adenocalymma inunda- 
tum (cartilaginous margins on leaflets) and Tyn- 
nanthus croatianus A. Gentry of Panama (tendrils 
with trifid tips). 

Arrabidaea mollissima (Kunth in H.B.K.) Bu- 
reau & K. Shum. in Mart., Fl. Bras. 8 (2): 46. 
1896. Bignonia mollissima Kunth in H.B.K., 
Nov. Gen. Sp. 3: 133. 1819. B. littoralis Kunth 
in H.B.K., Nov. Gen. Sp. 3: 139. 1819. A. mol- 
licoma Blake, Cont. Gray Herb. 52: 92. 1917. 
A. isthmica Standl., J. Wash. Acad. Sci. 18: 337. 
1925, pro parte. Figure 26. 



BURGER: FLORA COSTARICENSIS 



113 



Lianas (rarely shrub-like), stems to 3 cm diam., 
tendrils to 12 cm long, leafy stems 2-5 mm diam., 
minutely puberulent with thin straight hairs, gla- 
brescent, terete, glandular fields rarely present; 
pseudostipules 2-3 mm long, conical. Leaves de- 
ciduous in dry season, 2- or 3-foliolate, petioles 
25-80 mm long, 1.1-2.8 mm diam., densely pu- 
berulent with straight simple or gland-tipped hairs 
0.3-0.8 mm long, petiolules 6-35 mm long; leaf- 
let blades 4-13 cm long, 2-10 cm wide, broadly 
ovate to ovate-orbicular, apex caudate-acuminate 
or short-acuminate, base rounded and truncate to 
subcordate, drying grayish and chartaceous, upper 
surface pubescent with straight hairs 0.5-0.8 mm 
long, lower surface densely puberulent with soft 
hairs 0.3-0.7 mm long, venation subpalmate, 2 
veins 4-6/side. Inflorescences terminal or axil- 
lary to leafless nodes, 3-25 cm long, to 18 cm 
wide, densely puberulent throughout, flowers in 
distal groups of 2-5, pedicels 3-7 mm long. 
Flowers with cupular calyx 4-6 mm long, 3-4 
mm diam., densely puberulent, margin subentire 
or with 5 minute (0.3 mm) teeth; corolla 26-54 
mm long, pink-lavender to red-purple and white 
in the throat (rarely white throughout), densely 
puberulent externally, tube 8-16 mm diam. at 
mouth, lobes 6-12 mm long; filaments ca. 16 and 
12 mm long, thecae 4-5 mm long. Fruits 15-28 
cm long, 11-15 mm wide, valves flat, yellowish 
brown with minutely puberulent surface (soft to 
the touch), midvein apparent or not; seeds 11-13 
mm long, 30-45 mm wide, central area ca. 12 
mm wide, distal tranparent area of wings ca. 6 
mm wide. 

Common deciduous lianas in seasonally very 
dry deciduous forest formations and open vege- 
tation, 2-300 m elevation. Flowering in December 
to early March (with the great majority of flow- 
ering collections made in January); fruiting in 
January-June. This species ranges from Mexico, 
along the Pacific Coast, to Colombia and Vene- 
zuela. 

Arrabidaea mollissima is recognized by its 
densely soft pubescence, broadly ovate or round- 
ed leaf blades, flowering when leafless, puberulent 
purple or pink corollas, long, narrow, softly pu- 
berulent fruits, and two-winged seeds transparent 
at the lateral margins. Simple tendrils and restric- 
tion to areas of deciduous forest are additional 
distinctions. These plants can be very striking 
when in full flower, after the leaves have been 
shed. Compare this species with A. corallina (with 
glabrous fruits). 



Arrabidaea patellifera (Schldl.) Sandwith, Kew 
Bull. 22: 413. 1968. Bignonia patellifera 
Schldl., Linnaea 8: 516. 1833. Petastoma pa- 
telliferum (Schldl.) Miers, Proc. R. Hort. Soc. 
3: 195. 1863. P. breviflorum Standl., J. Arnold 
Arbor. 11: 128. 1930. Figure 26. 

Lianas, to over 10 m high and 5 cm diam., 
tendrils to 16 cm long, leafy stems 2-8 mm diam., 
densely puberulent with curved hairs ca. 0.2 mm 
long or subglabrous, terete, an interpetiolar line 
often present, gland fields absent at the node; 
pseudostipules rarely present and leaf-like, round- 
ed. Leaves simple or 2-foliolate, petioles 12-35 
mm long (shorter in compound leaves), 1.2-2.3 
mm diam., densely puberulent, petiolules 16-30 
mm long; leaflet blades (5-)6.5-14(-16) cm long, 
(3-)4-8(-12) cm wide, ovate to ovate-oblong or 
ovate-orbicular, apex acuminate, base narrowed or 
rounded and truncate, drying chartaceous, upper 
surface with scattered straight hairs 0.2-0.5 mm 
long, puberulent beneath with thin straight hairs 
ca. 0.4 mm long, venation pinnate to palmate, 2 
veins 3-6/side, ascending. Inflorescences axillary 
or often terminal and 3-parted, 6-28 cm long, to 
30 cm wide near base, peduncles to 8 cm long, to 
4 mm diam., densely puberulent, pedicels 4-7 
mm long. Flower buds white apically, calyx 
broadly spreading (saucer-like), 1-4 mm long, 4- 
7 mm wide, margin entire and undulate, sparsely 
puberulent externally; corolla 23-40 mm long, 
red-violet to magenta or lavender (pink-white in 
the throat), glabrous externally except for the 
lobes, narrowed (2 mm) base of the tube 6-8 mm 
long, mouth 8-1 1 mm wide, lobes 5-1 1 mm long, 
ca. 7 mm wide; filaments ca. 12 and 9 mm long, 
thecae 1.5-2 mm long. Fruits 1 1-39 cm long, 9- 
14 mm wide, linear, surface of valves flat, smooth, 
midvein and margins slightly raised; seeds 8-10 
mm long, 20-36 mm wide, central area 1 1 mm 
wide, wings not clearly differentiated. 

Common evergreen lianas in seasonally very 
dry deciduous and partly deciduous forests of the 
Pacific slope (rarely collected in evergreen forests 
of the Caribbean slope), 1-1200 m elevation. 
Flowering primarily in July-October with the ma- 
jority of collections made in August; fruiting in 
January-March. This species ranges from Mexico 
to Brazil. 

Arrabidaea patellifera is recognized by its 
climbing habit with simple tendrils, simple or bi- 
foliolate puberulent leaves, July-October flower- 
ing period, saucer-like (patelliform) calyx, laven- 
der to magenta corollas, and linear flattened fruits 



114 



FIELDIANA: BOTANY 



with thin-winged seeds. The frequent presence of 
simple leaves, strongly ascending secondary veins 
in a subpalmate pattern, domatia, and the occa- 
sional presence of rounded leaf-like little pseu- 
dostipules are additional distinctions. The flower 
buds are unusual with their pubescent distal co- 
rolla lobes whitish, in contrast to the glabrous ma- 
genta tube. 

Arrabidaea pubescens (L.) A. Gentry, Brittonia 
25: 239. 1973. Bignonia pubescens L., Sp. PI., 
ed. 2, 2: 870. 1763. Petastoma pubescens (L.) 
Miers, Proc. R. Hort. Soc. 3: 195. 1863. A. lun- 
dellii Standl.. Publ. Field Mus. Bot. 8: 48. 1930. 

Lianas to 20 m high and 5 cm diam., tendrils 
to 7 cm long, leafy stems 2-7 mm diam., grayish 
with minute (0.05-0. 1 mm) appressed hairs, glan- 
dular fields usually present at the nodes; pseudo- 
stipules small or absent. Leaves 2- or 3-foliolate. 
petioles 10-22 mm long, 0.9-2.3 mm diam., mi- 
nutely appressed puberulent, grayish, petiolules 
12-23 mm long; leaflet blades 4-13 cm long, 
2.3-8 cm wide, ovate to ovate-elliptic or narrowly 
ovate, apex acuminate, base obtuse or rounded 
and truncated, drying chartaceous and grayish be- 
neath, upper surface subglabrous or with minute 
(< 0.1 mm) hairs, lower surface with minute or 
longer (0.4 mm) hairs, 2 veins 4-6 side. Inflo- 
rescences to 35 cm long, open terminal or axillary 
panicles, peduncles ca. 6 cm long, densely ap- 
pressed puberulent, pedicels 3-5 mm long, ca. 0.4 
mm diam., grayish puberulent. Flowers with ca- 
lyx 3-5 mm long, 2.5-4 mm diam., tubular-cu- 
pulate, margin subentire, grayish puberulent; co- 
rolla 18-33 mm long, funnelform-campanulate, 
lavender to lilac, minutely puberulent externally, 
5-9 mm wide at mouth, lobes 8-10 mm long; 
filaments ca. 10 and 8 mm long. Fruits (7-) 14- 
27 cm long, 9-1 1 mm wide, linear-oblong, 
smooth or slightly pitted, midrib and margins 
raised; seeds 6-9 mm long, 17-32 mm wide, cen- 
tral area 5-6 mm wide, wings transparent. 

Arrabidaea pubescens has not been collected 
from the region between northeastern Guatemala 
and central Panama, but it may occur in Costa 
Rica. This species is distinguished by the simple 
tendrils, nodes with gland fields, minute dense ap- 
pressed hairs that give the undersides of the leaves 
and petioles a grayish appearance, bi- or tri-fo- 
liolate leaves, and linear fruits. The smaller lilac 
corollas produced in July-September, truncated 
grayish calyx tubes, and seeds with central brown 
area clearly demarcated from the transparent 



wings are other noteworthy characteristics. This 
species prefers seasonally dry lowland forests 
along the Caribbean coast; it ranges disjunctly 
from Tamaulipas, Mexico, to Brazil. 

Arrabidaea verrucosa (Standl.) A. Gentry, Sel- 
byana 2: 43. 1977. Adenocalymma verrucosa 
Standl.. Publ. Field Columb. Mus., Bot. Ser. 4: 
323. 1929. Martinella verrucosa (Standl.) 
Standl., Contrib. Arnold Arbor. 5: 138. 1933. 
Scobinaria verrucosa (Standl.) Seibert, Carne- 
gie Inst. Washington Publ. 522: 408. 1940. Fig- 
ure 18. 

Lianas to 35 m high, to 10 cm diam., tendrils 
to 20 cm long, leafy stems 2-7 mm diam., sparse- 
ly and minutely (0.1 mm) puberulent (more rarely 
with hairs to 1 mm long), gland fields and inter- 
petiolar lines present or absent at nodes. Leaves 
2-foliolate, petioles 14-23(-55) mm long, 0.7-2.3 
mm diam., minutely papillate puberulent, petio- 
lules 12-28(-45) mm long; leaflet blades 6-17 
(-22) cm long, 3-9(-l 1 ) cm wide, ovate to ovate- 
elliptic, narrowly elliptic or elliptic-oblong, apex 
acute to acuminate, base acute to slightly rounded, 
drying chartaceous and often dark brown, upper 
surface subglabrous with minute hairs on the mid- 
vein, lower surface with small (0.2-0.5 mm) 
straight hairs or branched along the veins, larger 
hairs often present in vein axils beneath (domatia), 
venation pinnate or subpalmate, 2 veins 3-5/side, 
strongly ascending. Inflorescences axillary, 5-15 
cm long, few-flowered panicles, peduncles 3-5 
cm long, sparsely papillate-puberulent, lenticel- 
late, pedicels 6-14 mm long, ca. 0.8 mm diam. 
Flowers with tubular-campanulate calyx, 14-23 
mm long, 4-1 1 mm diam., glabrous or pubescent 
with hairs to 0.7 mm long near the base, margin 
2-lipped or with shallow (1 mm) sinuses separat- 
ing the rounded lobes; corolla 35-60(-79) mm 
long, tubular-campanulate, lobes magenta to pur- 
ple with the tube white or pink, puberulent exter- 
nally, mouth 11-20 mm wide, lobes 8-18 mm 
long; filaments ca. 1 8 and 1 2 mm long, thecae 2- 
3 mm long. Fruits ( 1 2-)22-42(-5 1 ) cm long, 15- 
28 mm wide, linear-oblong, valves drying dark 
and verrucose-tubcrculate, tubercles 0.3-2 mm 
high; seeds 15-20 mm long, 38-57 mm wide, 
central area poorly delimited. 

Plants of evergreen rain forest formations on 
both the Caribbean and Pacific slopes, 1-700 
(-1300) m elevation. Flowering in May-Novem- 
ber (with the great majority of collections in July); 



BURGER: FLORA COSTARICENSIS 



115 



fruiting throughout the year. This species ranges 
from Mexico to Bolivia. 

Arrabidaea verrucosa is recognized by the 
climbing habit with simple tendrils, bifoliolate 
leaves, small inflorescences, long-tubular calyx, 
puberulent pink or white and purple corollas, and 
narrow fruits with dark rasp-like verrucose-tuber- 
culate valves. The wet-season flowering period 
and wet forest habitats are also characteristic. This 
material was placed under Scobinaria japurensis 
(DC.) Sandwith by Gentry in Flora of Panama 
(1973b), but he later decided that that name be- 
longs to a species restricted to Amazonian South 
America (see: Selbyana 2: 45, 1977). 



Callichlamys Miquel 

Lianas or shrub-like when young, climbing 
with simple tendrils, stems terete, with 4 phloem 
arms in cross-section, an interpetiolar line and 
glandular fields absent at the node, pseudostipules 
absent (pseudostipule-like structures present at the 
base of axillary shoots). Leaves opposite, 2- or 3- 
foliolate, terminal leaflet sometimes replaced by a 
tendril, petiolate, leaflets petiolulate, margins en- 
tire, venation pinnate. Inflorescences axillary 
(terminal) racemes, with 2-12 mostly opposite 
flowers, peduncles becoming woody, bracts sub- 
tending the pedicels lanceolate. Flowers large, ca- 
lyx tubular-inflated, with spongy texture, irregu- 
larly lobed and bilabiate; corolla tubular-campan- 
ulate, yellow, glabrous externally, lobes broadly 
rounded; stamens 4, of 2 lengths, anthers gla- 
brous, thecae divaricate, staminode absent; ovary 
2-locular, ovules in 4-8 series on each placenta, 
stigma simple, flattened. Fruits compressed ob- 
long capsules, valves woody and smooth, flat- 
tened parallel to the septum; seeds large, the 2 
thin lateral wings poorly differentiated from the 
central area. 

Callichlamys includes a single distinctive spe- 
cies ranging from Mexico to Brazil. 

Callichlamys latifolia (L. C. Rich.) K. Schum. in 
Engler & Prantl., Nat. Pflanzenfam. 4 (3b): 223. 
1894. Bignonia latifolia L. C. Richard, Act. 
Soc. Hist. Nat. Paris 1: 110. 1792. Tabebuia 
latifolia (L. C. Rich.) DC., Rev. Bign. (Bibl. 
Univ. Geneve) 15. 1838. T. speciosa Standl., 
Publ. Field Columb. Mus., Bot. Ser. 8: 49. 
1930. C. garnieri Standl. & L. O. Williams, 
Ceiba 3: 130. 1952. Figure 19. 



Lianas to over 30 m high and 8 cm diam. (rare- 
ly shrubs), leafy stems 3-9 mm diam., minutely 
puberulent at first but soon glabrescent with dark 
lenticels (0.5-1.5 mm) against a grayish surface. 
Leaves mostly 2-foliolate (also 3-foliolate), peti- 
oles 3-17 cm long, 1.7-4.5 mm diam., at first 
minutely puberulent but later resembling the 
stems, petiolules 8-35 mm long, sulcate above; 
leaflet blades 7-24(-36) cm long, 4-14(-20) cm 
wide, broadly elliptic to broadly ovate-elliptic or 
elliptic-oblong, apex acuminate to caudate-acu- 
minate, narrowed tip to 3 cm long, base obtuse to 
rounded, drying stiffly chartaceous, glabrescent 
on both surfaces or with small (0.4-0.5 mm) 
branched/stellate hairs beneath, often with dense 
hairs in vein axils beneath (domatia), 2 veins 5- 
1 I/side. Inflorescences 3-17 cm long, peduncles 
4-14 mm long, 2-3 mm diam., minutely puber- 
ulent with appressed brownish hairs, bracts to 5 
mm long, caducous, pedicels 8-24 mm long, ca. 
1 mm diam., drying dark. Flowers with calyx 28- 
48(-65) mm long, 8-30(-42) mm diam., tubular- 
inflated, rounded at the base, greenish white to 
yellow, lobes 12-20 mm long; corolla 5-9(-ll) 
cm long, bright yellow, tube 15-22(-28) mm wide 
near the mouth, marked with reddish lines within, 
lobes 1.5-3 cm long; filaments ca. 15-20 and 25- 
30 mm long, thecae 2-3 mm long; stigma ca. 4 
X 1.8 mm. Fruits 12-32 cm long, 6-1 1 cm wide, 
ca. 1 cm thick, oblong or elliptic-oblong with 
rounded ends, flattened, surface smooth; seeds 
2.5-4 cm long, 6-1 1 cm wide, brownish. 

Plants of evergreen rain forest formations of 
both Caribbean and Pacific slopes and also in sea- 
sonally very dry deciduous forest areas, 5-600 
(-1000) m elevation. Flowering collections have 
been made in February-August in Costa Rica 
(flowering primarily in October-November in 
Panama). The species ranges from Veracruz, 
Mexico, to Brazil. 

Callichlamys latifolia is recognized by its 
climbing habit with simple tendrils, large spongy 
calyx, large yellow corolla, and flattened oblong 
woody fruits with large brown seeds. The grayish 
stems with dark lenticels, lack of gland fields and 
interpetiolar lines at nodes, and larger leaf blades 
often with domatia in vein axils (or with branched 
hairs beneath) are useful in identifying sterile ma- 
terial. Collections from dryer forests of the Pacific 
slope tend to have the undersurface of the leaflets 
covered with conspicuous branched hairs. These 
collections look quite different from those of the 
lowland rain forests with subglabrous leaves. The 
type of C. garnieri (Gamier 130 F, from the Si- 



116 



FIELDIANA: BOTANY 



erra de Managua, Nicaragua) has leaves with 
branched hairs and unusually large (15 cm) flow- 
ers subtended by reduced leaves on a terminal 
branchlet. However, the fact that no other similar 
collection has been seen suggests that Gentry 
(1994) was correct in considering it an aberrant 
individual of this species. 



Ceratophytum Pittier 

Lianas, climbing with tendrils, stems terete, 
with 4 phloem areas in cross-section, nodes with 
interpetiolar ridges and glandular fields; pseudo- 
stipules of subulate scales. Leaves opposite, 3- 
foliolate or 2-foliolate and the terminal leaflet re- 
placed by a tendril or tendril scar, leaflets petiol- 
ulate, margins entire, venation pinnate, webbed 
tissue (domatia) sometimes present in vein axils. 
Inflorescences terminal, corymbose panicles or 
condensed fasicles, with 1-20 flowers, peduncles 
lepidote, bracts small, pedicels well developed. 
Flowers with tubular calyx, coriaceous, apex 
truncated, with short-linear glandular fields near 
the rim; corolla tubular-funnelform and slightly 
2-lipped, yellow to yellowish white, minutely pu- 
berulent externally, 5-lobed; stamens 4, of 2 
lengths, thecae straight, divaricate, a staminode 
present; disc cupular; ovary 2-locular, ovules in 
6-8 series on each placenta, stigma simple. Fruits 
oblong-linear capsules, tetrangular in cross-sec- 
tion, expanded and rounded at the base, tapering 
to the apex, valves parallel to the septum (but not 
flattened), smooth; seeds with 2 lateral papery 
wings, brownish, not clearly demarcated from the 
central area. 

Ceratophytum includes a single species ranging 
from Mexico and the Caribbean to Guyana and 
Bolivia. 

Ceratophytum tetragonolobum (Jacq.) Sprague 
& Sandw., Kew Bull. 1934: 222. 1935. Bigno- 
nia tetragonoloba Jacq., Fragm. Bot. 36, tab. 
40, fig. 2. 1800-1806. Anemopaegma toba- 
gense Urb., Fedde Repert. 14: 311. 1916. Ad- 
enocalymma heterophyllum Si ami I.. Publ. Field 
Columb. Mus., Bot. Ser. 8: 49. 1930, not 
Kranzl. C. tobagense (Urb.) Sprague & Sandw., 
Kew Bull. 1933: 322. 1933. Ad. standleyanum 
Lundell, Carnegie Inst. Washington Publ. 478. 
221. 1937, based on Ad. heterophyllum Standl. 
Figure 19. 



Lianas to 20 m high, stems up to 7 cm diam.. 
tendrils 1-3 mm diam., to 22 cm long, often trifid, 
leafy stems 1.3-7 mm diam., subglabrous, nodes 
with gland fields; pseudostipules 1-3 mm. Leaves 
2- or 3-foliolate. petioles 4-11 cm long, 1.2-2 
mm diam.. subglabrous with minute peltate hairs, 
petiolules 8-30 mm long; leaflet blades 6-18 cm 
long, 3.5-9(-15) cm wide, elliptic-oblong to 
broadly elliptic, apex short-acuminate or obtuse, 
base obtuse to rounded, drying thin-chartaceous, 
with scattered minute (0.1 mm) peltate hairs be- 
neath, 2 veins 3-7/side, minor venation often 
prominent above, domatia occasionally present. 
Inflorescences terminal, 1-8 cm long, with 2-20 
flowers, peduncles 1-10 mm long, bracts minute, 
pedicels 8-17 mm long, ca. 0.7 mm diam., with 
minute peltate hairs. Flowers with calyx 8-12 
mm long, 5-9 mm wide, tubular with subentire 
rim, appearing glabrous but with minute peltate 
hairs and ciliolate rim, with glands near the mar- 
gin; corolla 5-8 cm long, tubular-funnelform with 
tube gradually expanded to 16-21 mm wide at the 
mouth, white to yellow, with a dense covering of 
minute glandular hairs drying pale yellowish 
brown, lobes 15-22 mm long, rounded or obtuse; 
filaments 10-16 and 16-24 mm long, thecae 4 X 
1 mm; ovary ca. 5 mm long, with peltate hairs. 
Fruits 16-45 cm long. 25-37 mm wide, 15-26 
mm thick, linear-oblong, narrowed to the apex, 
yellowish and lenticellate; seeds 8-13 mm long, 
28-45 mm wide, lateral wings brown or gray, not 
clearly differentiated from central area, translu- 
cent. 

Uncommon evergreen climbers of deciduous, 
partly deciduous, or wet evergreen lowland rain 
forest formations, 1-500 m elevation. Flowering 
in January-June and October in Central America; 
fruiting throughout the year. This species ranges 
from Mexico to Bolivia. 

Ceratophytum tetragonolobum is recognized by 
its climbing habit, three-tipped tendrils, bi- or tri- 
foliolatc opposite leaves, compact terminal inflo- 
rescences, truncated calyx tube with distal glands, 
white or yellowish corolla minutely pubcrulent on 
the exterior, thick narrow woody fruits, and seeds 
with poorly differentiated lateral wings. Young 
stems with prominent glandular fields and usually 
four subulate pseudostipules at the nodes and oc- 
casional domatia are additional characteristics. 



Clytostoma Miers ex Bureau 

Lianas climbing with simple tendrils, stems 4- 
angled or terete, with 8 phloem areas in cross- 



BURGER: FLORA COSTARICENSIS 



117 



section, nodes lacking gland fields; pseudostipules 
usually a small condensed axis of subulate cata- 
phylls. Leaves 2-foliolate (rarely simple), with or 
without a terminal simple tendril, petiolate, mar- 
gins entire, venation pinnate. Inflorescences ter- 
minal or axillary, few-flowered fascicles, cymes, 
panicles, or the flower solitary, subtended by 
bracts similar to the cataphylls. Flowers with cu- 
pular or campanulate calyx truncated distally and 
minutely 5-denticulate or with 5 linear lobes, 
subglabrous to minutely lepidote or puberulent; 
corolla tubular-funnelform, white to purple, glan- 
dular-puberulent or with minute peltate hairs ex- 
ternally; stamens 4, included, filaments of 2 
lengths, anthers glabrous, thecae straight and di- 
varicate, a staminode present; disc absent; ovary 
2-locular, ovules 2(-4)-seriate in each locule. 



Fruits ellipsoid to suborbicular capsules, valves 
convex or flattened parallel with the septum, 
woody and echinate with curved spines; seeds 
transverse-oblong, corky, flattened but not 
winged. 

Clytostoma includes nine South American spe- 
cies and one that ranges from Mexico to Brazil. 
Echinate woody fruits and flowers lacking a nec- 
tariferous disc are distinctive characteristics. Cly- 
tostoma callistegioides (Cham.) Bureau ex Gri- 
sebach of southern South America was collected 
in a Costa Rican garden in 1935 (Brenes & Ze- 
ledon 99; Standley, 1938), but we have seen no 
further evidence of its presence in Central Amer- 
ica. It differs from all our other species of Big- 
noniaceae in Costa Rica by having prominent lin- 
ear calyx lobes arising from a truncated calyx 
margin (see key below). 



Key to the Species of Clytostoma 

la. Larger leaf blades > 12 cm long; calyx lobes 0.2-0.6 mm long; native C. binatum 

Ib. Larger leaf blades < 10 cm long; calyx lobes 1-4 mm long, subulate to linear; garden ornamental 
(not included in descriptions, see above) C. callistegioides 



Clytostoma binatum (Thunb.) Sandw., Recueil 
Trav. Bot. Neerl. 34: 235. 1937. Bignonia bi- 
nata Thunb., PI. Bras. 3: 35. 1821. Adenoca- 
lymma ocositensis J. D. Smith, Bot. Gaz. 18: 
209. 1893. C. isthmicum Pittier, Contrib. U.S. 
Natl. Herb. 18: 257, pi. 106. 1917. Petastoma 
ocositense (J. D. Smith) Kranzl., Fedde Repert. 
17: 61. 1935. Figure 18. 

Lianas to 5 cm diam., sometimes forming 
clumps 1-2 m high, tendrils to 12 cm long, leafy 
stems 2-7 mm diam., rounded-tetrangular, gla- 
brous or very minutely (0.05 mm) puberulent, 
lenticellate; pseudostipules forming an axillary 
cone 3-5 mm long. Leaves 2-foliolate, petioles 
6-18(-31) mm long, 1-1.5 mm diam., petiolules 
4-6(-21) mm long, usually glabrous; leaflet 
blades 7-16(-19) cm long, 2-6(-8) cm wide, el- 
liptic to elliptic-oblong or oblong, apex acumi- 
nate, base acute to cuneate, drying chartaceous 
and with minor venation raised on both surfaces, 
usually glabrous on both surfaces, 2 veins 5-87 
side. Inflorescences mostly axillary, 0.4-8 cm 
long, peduncle subtended by subulate bracts (sim- 
ilar to the pseudostiples), pedicels 6-32 mm long, 
0.4-0.8 mm diam., glabrous. Flowers with cu- 
pular calyx 4-8 mm long, 3-5 mm diam., gla- 
brous, with many longitudinal veins or smooth, 



truncate or with teeth 0.2-0.6 mm long; corolla 
45-80 mm long, lilac to blue-purple (white), 
sparsely and minutely (0.05 mm) puberulent ex- 
ternally, tube 12-24 mm diam. at the mouth, 
throat white internally, lobes 6-16 mm long; fil- 
aments ca. 10-16 and 16-20 mm long, thecae ca. 
3 mm long; ovary covered by thick rounded hairs. 
Fruits 5-9 cm long, 3-6 cm wide, broadly elliptic 
to suborbicular, discoid-lenticular, surfaces of the 
valves covered with echinate spines to 8 mm long, 
slender-tipped and slightly curved from a broad 
base; seeds 12-19 mm long, 19-24 mm wide, 
suborbicular, brown. 

Evergreen or deciduous lianas found mostly in 
swamp forests, river edges, and seasonally inun- 
dated areas in both wet evergreen and dry decid- 
uous forest formations, 1-200 m elevation. Flow- 
ering throughout the year but most frequently in 
September-November. This species ranges from 
Mexico to Brazil. 

Clytostoma binatum is recognized by its climb- 
ing habit with simple tendrils, bifoliolate leaves, 
short few-flowered inflorescences, essentially gla- 
brous calyx cups with minute teeth distally, lilac 
to blue-purple (rarely white) corolla sparsely mi- 
nutely puberulent on the exterior, short rounded 
fruit covered with woody echinate projections, 



118 



FIELDIANA: BOTANY 



and wingless seeds. The pseudostipule resembles 
a minute bromeliad plantlet in leaf axils and is 
helpful in recognizing this species. The seeds are 
apparently adapted to water dispersal; conse- 
quently, the plants are restricted to low elevations. 



Crescentia Linnaeus 

Small to medium-size trees, with many thick 
twisted branches, leafy stems becoming terete 
with enlarged nodes, gland Melds absent. Leaves 
alternate or more often in fascicles of 3-9 on con- 
densed short-shoots in the axils of fallen leaves, 
simple or 3-foliolate (1- or 2-foliolate), petioles of 
compound leaves winged and leaf-like, blades 
with entire margins, subglabrous or sparsely pu- 
berulent, venation pinnate. Inflorescences borne 
on older thick branches or trunks, peduncles ab- 
sent, flowers 1-3 in sessile fascicles, pedicels 
short and thick. Flowers with large tubular calyx 
that usually splits into 2 halves at maturity, thick, 
glabrous or minutely puberulent; corolla broadly 
campanulate and bilabiate, narrowed only at the 
base, thick textured, usually with minute peltate 
hairs on the outer surface, tube with a transverse 



fold midway across the lower side, distally 5- 
lobed or variously erose along the margin; sta- 
mens 4, subexserted, filaments of 2 lengths, disc 
large, staminode present; ovary lepidote, 1-locular 
with ovules multiseriate on 4 parietal placentas, 
stigma flat and expanded. Fruits large pepos (a 
gourd-like calabash) with a hard corky pericarp, 
subglobose to ovoid, pulpy within; seeds angular, 
without wings, less than 10 mm long, embedded 
in the pulp. 

Crescentia is a genus of six species in Mexico, 
the West Indies, Central America, and (one spe- 
cies) Amazonia. The genus is distinguished by the 
tree habit, mostly fasciculate leaves on alternate 
short-shoots, few large thick-textured cauliflorous 
flowers, ovaries with a single locule and parietal 
placentation, and rounded indehiscent fruits with 
seeds embedded in pulp. The flowers are foul- 
smelling and adapted for bat pollination. The 
pulp-containing fruits are dispersed by mammals. 
The hard outer shell of the fruits has been used 
to make cups, ladies, and bowls (see Standley & 
Williams, 1974). The differentiation of the two 
Central American species is not complete, and hy- 
brids have been reported. (See Flora Neotropica 
treatment; Gentry, 1980.) 



Key to the Species of Crescentia 

la. Stems with 3-foliolate leaves, simple leaves may also be present; fruits up to 10 cm diam. and 
globose; wild plants (rarely cultivated) C. alata 

Ib. Stems lacking 3-foliolate leaves, all the leaves simple; fruits > 13 cm diam. and globose to ovoid; 
plants widely cultivated C. cujete 



Crescentia alata Kunth in H.B.K., Nov. Gen. Sp. 
3: 158. 1819. Parmentiera alata (Kunth in 
H.B.K.) Miers, Trans. Linn. Soc. 26: 166. 1868. 
C. ternata Sesse & Moc., La Naturaleza, ser. 2. 
1 (append.); 94. 1889. Figure 11. 

Trees to 10 m tall, trunks to 25 cm diam., leafy 
stems 3-18 mm diam., glabrous, the alternate 
nodes becoming enlarged and prominent and sub- 
tending the condensed leafy short-shoots. Leaves 
mostly in fascicles of 3-9, simple or 3-foliolate 
(rarely 1- or 2-foliolate), simple leaves sessile, 
compound leaves with winged petioles 2.5-1 1 cm 
long, 4-11 mm wide, petiolules absent; leaf 
blades 4-15 cm long, 5-35 mm wide, oblanceo- 
late to narrowly obovate (small leaves often 
rounded), leaflet blades 1.5-10 cm long, 4-23 mm 
wide, oblanceolate, apex rounded or emarginate. 



gradually narrowed to the acute or cuneate base, 
drying subcoriaceous and grayish green, glabrous. 
Inflorescences of 1-3 flowers borne directly on 
the surfaces of larger stems and trunks, pedicels 
5-10 mm long, 1.3 mm diam., subtended by small 
(1.5 mm) bracts, glabrous or with minute peltate 
hairs. Flowers with unpleasant aroma, calyx 14- 
28 mm long, 9-14 mm diam., usually splitting 
into 2 subequal halves, glabrous or with few mi- 
nute peltate hairs, greenish; corolla 4-6 cm long, 
tube narrowed only near the base, 22-30 mm 
diam.. greenish purple to reddish white, surface 
glabrous or with minute peltate glands, 5-lobed or 
the margin irregularly toothed; filaments ca. 30 
and 1 8 mm long, thecae 5-6 mm long. Fruits 7- 
10 cm diam., globose, greenish, surface smooth; 
seeds 6-7 mm long, 7-9 mm wide. 

Plants of seasonally dry deciduous forest for- 



BURGER: FLORA COSTARICENSIS 



119 



inations of the Pacific lowlands, 5-800 m eleva- 
tion. Flowering throughout the year (collected 
mostly in January-September). This species rang- 
es from Mexico along the Pacific slope to north- 
western Costa Rica (introduced in Panama). 

Crescentia alata is recognized by the mix of 
simple and trifoliolate leaves with all the larger 
blades being oblanceolate or narrowly obovate, 
the few flowers coming directly from the trunk, 
thick calyx usually splitting in two, and large glo- 
bose gourd-like fruits borne from large branches 
and trunks. The trifoliolate leaves, with sessile ob- 
lanceolate leaflets and winged petiole, resemble a 
Christian cross and were mentioned in early Span- 
ish accounts of Mexico (Standley & Williams, 
1974, p. 187). The trees are noteworthy for their 
short compact form and dense branching, often 
contrasting with the flat, open, grazed land around 
them. The dense branching is often the habitat for 
many epiphytes. There may be hybridization be- 
tween this species and the widely planted C. cu- 
jete. This species is called morro or jicaro, but 
jicaro is also the name of the following species. 

Crescentia cujete L., Sp. PI. 2: 626. 1753. Figure 
11. 

Trees to 10 m tall, with dense rounded crowns, 
trunks to 40 cm diam., branches often crooked 
with mostly thick branchlets, leafy stems 6-30 
mm diam., glabrous, nodes becoming thickened 
and subtending the leafy short-shoots. Leaves 
usually fasciculate on elevated alternate nodes, 1- 
15 and varying in size at the same node, unifoli- 
olate, sessile, glabrous; leaf blades 3-22(-26) cm 
long, l-6(-7.5) cm wide, oblanceolate to narrow- 
ly obovate, apex acute to acuminate or rounded 
(emarginate), gradually narrowing to the acute or 
cuneate base, usually drying subcoriaceous and 
pale gray, glabrous or with minute simple or 
branched hairs along the midvein beneath, 2 
veins 5-14/side. Inflorescences of 1 or 2 flowers 
borne on older branches and trunks, pedicels 9- 
30 mm long, 0.7-1.7 mm diam., glabrous or with 
few minute peltate hairs. Flowers with calyx 18- 
34 mm long, splitting into 2 subequal lips 12-24 
mm wide; corolla 55-70 mm long, dull white to 
yellowish white, with purple lines, tube 18-25 
mm diam., lobes triangular with narrow tip, sub- 
glabrous or with minute peltate hairs externally; 
filaments ca. 30 and 10 mm long, anthers 6-9 mm 
long, slightly divergent. Fruits 13-25 cm diam. 
or to 30 cm long, subglobose or ovoid, surface 



smooth, subglabrous; seeds 7-8 mm long, 4-6 
mm wide. 

Widely cultivated plants in regions of both de- 
ciduous and wet evergreen forest formations; 1- 
1200 m elevation. Flowering irregularly through- 
out the year. This species may have been indige- 
nous to southeastern Mexico, but it is now culti- 
vated throughout tropical America, and its 
original range is unknown. 

Crescentia cujete is recognized by its fascicles 
of narrow simple leaves from alternate thickened 
nodes, oblanceolate blades, few flowers borne di- 
rectly on larger stems and trunks, large calyx usu- 
ally splitting into two, large thick white to yellow- 
ish corolla, and larger globose gourd-like fruit 
(calabash). The hard shell of the pericarp is often 
used as a cup, ladle, or other kitchen utensil; the 
round, dried, empty fruits are often incised or 
painted for ornament. The pulp of the fruit has 
been used as a purgative, according to Standley 
(1938), who reported that the oval fruits are called 
guacales and the globose ones jicaras. The wood 
is hard but not durable. Jicaro, calabacero, cala- 
bash, and wild calabash are common names. 



Cydista Miers 

REFERENCE W. D. Hauk, A review of the ge- 
nus Cydista. Ann. Missouri Bot. Gard. 84: 815- 
840. 1997 (1998). 

Lianas climbing with simple tendrils, stems 
terete to tetrangular, with 8-16 phloem areas in 
cross-section, nodes usually with interpetiolar 
lines, gland fields absent at the nodes; pseudos- 
tipules absent or leaf-like. Leaves opposite, sim- 
ple or 2-(4-) foliolate, petiolate, margins entire, 
venation pinnate, glands or gland fields often 
present in basal vein axils beneath. Inflores- 
cences terminal or axillary, cymose racemes or 
panicles (thyrses), often with fewer than 15 flow- 
ers, bracts minute or absent, flowers pedicellate. 
Flowers with cupular or campanulate calyx, gla- 
brous to glandular puberulent or with minute pel- 
tate hairs, glands present or absent distally, mar- 
gin truncate and entire to 5-lobed or irregularly 
split; corolla funnelform to tubular-campanulate, 
2-lipped, lavender to purple or white, usually 
with minute glandular or peltate hairs externally, 
with 2 upper and 3 lower lobes, broadly rounded; 
stamens 4, included, filaments of 2 lengths, an- 
thers glabrous, thecae divaricate; disc absent; 
ovary with glandular or peltate hairs on the sur- 



120 



FIELDIANA: BOTANY 



face, 2-locular, ovules in 2 series in each locule, 
stigma 2-lobed. Fruits linear to narrowly oblong 
woody capsules, valves 2, flattened parallel to 
the septum, septicidally dehiscent, surface usu- 
ally smooth, glabrate to puberulent; seeds flat 
and transversely oblong, the central body of the 
seed not clearly differentiated from the 2 lateral 
wings. 

Cydista is a Neotropical genus of six species, 
ranging from Mexico and the West Indies to Bra- 
zil and Bolivia. Five species have been found in 



Costa Rica, but two are known from only single 
collections. The mix of simple and bifoliolate 
leaves, presence of glands or gland fields in basal 
vein axils (on underside of leaf), narrow capsules, 
and brown two-winged seeds with central area not 
clearly differentiated help distinguish the genus. 
The absence of a disc is consistent with reports 
that the flowers do not produce nectar and deceive 
pollinators by multiple "big-bang" flowering of a 
few days. Compare species of Arrabidaea and 
Clytostoma. 



Key to the Species of Cydista 

la. Fruits linear or linear-oblong, 10-25 mm wide, to 45 cm long; commonly encountered plants in 

areas of dry deciduous, partly deciduous, or (less often) evergreen forests 2 

Ib. Fruits narrowly oblong, 27-43 mm wide, to 24 cm long; more rarely encountered plants of evergreen 

or partly deciduous forests 4 

2a. Stems with persisting rounded pseudostipules 3-15 mm long [leaf- venation mostly palmate; 

flowering mostly in June-August; fruits with thin flat valves] C. diversifolia 

2b. Pseudostipules usually absent or not broadly ovate 3 

3a. Plants usually flowering in April-May while leafless, in dry deciduous forest formations; calyx 
usually split into 2 or 3 lobes; fruits with 2 longitudinal ridges on the flat valve surface; leaves 
simple or 2-foliolate, leaf venation mostly palmate-subpalmate, lower leaf surfaces never dense- 
ly hirtellous C. heterophylla 

3b. Plants flowering throughout the year with leaves fully expanded, in both deciduous and ever- 
green habitats; calyx usually entire distally; fruits smooth and flat on the valve surfaces; leaves 
usually only 2-foliolate, venation pinnate to subpalmate; leaf surfaces densely hirtellous to 

pilose or subglabrous and lustrous C. aequinoclialis 

4a. Valves of fruit with smooth surface; lower leaf surface without glands or gland fields in basal vein 

angles; leaflet blades mostly 2-5 cm wide; Mexico to Costa Rica C. potosina 

4b. Valves of fruit with longitudinally wrinkled surface; lower leaf surface usually with glands in basal 
vein angles; leaflet blades mostly 5-15 cm wide; Costa Rica to Bolivia C. lilacina 



Cydista aequinoctialis (L.) Miers, Proc. R. Hort. 
Soc. 3: 191. 1863. Bignonia aequinoctialis L., 
Sp. PI. 2: 623. 1753. B. sarmentosa Bertol., Fl. 
Guatimal. 25. 1840. B. sarmentosa var. hirtella 
Benth., Bot. voy. Sulphur 128. 1845. C. sar- 
mentosa (Bertol.) Miers, Proc. R. Hort. Soc. 3: 
192. 1863. Levya nicaraguensis Bureau ex 
Baillon, Hist. PI. 10: 29. 1888. Arrabidaea gua- 
temalensis K. Schum. & Loes., Bot. Jahrb. Syst. 
23: 129. 1896. Arrabidaea pseudochica 
Kra'nzl., Repert. Spec. Nov. Regni Veg. 17: 19. 
1921. Anemopaegma tonduzianum Kra'nzl., 
Fedde Repert. 17: 116. 1921. Cydista pubescens 
Blake, Contrib. U.S. Natl. Herb. 24: 23. 1922. 
Arrabidaea isthmica Standl., J. Wash. Acad. 
Sci. 15: 461. 1925, pro parte (leaves). C. ae- 
quinoctialis var. hirtella (Benth.) A. Gentry, 
Ann. Missouri Bot. Card. 60: 838. 1973. Figure 
24. 



Lianas to over 20 m high, to 8 cm diam., ten- 
drils 4-21 cm long, leafy stems 2-7 cm diam., 
glabrous to sparsely or densely pilose with 
straight hairs to 1 mm long (in var. hirtella), 
young stems tetragonal; pseudostipules usually in- 
conspicuous (leaf-like to 4 mm long), caducous. 
Leaves simple or 2-foliolate, petioles 1 1-47 mm 
long, 1 .2-2 mm diam., petiolules 9-40 mm long, 
glabrous to sparsely or densely pilose (var. hirtel- 
la); leaflet blades 6-16 cm long, 3-10 cm wide, 
ovate-elliptic to elliptic, apex acuminate, base ob- 
tuse or rounded and subtruncate, drying stiffly 
chartaceous, glabrous to sparsely puberulent 
above, minutely puberulent on the veins to dense- 
ly pubescent beneath (in var. hirtella), 2 veins 4- 
7/side, basal veins often strongly ascending (sub- 
palmate). Inflorescences terminal or axillary, 
short (3-15 cm) panicles of 3-15 flowers (flowers 



BURGER: FLORA COSTARICENSIS 



121 



rarely solitary), peduncles 1.5-10 cm long, 
sparsely to densely puberulent, pedicels 5-25 mm 
long. Flowers with calyx 4-9 mm long, 4-7 mm 
wide, cupular, margin entire or with minute teeth, 
surface glabrous to densely puberulent (in var. 
hirtella), sometimes with gland fields near the dis- 
tal margin; corolla (25-)35-75 mm long, funnel- 
form-campanulate, magenta to pink or white, 
throat often yellowish or with purplish lines with- 
in, minutely lepidote externally, tube (6-) 11 -22 
mm diam. at the mouth, lobes 12-24 mm long, 
14-25 mm wide; filaments 11-18 and 8-13 mm 
long, thecae 3-5 mm long. Fruits 21-45 cm long, 
17-24 mm wide, linear-oblong, ends rounded, 
valves flat with slightly raised submarginal ridges, 
drying dark or blackish, smooth; seeds (8-) 12-20 
mm long, (26-)41-70 mm wide, pale brown 
throughout, sometimes transparent at the tips. 

Common evergreen lianas of lowland wet ev- 
ergreen forest areas, partly deciduous forests, and 
seasonally very dry deciduous formations, 0-800 
m elevation. Flowering throughout the year but 
with a peak in April-May; fruiting mostly in the 
dry season (January-May). This species ranges 
from Mexico and the West Indies to Brazil, Bo- 
livia, and Paraguay. 

Cydista aequinoctialis is recognized by the 
climbing habit with simple tendrils, opposite sim- 
ple or bifoliolate leaves, short, few-flowered inflo- 
rescences, usually entire calyx cups, pink to ma- 
genta or white corollas minutely lepidote on the 
exterior, long narrow flat fruits, and seeds with 
uniform brownish coloring. These plants are usu- 
ally leafless when flowering and fruiting (Gentry, 
1973a). An important identifying characteristic is 
that gland fields are sometimes present in the axils 
of secondary veins on the lower leaf surface. This 
species is extremely variable in leaflet form, flow- 
er size, and vesture; the more densely pubescent 
individuals (formerly called C. sarmentosa or C. 
pubescens) have been designated var. hirtella 
(Bentham) A. Gentry. The more puberulent vari- 
ety is usually encountered in dry vegetation, and 
the glabrous variety is more common in evergreen 
formations. 

Cydista diversifolia (Kunth in H.B.K.) Miers, 
Proc. R. Hort. Soc. 3: 192. 1863. Bignonia di- 
versifolia Kunth in H.B.K., Nov. Gen. Sp. 
quart, ed. 3: 133, folio ed. 3: 104. 1819. Pleon- 
otoma diversifolium (Kunth in H.B.K.) Bureau 
& K. Schum. in Mart., Fl. Bras. 8(2): 274. 
1897. Figure 24. 



Lianas to 5 cm diam, to 20 m high, tendrils 
11-21 cm long, leafy stems 2-7 mm diam., 
strongly tetragonal with 4 longitudinal ridges, 
subglabrous or sparsely pubescent with thin hairs 
to 0.4 mm long; pseudostipules 3-15 mm long, 
leaf-like, rounded-ovate. Leaves 2-foliolate or 
sometimes simple, petioles 18-45 mm long, ca. 1 
mm diam., surface similar to stems, petiolules 10- 
31 mm long; leaflet blades 4-13 cm long, 2.5-8 
cm wide, ovate to broadly ovate, apex acuminate 
with tip to 13 mm long, base rounded and sub- 
cordate to shallow-cordate, drying chartaceous, 
upper surface subglabrous, lower surface glabrous 
to sparsely pubescent with thin hairs ca. 0.4 mm 
long, venation palmate with 3 major veins or sub- 
palmate with 3 or 4 2 veins/side, gland fields of- 
ten present in the basal vein axils beneath. Inflo- 
rescences terminal or axillary, 6-20 cm long, 
panicles with 4-25 flowers, peduncles to 3-9 cm 
long, ca. 1.5 mm diam., similar to stems, lateral 
branches 8-25 mm long, bracts 0.4-1 mm long, 
pedicels 4-12 mm long. Flowers with calyx cups 
3.5-5 mm long, ca. 4 mm diam., entire or split 
along the margin, with minute peltate hairs or thin 
hairs along the margin; corolla 22-46 mm long, 
funnelform-campanulate, magenta to lavender or 
bluish purple, subglabrous externally, tube 8-16 
mm diam. at the mouth, throat white with purple 
stripes within, lobes 7-22 mm long, broadly 
rounded; filaments ca. 16 and 13 mm long, thecae 
2-2.5 mm long. Fruits 27-41 cm long, 10-16 
mm wide, linear, valves flat and smooth, dark 
brown; seeds 10-13 mm long, 28-57 mm wide, 
pale yellowish brown throughout, wings thin. 

Common evergreen vines in seasonally dry de- 
ciduous formations (rarely in evergreen forests), 
5-500 m elevation. Flowering throughout the year 
but most often collected in June-August; fruiting 
in November-March. This species ranges from 
Mexico and the West Indies to Colombia and Ven- 
ezuela. 

Cydista diversifolia is recognized by its climb- 
ing habit with simple tendrils, usually bifoliolate 
leaves with rounded subcordate base and palmate 
venation, open-paniculate inflorescences, wet-sea- 
son flowering, bright lavender to bluish purple co- 
rollas, long narrow flattened fruits, and uniformly 
colored thin seeds. The persisting rounded pseu- 
dostipules (four per node), square stems with 
prominent ridges and hollow in early stages, and 
gland fields in the basal vein axils are additional 
vegetative features. 

Cydista heterophylla Seibert, Carnegie Inst. 
Wash. Publ. 522: 417. 1940. Bignonia lepidota 



122 



FIELDIANA: BOTANY 



Seem., Bot. voy. Herald 179. 1854, non Kunth 
in H.B.K. Figure 24. 

Lianas to 10 m high, to 10 cm diam., tendrils 
to 18 cm long, leafy stems 1.8-5 mm diam., te- 
rete, glabrescent, nodes with or without interpe- 
tiolar lines; pseudostipules 1-2 mm long, conical. 
Leaves simple or 2-foliolate (with or without ten- 
drils), petioles 15-50 mm long, glabrous or 
sparsely puberulent with minute peltate hairs, lon- 
gitudinally striate, petiolules 5-50 mm long; leaf- 
let blades 5-14(-17) cm long, 3-9(-l 1) cm wide, 
broadly to narrowly ovate, apex bluntly obtuse to 
short-acuminate, base rounded and truncate to 
subcordate, drying thin-chartaceous, subglabrous 
on both surfaces, venation palmate with 3 major 
veins or subpalmate, 2 veins 3-5/side, glandular 
fields usually present in basal vein axils beneath. 
Inflorescences mostly axillary to fallen leaves, 
1.5-7(-14) cm long, racemose, peduncles 8-14 
mm long, 0.7-1.8 mm diam., densely covered 
with peltate (lepidote) hairs, pedicels 6-13 mm 
long, vesture like the peduncle. Flowers with ca- 
lyx 3.5-6 mm long, 4-5 mm diam., densely cov- 
ered with peltate hairs (glands sometimes pres- 
ent), margin entire or split into several lobes; co- 
rolla 47-66 mm long, funnelform-campanulate, 
lavender to purple-magenta, glabrous or with mi- 
nute peltate hairs externally, tube 11-17 mm wide 
at the mouth, lobes 9-20 mm long, to 22 mm 
wide; filaments ca. 18 and 12 mm long, thecae 4- 
5 mm long. Fruits 18-33 cm long, 14-25 mm 
wide, linear-oblong, with 2 raised longitudinal 
ridges and raised edges on each flattened valve, 
smooth; seeds 8-13 mm long, 34-64 mm long, 
wings with transparent tips. 

Deciduous lianas of the seasonally very dry de- 
ciduous forest formations and drier areas within 
evergreen formations, 5-300 m elevation (to 1000 
m in Honduras). Flowering in April-May, fruiting 
in November-March. This species ranges from 
Mexico to northern Colombia. 

Cydista heterophylla is recognized by its climb- 
ing habit with simple tendrils, the simple or bi- 
foliolate opposite leaves, short racemose inflores- 
cences with vesture of minute peltate hairs, lav- 
ender or purple corollas, long narrow fruit with 
two central longitudinal ridges and raised edges, 
and winged seeds with transparent distal tips. 
Stem tips of young plants often have four leaves 
(two pairs of simple leaves). The gland fields in 
the axils of basal veins may be difficult to see. 
The short flowering period when leaves are usu- 
ally absent and restriction to deciduous forest ar- 
eas are additional characteristics. 



Cydista lilacina A. Gentry, Mem. New York Bot. 
Card. 29: 277. 1978. Figure 24. 

Lianas climbing with simple tendrils (not 
seen), leafy stems ca. 4 mm diam., terete, gla- 
brous or with few minute hairs, drying black; 
pseudostipules 1-2 mm long. Leaves opposite or 
subopposite, 2-foliolate or simple, petioles 17-50 
mm long, 1.4-3.3 mm diam., subglabrous, drying 
dark, petiolules 9-23 mm long; leaflet blades 5- 
19(-27) cm long, 3-13(-17) cm wide, ovate to 
ovate-oblong, apex obtuse with a small acutp tip, 
base obtuse to rounded and often unequal, drying 
stiffly chartaceous and dark brown, subglabrous 
above and below, venation pinnate, 2 veins 4-67 
side, glands usually present at the basal vein axils 
beneath. Inflorescences terminal, ca. 23 cm long, 
panicles with well-separated opposite lateral 
branches to 4 cm long, peduncle 4-9 cm long, 2- 
3 mm diam., glabrous, drying dark, flowers in dis- 
tal groups of 2 or 3, pedicels 5-14 mm long. 
Flowers with calyx 4-7 mm long, cupular with 5 
prominent (0.5-1.2 mm) teeth or entire, subgla- 
brous but with rounded discoid flat glands; co- 
rolla 40-55 mm long, tubular-funnelform, lav- 
ender with a white throat, tube 10-15 mm diam. 
at mouth, glabrous or appressed puberulent exter- 
nally, lobes to 20 mm long; filaments ca. 20 and 
13 mm long, thecae 3-4 mm long; ovules 4-se- 
riate. Fruits 14-32 cm long, 3-4 cm wide, ob- 
long, valves flattened, margins rounded, midvein 
flat, surface wrinkled; seeds 15-21 mm long, 35- 
58 mm wide, brownish. 

Plants of lowland evergreen rain forest forma- 
tions of the Pacific slope, 10-100 m elevation. A 
collection from Reserva Biol6gica Carara, flow- 
ering in early February (Ziiniga 90), is the only 
record known from Central America. Otherwise, 
this species ranges from northern South America 
to Brazil and Bolivia. 

Cydista lilacina is recognized by its climbing 
habit, simple or bifoliolate leaves, terminal pani- 
cle, calyx with round flat glands, and oblong fruits 
with flattened wrinkled surface. The tendency to 
dry dark (especially stems and inflorescence 
axes), general lack of puberulence, and lowland 
wet forest habitat are additional characteristics. 

Cydista potosina (Schum. & Loes.) Loes., Re- 
pert. Spec. Nov. Regni Veg. 16: 209. 1919. Ar- 
rabidaea potosina Schum. & Loes., Bull. Herb. 
Boissier 3: 618. Clytostoma mayanum Si ami I.. 
Carnegie Inst. Wash. Publ. 461. 86. 1935. Fig- 
ure 19. 



BURGER: FLORA COSTARICENSIS 



123 



Lianas, tendrils to 15 cm long, slender, leafy 
stems 1.7-5 mm diam., glabrous or with minute 
peltate hairs, tetragonal with 4 prominent longi- 
tudinal ridges, interpetiolar ridge usually present; 
pseudostipules linear or triangular, 1-8 mm. 
Leaves 2-foliolate, petioles 12-37 mm long, 1- 
1.5 mm diam., sparsely puberulent, striate, petio- 
lules 9-40 mm long; leaflet blades 4.5-15 cm 
long, 2-8 cm wide, ovate to ovate-elliptic or 
ovate-oblonge, apex acute to acuminate, base ob- 
tuse or rounded, drying chartaceous and brown, 
upper surface glabrous, lower surface glabrous or 
with short (0.2-0.4 mm) thin hairs along the 
veins, venation pinnate, 2 veins 3-6/side. Inflo- 
rescences axillary or terminal, 2-10 cm long, ra- 
cemes with 2-9 flowers, peduncles 5-85 mm 
long, glabrous or with minute (0.05 mm) peltate 
hairs, pedicels 5-18 mm long. Flowers with calyx 
5-7 mm long, 4-6 mm diam., narrow-cupular, 
drying dark, subglabrous, sometimes with round- 
ed flat glands distally, margin entire with 5 tri- 
angular minute lobes or irregular with pale edge; 
corolla 43-57 mm long, funnelform-campanulate, 
lavender or white, with minute glandular hairs ex- 
ternally, tube 1 1-16 mm diam. at the mouth, lobes 
9-16 mm long, broadly rounded; filaments ca. 17 
and 20 mm long, thecae ca. 3 mm long. Fruits 
17-24 cm long, 27-43 mm wide, narrowly ob- 
long, narrowed at the apex, valves flat with the 
margins elevated, midvein flat or elevated, surface 
smooth and dark; seeds 14-23 mm long, 45-70 
mm wide, brownish but transparent at the wing 
tips. 

Plants of wet lowland evergreen and partly de- 
ciduous forest formations on both Caribbean and 
Pacific slopes, 5-400 m elevation (to 800 m in 
northern Central America). Flowering in May- 
August. This species ranges from Veracruz, Mex- 
ico, to Costa Rica but is rarely collected along the 
Pacific slope of Central America. 

Cydista potosina is recognized by its climbing 
habit with simple tendrils, bifoliolate leaves, 
short, few-flowered racemes, lavender or white 
corollas, and narrowly oblong fruits with winged 
seeds. The lack of gland fields in the axils of basal 
veins (lower leaf surface), consistently pinnate ve- 
nation, and elevated ridges on leafy stems help 
distinguish this species from some of its conge- 
ners. 

Dendrosicus species are now placed in Amphi- 
tecna. 



Distictella Kuntze 

Lianas (shrubs), climbing with distally coiled 
and trifid tendrils, stems with 4 phloem areas in 
cross-section, branchlets terete, nodes lacking in- 
terpetiolar lines and glandular fields; pseudostip- 
ules short and inconspicuous. Leaves opposite, 
petiolate, 3-foliolate or 2-foliolate with terminal 
tendril, petiolules well developed, margins entire, 
venation pinnate, domatia absent, gland fields of- 
ten present in the basal vein axils beneath. Inflo- 
rescences terminal or axillary, racemes or race- 
mose panicles with short lateral branches, rachis 
minutely puberulent, bracts small and caducous. 
Flowers with cupulate calyx, minutely puberu- 
lent, margin truncated and subentire, glandular 
fields often present near the edge; corolla tubular- 
campanulate, white, minutely and densely puber- 
ulent externally, lobes 5; stamens 4, of 2 lengths, 
anthers glabrous, thecae straight and divaricate, a 
staminode present; disc annular; ovary 2-locular, 
ovules in 4-8 series on each placenta, stigma sim- 
ple. Fruits laterally compressed or biconvex cap- 
sules, the 2 valves flattened parallel to the septum, 
thick and woody; seeds flat, woody or with thin 
wings on 2 lateral sides and distally, usually 
brown. 

Distictella is a genus of 13 South American 
species and one species that ranges from Costa 
Rica to Brazil. 

Distictella magnoliifolia (Kunth in H.B.K.) 
Sandw., Lilloa 3: 460. 1938. Bignonia magno- 
liaefolia Kunth in H.B.K., Nov. Gen. Sp. 3: 
136. 1819. Figure 21. 

Lianas to 30 m high and 8 cm diam., tendrils 
to 18 cm long, 0.6-2.3 mm diam., leafy stems 3- 
8 mm diam., terete, minutely (0.05-0.1 mm) pu- 
berulent with reddish brown hairs. Leaves 2-fo- 
liolate, petioles 8-55 mm long, 1.3-2.7 diam., 
petiolules 6-23 mm long, often thickened below 
the blade, minutely puberulent; leaf blades 8-27 
cm long, 4-12 cm wide, elliptic to elliptic-oblong 
or ovate-elliptic, apex short-acuminate, base ob- 
tuse to cuneate, drying chartaceous, glabrous 
above, minutely puberulent on the veins beneath, 
2 veins 5-8/side, glands often present in basal 
vein axils. Inflorescences terminal or axillary, 
10-28 cm long, racemose panicles, rachis densely 
minutely puberulent, brownish, 2 peduncles (lat- 
eral branches) 6-15 mm long, 1.5-2 mm diam., 
bracts 2-4 mm long, pedicels 4-8 mm long. 
Flowers with calyx 9-13 mm long, 8-11 diam., 



124 



FIELDIANA: BOTANY 



cupular with entire margin, minutely papillate pu- 
berulent, longitudinal gland fields 2-3 mm long 
often present distally; corolla 42-65(-73) mm 
long, 30-45 mm wide at the lobes, white with 
yellow throat, densely minutely glandular puber- 
ulent externally, tube 10-17 mm diam. above the 
narrowed (3 mm) basal portion, lobes 10-18 mm 
long, rounded distally and narrowed near the base; 
filaments ca. 22 and 16 mm long. Fruits 7.5-23 
cm long, 32-55 mm wide, 15-20 mm thick, nar- 
rowly oblong to elliptic-oblong, valves woody, 
surface smooth; seeds 19-24 mm long, 28-38 
mm wide, oblong to suborbicular, brown, body of 
seed 14-17 mm wide, not differentiated. 

Rarely collected plants of lowland Caribbean 
evergreen rain forest formations ca. 100 m ele- 
vation, fruiting in November. The species ranges 
from northeastern Costa Rica (Morales et al. 3212 
from La Selva) to Peru and Brazil. 

Distictella magnoliifolia is recognized by its 
climbing habit with distally trifid tendrils, oppo- 
site bifoliolate leaves, terminal raceme-like inflo- 
rescences, minutely puberulent calyx cups with 
entire margins and distinct glandular areas, large 
white corollas, woody oblong fruit, and flat brown 
seeds in which the opaque wings extend around 
three sides of the central area. The above descrip- 
tion is based on South American material. 

Gibsoniothamnus is now placed in the Schle- 
geliaceae (q.v.). 



Godmania Hemsley 

Small to medium-size trees, bark smooth to 
longitudinally ridged, interpetiolar line and glan- 
dular field absent at the nodes. Leaves opposite, 
long-petiolate, palmately 5-9 foliolate, basal leaf- 
lets smaller than distal, leaflets petiolulate, mar- 
gins entire (serrate on juvenile shoots), venation 
pinnate. Inflorescences terminal, compact many- 
branched corymbose panicles with many flowers, 
bracts small, puberulent, pedicels well developed. 
Flowers with small (< 2 mm) broadly campanu- 
late calyx, subentire or with 5 small lobes; corolla 
urceolate-campanulate and slightly 2-lipped, yel- 
low and brownish, puberulent externally, lower 
lip 3-lobed, lobes valvate in bud and triangular; 
stamens 4, of 2 lengths, included, filaments and 
anthers pubescent, thecae divaricate, a staminode 
present; disc annular-pulvinate; ovary minutely 
puberulent, 2-locular, ovules multiseriate in each 
locule. Fruits linear-cylindric capsules, twisted, 



dehiscing loculicidally (perpendicular to the sep- 
tum), valves coriaceous; seeds with 2 lateral 
membranaceous wings. 

Godmania is a Neotropical genus of two spe- 
cies characterized by the smaller yellowish flow- 
ers with valvate lobes, twisted linear fruits, and 
palmately compound opposite leaves. A second 
species occurs in the caaiinga formations of Bra- 
zil. The genus is a member of tribe Tecomeae and 
related to Tabebuia (Gentry, 1992). 

Godmania aesculifolia (Kunth in H.B.K.) Stand). 
in Standl. & Calder6n, Lista Prelim. PI. El Sal- 
vador 200. 1925. Bignonia aesculifolia Kunth 
HI H.B.K., Nov. Gen. Sp. 3: 140. 1819. Tecoma 
fuscata Mocino ex DC., Prodr. 9: 221. 1845. 
Cybislax macrocarpa Benth. in Benth. & 
Hook., Gen. PI. 2: 1043. 1876. G. macrocarpa 
(Benth.) Hemsl., Diag. PI. Nov. Mex. 2: 35. 
1879. Figure 12. 

Trees or shrubs, 3-7(-20) m tall, leafy stems 
3-8 mm diam., minutely (0.1-0.2 mm) puberu- 
lent, brown, terete, becoming grayish and lenti- 
cellate. Leaves 1 1-38 cm long, with 5, 7, or 9 
leaflets, petioles 4.5-18 cm long, 2-3 mm diam., 
petiolules 2-24 mm long, usually densely puber- 
ulent; leaflet blades 2.5-16 cm long, 1.5-7 cm 
wide, elliptic-obovate, obovate, narrowly ellip- 
tic-oblong or oblanceolate, apex short caudate- 
acuminate to gradually acuminate, margin entire 
(with teeth on juvenile shoots), base acute to cu- 
neate, drying stiffly chartaceous, sparsely to 
densely puberulent with thin straight hairs 0.1- 
0.3 mm long beneath, punctate on both surfaces, 
2 veins 6-13/side, loop-connected distally. In- 
florescences terminal, 5-10 cm long, 6-10 cm 
wide, often a flat-topped panicle, 1 peduncle 2- 
9 mm long, 2 peduncles 12-15 mm long, dense- 
ly puberulent, bracts ca. 0.7 mm long, pedicels 
2-9 mm long, ca. 0.5 mm diam. Flowers with 
calyx 1-2 mm long, 2-3 mm wide, broadly ob- 
conic or campanulate, minutely puberulent, sub- 
entire or 5-lobed (lobes ca. 0.2 mm and obscure); 
corolla 12-16 mm long, urceolate to campanu- 
late and slightly 2-lipped, yellow with red-brown 
marking on the upper lobes, tube 5-7 mm diam. 
for most of its length, densely minutely puberu- 
lent externally, lobes 1-5 mm long; filaments ca. 
9 and 7 mm long, thecae less than 1 mm long; 
style ca. 9 mm long. Fruits 28-60(-100) cm 
long, 10-14 mm wide, 8-14 mm thick, long-lin- 
ear, somewhat flattened and becoming coiled or 
twisted, with ca. 6 prominent longitudinal ridg- 



BURGER: FLORA COSTARICENSIS 



125 



es on each face; seeds 7-11 mm long, 50-135 
mm wide, central area well differentiated, 6-8 
mm long, 10-12 mm wide, wings translucent. 

Infrequent deciduous trees of deciduous and 
partly deciduous forests of the Meseta Central 
and Pacific slope, 5-1200 m elevation. Flower- 
ing in February-July; fruiting in November- 
March. This species ranges from Mexico to Bo- 
livia. 

Godmania aesculifolia is recognized by the 
tree habit, opposite palmately compound leaves 
with three to nine leaflets, the short, many- 
branched terminal inflorescences with small 
crowded flowers, short calyx, and campanulate 
or urceolate yellow corolla. The long linear spi- 
rally coiling fruits and seeds with clearly differ- 
entiated translucent membranaceous lateral 
wings are additional characteristics. The branch- 
es often have an unpleasant odor when broken. 



Jacaranda Jussieu 

Trees (shrubs in Brazil), stems quadrangular or 
terete, nodes without gland fields or interpetiolar 
lines; pseudostipules absent. Leaves opposite, 
usually bipinnate (pinnate to simple in Brazilian 
spp.), often large, petiolate, lateral pinnae oppo- 
site or alternate, leaflets sessile or subsessile, often 
strongly asymmetric with 1 side broader than the 
other, margin entire or with distal teeth, base usu- 
ally asymmetric, glabrous or puberulent, venation 
pinnate. Inflorescences terminal, axillary or from 
older leafless nodes, panicles or thyrses (racemes) 
with many flowers, pedunculate, lateral branches 



usually opposite and not crowded, pedicels sub- 
tended by enlarged ebracteate nodes. Flowers 
showy, calyx broadly cupulate to tubular-campan- 
ulate, short, 5-lobed to denticulate or subentire; 
corolla tubular-campanulate or campanulate-fun- 
nelform above a narrowed base, slightly bilabiate, 
blue or lavender to magenta (white), densely pu- 
berulent to glabrous externally, the tube contract- 
ed near the base; stamens 4, filaments of 2 
lengths, anthers glabrous, unequal or with only 1 
theca, staminode usually exceeding the stamens 
with glandular hairs along its length and at the 
apex, pollen 3-colpate; disc pulviniform, ovary 
glabrous or puberulent, 2-locular, ovules on ca. 8 
series in each locule. Fruits capsules, oblong or 
rounded in outline, strongly flattened perpendic- 
ular to the septum and dehiscing parallel to the 
plane of compression, surfaces glabrous or lepi- 
dote; seeds thin, flat, with translucent or brown 
circumferential wings (wider in width than in 
length), seed area and membranaceous wing dif- 
ferentiated or not. 

Jacaranda is a genus of 49 species ranging 
from Mexico to Argentina (Gentry & Morawetz 
in Gentry, 1992), belonging to the tribe Teco- 
meae. The beauty of the trees when in flower has 
made a few of the species ornamental favorites, 
now planted throughout the tropics and subtrop- 
ics. The genus stands out within the family be- 
cause of its tree habit, opposite bipinnate leaves 
with usually many small leaflets, bluish or lav- 
ender flowers, and flat, rounded woody fruits. The 
long staminode, often adorned with glandular 
hairs, is another important generic distinction (it 
can be mistaken for a style). 



Key to the Species of Jacaranda 

la. Calyx 4-7 mm long and tubular; leaflets 20-80 mm long, 6-25 mm wide [corolla densely puberulent 
externally; fruits with flat rounded margin; trees to 45 m tall; wet forests of both the Caribbean and 
Pacific slopes] J. copaia 

Ib. Calyx 1-3 mm long and cupular or saucer-like; leaflets 6-24 mm long, 1-10 mm wide 2 

2a. Corolla mostly glabrous externally; leaflets 3.5-10 mm wide; fruits 5-12 cm long, oblong or rounded 
with strongly undulating lateral margins; growing wild in evergreen forests of the Pacific slope . . 
J. caucana 

2b. Corolla minutely puberulent externally; leaflets 1-4 mm wide; fruits 4-7 cm long, suborbicular 
with flattened or slightly undulating margins, widely planted for ornament in evergreen and partly 
deciduous areas J. mimosifolia 



126 



FIELDIANA: BOTANY 



Jacaranda caucana Pittier, Contrib. U.S. Natl. 
Herb. 18: 258. 1917. J. ficifolia D. Don, sec 
Seem., Bot. Voy. Herald 181. 154, non Don. J. 
trianae Kranzl., Fedde Repert. 17: 226. 1921. 
J. caucana ssp. sandwithiana A. Gentry, Ann. 
Missouri Bot. Card. 60: 858. 1973 (1974). Fig- 
ure 14. 

Trees 8-28 m tall, trunks up to 1 m diam., 
leafy stems 3-12 mm diam., glabrous or with few 
minute peltate hairs, young stems drying dark 
with narrow whitish lenticels, older stems gray. 
Leaves bipinnate, 20-45 cm long, petioles 4-8 
cm long, 1.8-4 mm diam., sparsely minutely pu- 
berulent, rachis sulcate above, with lateral wings 
ca. 0.5 mm wide, lateral pinnae 7-18 pairs, 4-17 
cm long, petiolules 3-10 mm long; leaflet blades 
8-28 mm long, 3.5-10 mm wide (terminal leaflets 
to 37 X 16 mm), asymmetrically oblong-rhombic, 
apex obtuse with a small apiculate tip, base with 
narrowly cuneate and broadly obtuse sides, drying 
chartaceous, glabrous above with major veins im- 
pressed, grayish beneath with thin hairs 0.2-0.4 
mm long 2 veins 4-7/side. Inflorescences usu- 
ally borne at distal leafless nodes, 6-15 cm long, 
paniculate with 3-30 flowers, peduncles 3-25 mm 
long, 0.7-2 mm diam., glabrous or sparsely pu- 
berulent, drying dark, pedicels 0.5-3 mm long 
(ca. 6 mm including subtending node). Flowers 
with calyx ca. 1 .5-3 mm long, 1 .8-4 mm wide at 
the apex, broadly campanulate, sparsely papillate 
puberulent, bluntly 5-toothed; corolla 35-48 mm 
long, tubular-campanulate and often bent almost 
90 near the base, purple-blue, subglabrous exter- 
nally, tube 10-17 mm diam. near the mouth, lobes 
ca. 10 X 12 mm; filaments ca. 14 and 13 mm 
long, thecae 1.5-2 mm long, staminode 22-25 
mm long with hairs to 1 mm long. Fruits 6-12 
cm long, 4-7 cm wide, oblong-elliptic (narrowed 
at base and apex), 10-17 mm thick, margins thin- 
ner and undulating (2 lobes/side); seeds 8-19 mm 
long, 23-42 mm wide, wings brown or slightly 
translucent, not clearly differentiated. 

Trees of evergreen forest formations of the 
southern Pacific slope (in Costa Rica), 5-1000 m 
elevation. Flowering in March-May; fruiting in 
November-March. The species ranges from 
southwestern Costa Rica to Venezuela. 

Jacaranda caucana is recognized by its tree 
habit, opposite bipinnate leaves with small asym- 
metric leaflets, flowers with small, widely flaring 
calyx cups, purple-blue corollas mostly glabrous 
on the exterior, and the flat woody fruits with un- 
dulating margin. Costa Rican material is placed in 



subspecies sandwithiana, having shorter or un- 
developed calyx lobes and a corolla tube that is 
glandular-pilose externally. Four subspecies were 
recognized by Gentry and Morawetz (Gentry, 
1992). 

Jacaranda copaia (Aublet) D. Don, Edinburg 
Phil. J. 9: 267. 1823. Bignonia copaia Aublet. 
Hist. PI. Guiane Fr. 2: 650. tab. 262, fig. 1, tab. 
265. 1775. J. spectabilis Mart, ex DC., Prodr. 
9: 229. 1845. J. superba Pittier. Bol. Soc. Ve- 
nez. Ci. Nat. 6: 19. 1940. J. copaia ssp. spec- 
tabilis (Mart, ex DC.) A. Gentry, Rhodora 79: 
441. 1977. Figure 14. 

Trees to 45 m tall, trunks to ca. 50 cm diam., 
with few upright branches forming a plume-like 
large-leaved crown when young, leafy stems 4- 

12 mm diam., glabrous, subtetragonal, lenticel- 
late. Leaves 30-160 cm long, to 60 cm wide, pet- 
ioles 4-29 cm long, 2.5-10 mm diam., minutely 
papillate puberulent or muricate, terete, rachis 
narrowly sulcate above (without wings), pinnae 
7-35 cm long, petiolules 0-3 mm long; leaflet 
blades 20-80 mm long, 6-18(-25) mm wide, 
asymmetrically rhomboid-elliptic to elliptic-ob- 
long, apex acuminate, base asymmetric with nar- 
rowly cuneate and obtuse sides, drying dark and 
chartaceous, upper surface with minute hairs on 
the midvein, sparsely minutely puberulent on the 
veins beneath, 2 veins 3-7/side. Inflorescences 
to 48 cm long, open pyramidal panicles with lat- 
eral branches to 30 cm long (sometimes with 3 
terminal thyrses), peduncles to 8 cm long, 3-5 
mm diam., minutely puberulent or with peltate 
hairs. Flowers with calyx tube 4-7 mm long, 3- 
5 mm diam, densely minutely (0.1 mm) puberu- 
lent, irregularly lobed (0.3 mm) or subentire; co- 
rolla 23-50 mm long, tubular-campanulate above 
the narrowed (8-10 mm) base, blue or lavender 
(white in the throat), densely minutely puberulent 
externally, tube 8-14 mm diam. distally, lobes 3- 

13 mm long; filaments ca. 12 and 9 mm long, 
thecae ca. 2 mm long, staminode 24-27 mm long, 
pubescent, bifurcate at tip. Fruits 4.5-12 cm long, 
3-6 cm wide, 5-8 mm thick, oblong-rounded, sur- 
face smooth but minutely muricate; seeds 11-14 
mm long, 21-30 mm wide, seed ca. 5 mm diam., 
wings translucent with brown rays. 

Trees of rain forests on both Caribbean and Pa- 
cific slopes, 10-600 m elevation. Flowering in 
February- April; fruiting in July-November. This 
species ranges from southern Mexico to Brazil 
and Bolivia. 



BURGER: FLORA COSTARICENSIS 



127 



Jacaranda copaia is distinguished by the tall 
tree habit, opposite bipinnate leaves, strongly 
asymmetric leaflets, tubular calyx, bluish lavender 
corolla densely puberulent externally, flat oblong 
woody fruit, and seeds with brown rays in the 
otherwise transparent wings. When mature, this is 
one of the most beautiful trees of southern Central 
America (Standley, 1938). As a young tree, this 
species has a slender unbranched trunk with as- 
cending distal branches forming a terminal tuft 
with very large bipinnate leaves. The mimosoid 
legume Schizolobium parahybum (Veil.) Blake is 
quite similar in this regard; both species are vi- 
sually striking and often seen in lowland rain for- 
est secondary growth (Gentry, 1973b). Gallinazo 
is a common name. 

Jacaranda mimosifolia D. Don, Bot. Reg. 8: tab. 
631. 1822. J. ovalifolia R. Br., Bot. Mag. tab. 
2327. 1822. Figure 14. 

Trees, usually small to medium size (8-15 m 
tall), stems 4-9 mm diam., usually glabrous, te- 
rete, with elongate lenticels. Leaves 13-34 cm 
long, petioles to 4 cm long, 1.8-2.6 mm diam., 
lateral pinnae 3-10 cm long, with narrow wings 
0.3-0.5 mm wide, with 13-41 opposite or alter- 
nate, subsessile leaflets; leaflet blades 5-14 mm 
long, 1-4 mm wide (terminal leaflets to 22 X 6 
mm), elliptic to elliptic-obovate, apex acute to 
acuminate, slightly asymmetric at the obtuse/cu- 
neate base, glabrous above, glabrous or puberu- 
lent along the edge and major veins beneath, 2 
veins 3-7/side. Inflorescences terminal, 9-38 cm 
long, open panicles with well-separated opposite 
branches, peduncles 34-88 mm long, 1-4 mm 
diam., pedicels 1-2 mm long (to subtending 
node), flowers usually in distal cymes. Flowers 
with calyx cup 1-2 mm long, ca. 2 mm wide, 
sparsely minutely puberulent, margin with 5 tri- 
angular or acute teeth 0.2-1 mm long; corolla 
30-45 mm long, tubular-campanulate, bluish lav- 
ender or lavender, densely puberulent near the 
base with hairs 0.1-0.3 mm long, less densely pu- 
berulent distally, tube 9-13 mm diam. near the 
mouth, lobes 3-8 mm long; filaments ca. 16 and 
13 mm long, thecae ca. 2 mm long, staminode 
20-25 mm long. Fruits 30-70 mm long, 37-60 
mm wide, 12-20 mm diam., orbicular to oblong- 
rounded, lenticular with thinner flattened edges, 
surface smooth and slightly muricate; seeds 9-13 
mm long, 11-19 mm wide, seed ca. 7 X 6 mm, 
clearly differentiated from the transparent circum- 
ferential wing. 

Jacaranda mimosifolia, native to northern Ar- 



gentina and adjacent Bolivia, is widely planted 
throughout the tropics and subtropics as an orna- 
mental tree. In Central America it is usually 
grown between 500 and 1400 m elevation, flow- 
ering in January-July, with peak flowering in 
April-May. Standley and Williams (1974) de- 
scribed the colorful effect of these showy trees in 
Guatemala. The very small leaflets on opposite 
bipinnate leaves, bluish lavender flowers bloom- 
ing in the dry season, and hard, rounded, lens- 
shaped fruits help distinguish this species. Jaca- 
randa and Jacaranda are the Spanish and English 
names. 



Kigelia DeCandolle 

Medium-size trees up to 20 m tall, with broad 
spreading crowns and inflorescences hanging 
from the lower branches. Leaves opposite, peti- 
olate, imparipinnate with 3-1 1 leaflets, the ter- 
minal leaflet larger than the laterals and borne on 
a longer petiolule, margin entire to dentate, ve- 
nation pinnate. Inflorescences panicles with few 
short lateral branches from an elongate central ra- 
chis, long-pedunculate and pendulous, bracts 
small, pedicels well developed. Flowers large, ca- 
lyx campanulate or cupular, splitting into 2-5 
lobes, coriaceous, usually glabrous; corolla cam- 
panulate-funnelform and distally curved, glabrous 
on the exterior, 5-lobed and somewhat 2-lipped, 
deep purple to dark wine-red (orange-red); sta- 
mens 4, of 2 lengths, subexserted at the mouth of 
the corolla, thecae slightly divergent; disc annular; 
ovary lepidote, 1-locular with 2 parietal placentas 
or bilocular in the lower half, style short, stigma 
simple. Fruits indehiscent, pendulous, cylindric- 
oblong, fibrous-woody; seeds many, without 
wings. 

Kigelia is now believed to comprise a single 
very variable species, native to savannas and for- 
ests in central tropical Africa. (In the past some 
authors recognized as many as ten species.) These 
trees are occasionally grown as unusual exotics in 
tropical parks and gardens. 

Kigelia pinnata (Jacq.) DC., Rev. Bign. (Bibl. 
Univ. Geneve) 24. 1838. Crescentia pinnata 
Jacq., Collect. 3: 203, tab. 18. 1791. K. africana 
DC., Fl. Nigrit. 463. 1849. 

Trees 6-12 m (rarely 20 m) tall, twigs often 
hollow, leafy stems 6-14 mm diam., glabrous or 
minutely puberulent, glandular fields absent at the 
nodes. Leaves to 50 cm long, with usually 7 or 9 



128 



FIELDIANA: BOTANY 



leaflets, petioles 6-14 cm long, ca. 3 cm diam., 
petiolules of lateral leaflets 1-4 mm long; leaf 
blades (2-)5-16 cm long, (1.5-)3-7 cm wide, el- 
liptic-oblong to elliptic-obovate, apex obtuse to 
short-acuminate, margin entire or dentate distally, 
base asymmetric in lateral leaflets (rounded/cu- 
neate), drying stiffly chartaceous, glabrous or with 
minute (0.1 mm) straight hairs, 2 veins 7-1 I/side. 
Inflorescences terminal and pendulous, to 40 cm 
long, peduncles ca. 20 cm long, 3 mm diam., with 
lateral branches to 9 cm long and 2.5 mm diam., 
pedicels 10-14 mm long. Flowers held horizon- 
tally, calyx 2-3 cm long, 12-20 mm wide at the 
apex, truncated at the base, lobes to 12 mm long, 
coriaceous; corolla 6-10 cm long, tube ca. 14 mm 
diam. at base, narrowing to 1 1 mm and then open- 
ing to 4-6 cm wide, dark maroon or wine-red; 
anthers ca. 8 mm long; ovary 10-11 mm long, 3 
mm diam. Fruits 25-50(-80) cm long, 5-12 cm 
diam., tubular-oblong, pendulous, glabrous on the 
exterior. 

Kigelia pinnata is an African tree rarely en- 
countered in Neotropical parks and gardens (our 
only Costa Rican collections come from Golfito). 
The pendulous raceme-like inflorescences with 
large dark wine-red or maroon flowers held hor- 
izontally and the large pendulous sausage-like 
fruits make this a unique species. These plants are 
adapted for bat pollination, with inflorescences 
long-pendulous beneath the lower branches, and 
night-blooming flowers with thick tissues and foul 
odor. 



Lundia DeCandolle 
Nomen conservandum 

Lianas climbing with tendrils (simple or trifid 
near the tip), stems with 4 phloem areas in cross- 



section, branchlets terete, nodes often wider than 
the stems, with interpetiolar gland fields; pseu- 
dostipules small or absent. Leaves opposite, 2- or 
3-foliolate. petiolate, blades usually somewhat 
rounded near the base, margin entire, venation 
palmate or subpalmate, tufts of hairs (domatia) of- 
ten present in the proximal vein axils beneath. In- 
florescences axillary or terminal, usually less than 
20 cm long, paniculate, pedunculate, bracts mi- 
nute or absent, flowers pedicellate. Flowers with 
calyptrate-conical calyx buds, calyx margin entire 
or splitting, puberulent; corolla tubular-funnel- 
form, white to rose or magenta, puberulent exter- 
nally; stamens 4, included, filaments of 2 lengths, 
anthers puberulent, thecae divaricate and straight, 
staminode small; disc absent; ovary densely pu- 
berulent, 2-locular, ovules many in 2-6 series in 
each locule. style puberulent or glabrous. Fruits 
capsules, linear and smooth, valves flattened par- 
allel to the septum, surface densely puberulent, 
median vein and margins usually elevated, de- 
hiscing septicidally; seeds flat, thin, transverse-ob- 
long, with 2 lateral membranaceous wings trans- 
parent at the tips. 

Lundia is a genus of 12 species ranging from 
Mexico to Brazil and Bolivia. The genus is dis- 
tinguished by its unusual calyx, externally puber- 
ulent white to rose or purple corollas, stamens 
with divaricate puberulent thecae (often in a 
straight line), and long narrow fruits with densely 
puberulent surface. The early flower buds are tu- 
bular with narrowly conical tip that is deciduous 
to produce a calyx with an entire margin, which 
then may split as growth continues. The calyx re- 
mains obconic at the base as opposed to the 
rounded base in Paragonia pyramidata, which 
may also have calyptrate calyces. Costa Rican 
species of Lundia are also vegetatively similar to 
species of Arrabidaea and Cydista. 



Key to the Species of Lundia 

la. Flowers white; style glabrous except at the base; calyx drying yellowish, entire or sometimes split; 

occasional plants of evergreen and partly deciduous forests L corymbifera 

Ib. Flowers pink or magenta; style puberulent near the tip or along its length; calyx drying brownish 

and often split to form 2 lips; rarely collected in evergreen forests L. puberula 



Lundia corymbifera (Vahl) Sandw., Recueil 
Trav. Bot. Ne~erl. 34: 229. 1937. Bignonia cor- 
ymbifera Vahl, Eclog. Am. 2: 45, pi. 17. 1798. 
B. umbrosa Kunth in H.B.K., Nov. Gen. Sp. 3: 
138. 1819. L. valenzuelae Dugand, Mutisia 10: 
7. 1952. Figure 23. 



Lianas to 3 cm diam., tendrils 4-14 cm long, 
leafy stems 1 .7-6 mm diam., minutely puberulent 
with thin hairs 0.1-0.2 mm long, longitudinally 
ridged (striate), gland fields prominent; pseudo- 
stipules conical. Leaves 2- or 3-foliolate, petioles 
8-64 mm long, 0.8-1.7 mm diam., minutely pu- 



BURGER: FLORA COSTARICENSIS 



129 



berulent, petiolules 7-42 mm long; leaflet blades 
5-14 cm long, 2-9 cm wide, ovate-elliptic to 
broadly ovate, apex caudate-acuminate to long- 
acuminate (tips ca. 15 mm long), base rounded 
and truncate to subcordate, slightly or clearly 
asymmetric, drying thin-chartaceous and brown, 
minutely puberulent on the midvein above, with 
whitish hairs 0.1-0.3 mm long beneath and larger 
hairs sometimes present in the vein axils (doma- 
tia), venation palmate or subpalmate, 2 veins 3- 
7/side. Inflorescences axillary or terminal, 5-10 
cm long, peduncles 3-4.5 cm long, ca. 1 mm 
diam., densely puberulent with short (0.1-0.2 
mm) yellowish hairs, pedicels 5-8 mm long. 
Flowers with calyx 3-6 mm long, 3-4 mm diam., 
cupulate-campanulate to obconic, margin entire or 
split on 1 or 2 sides, minutely puberulent; corolla 
26-43 mm long, tubular-funnelform, white, 
densely minutely puberulent externally, with yel- 
low ridges in the throat, lobes 4-12 mm long; 
filaments ca. 14 and 9 mm long, thecae 2-3 mm 
long, divaricate. Fruits 28-60 cm long, 15-20 
mm wide, valves thick, with raised midrib and 
edges, densely minutely velutinous; seeds 6-14 
mm long, 19-40 mm wide, central area ca. 10 
mm wide, lateral wings with darker interior and 
clear distal areas. 

Vines often found along stream edges in ever- 
green and partly deciduous forest formations of 
the Pacific slope, 10900 m elevation. Flowering 
in June-December; fruiting in February-March. 
This species ranges from central Costa Rica to 
Brazil and Bolivia. 

Lundia corymbifera is distinguished by its vin- 
ing habit with simple tendrils, opposite bi- or tri- 
foliolate leaves, leaflets rounded at the base, short 
inflorescences, unusual calyx buds, pubescent an- 
thers, white corolla puberulent on the exterior, and 
long linear fruits with two-winged seeds. The 
flower buds are ellipsoid and often have an apic- 
ulate apex that usually dehisces as a cap, leaving 
an entire margin. 

Lundia puberula Pittier, Contrib. U.S. Natl. 
Herb. 18: 258. 15 Sept. 1917. L dicheilocalyx 
Blake, Contrib. Gray Herb. 52: 94. 28 Sept. 
1917. L. schumanniana Kranzl., Fedde Repert. 
17: 120. 1921. L. colombiana Dugand, Caldasia 
4: 236. 1946. 

Lianas to 5 cm diam., tendrils 5-15 cm long, 
leafy stems 1.6-6 mm diam., minutely (0.1 mm) 
puberulent, longitudinally ridged or striate, pseu- 
dostipules inconspicuous. Leaves 2- or 3-folio- 



late, petioles 2.4-6 cm long, petiolules 10-20 mm 
long, 0.6-1 mm diam., minutely puberulent; leaf 
blades 6-12(-16) cm long, 3.5-7(-9) cm wide, 
ovate to ovate-oblong, apex long-acuminate (tips 
ca. 2 cm long), base rounded and truncate to cor- 
date, slightly asymmetric, drying thin-chartaceous 
and brown, sparsely puberulent above, minutely 
puberulent beneath and often with longer (0.2-0.6 
mm) hairs in the vein axils (domatia), venation 
palmate or subpalmate, 2 veins 3-5/side. Inflo- 
rescences axillary or terminal, 3-12 cm long, pe- 
duncles 15-55 mm long, 1-2 mm diam., minutely 
puberulent, bracts ca. 1 mm long, pedicels 38 
mm long, ca. 0.6 mm diam. Flowers with calyx 
4-8 mm long, 3-4 mm diam, tubular, entire or 
split on the margin, minutely puberulent with 
short (0.1 mm) hairs; corolla 30-58 mm long, 
tubular-campanulate or funnelform, slightly 2- 
lipped, pale pink to magenta, densely minutely 
puberulent externally, lobes 8-21 mm long; fila- 
ments 14-20 and 9-13 mm long, thecae 2-3 mm 
long. Fruits long linear capsules, 23-48 cm long, 
15-17 mm wide, valves flat with raised midvein, 
surface velutinous; seeds 9-12 mm long, 33-35 
mm wide, wings brown to transparent distally. 

Rarely collected vines in lowland evergreen 
rain forests of the Caribbean and Pacific slopes, 
0-900 m elevation. Flowering throughout the 
year. This species ranges from Mexico and Belize 
to Peru. 

Lundia puberula is recognized by its vining 
habit with simple tendrils, the opposite bi- or tri- 
foliolate leaves with long-acuminate tips and 
rounded base, the entire or split calyx, pubescent 
anthers, pink to magenta corollas minutely puber- 
ulent externally, and long slender fruits. The usual 
presence of tufted hairs in vein axils, gland fields 
at nodes, and lowland rain forest habitat are ad- 
ditional distinctions. This species is easy to con- 
fuse with some species of Arrabidaea and Cydis- 
ta. 



Macfadyena A. DeCandolle 

Lianas, tendrils with hardened curved trifid 
tips, stems terete, cross-section with ca. 8 phloem 
areas and fissured xylem, roots with swollen tu- 
bers, interpetiolar gland fields present or absent; 
pseudostipules small, lanceolate to ovate. Leaves 
evergreen or deciduous, opposite, 2-foliolate, pet- 
iolate, often with a tendril, margin entire, venation 
pinnate. Inflorescences axillary or terminal on 
short-shoots, 1 or 2 or 3-15 on short cymes or 



130 



FIELDIANA: BOTANY 



reduced panicles, pedicels subtended by nodes. 
Flowers with tubular or campanulate calyx, lobed 
or variously split to spathe-like, usually thin-tex- 
tured; corolla tubular-campanulate or tubular-fun- 
nelform, 2-lipped, bright yellow, glabrous exter- 
nally, with 5 rounded lobes; stamens 4, filaments 
of 2 lengths, anthers glabrous, thecae straight, di- 
varicate, staminode present; disc annular-pulvi- 
nate, ovary with minute peltate hairs to puberulent 
or glabrate, locules 2, ovules in 2-4 series in each 
locule. Fruits elongate linear capsules, valves 
parallel to the septum, flat, smooth, midvein 
slightly raised; seeds flat, transversely oblong. 



central area not clearly differentiated from the 2 
thin lateral wings. 

Macfadyena is a genus of four species ranging 
from Mexico and the West Indies to Argentina. 
The plants are dimorphic, with the leaf blades of 
young plants usually smaller and more lanceolate 
than the blades of more mature stems. In addition, 
these plants are unusual in that they arc able to 
climb up tree trunks using their short, three- 
clawed tendrils as grappling hooks; advcntitous 
roots also aid in adhering to their support. The 
flowers can vary greatly in size, even on the same 
vine. 



Key to the Species of Macfadyena 

la. Calyx usually split on 1 side (spathe-like) with apical tip curved backward; seeds 20-25 mm wide, 
brown throughout, fruits to 30 cm long; pseudostipules subulate-lanceolate; plants of lowland 
swamps below 100 m elevation in evergreen forest formations M. uncata 

Ib. Calyx subentire or variously lobed or split (not spathe-like); seeds 40-65 mm wide, translucent near 
the tips, fruits to 130 cm long; pseudostipules often ovate and striate; plants of well-drained sites 

in deciduous and partly deciduous or evergreen forest formations, 10-1200 m elevation 

M. unguis-cati 



Macfadyena uncata (Andr.) Sprague & Sandw., 
Recueil Trav. Bot. Neerl. 34: 215. 1937. Big- 
nonia uncata Andr., Bot. Repos. tab. 530. 1808. 
B. uncinata G. Meyer, Prim. Fl. Essequeb. 210. 
1818. M. uncinata (G. Meyer) A. DC., Prodr. 
180. 1845. M. guatemalensis Blake, Contrib. 
U.S. Natl. Herb. 24: 24. 1922. Figure 17. 

Lianas to 30 m high, tendrils with 3 stiff 
curved distal tips, leafy twigs 1-4 mm diam., mi- 
nutely puberulent or glabrous, terete, gland fields 
usually visible on young stems; pseudostipules ca. 
2 mm long, subulate-lanceolate. Leaves with pet- 
ioles 8-36 mm long, 1-1.7 mm diam., glabrous 
or sparsely puberulent with thin hairs ca. 0.2 mm 
long, petiolules 4-27 mm long; leaflet blades 5- 
20 cm long, 2-9 cm wide, narrowly ovate to ovate 
or ovate-elliptic, apex acuminate, often with a 
long (2 cm) narrow tip, base obtuse or cuneate 
(rounded), drying chartaceous and often dark, 
subglabrous or with very minute hairs above, mi- 
nutely puberulent and with appressed peltate hairs 
beneath, 2 veins 5-8/side. Inflorescences axil- 
lary to foliage leaves, flowers 1-3 in reduced 
cymes or panicles, peduncles 0-9 mm long, ca. 1 
mm diam., glabrous, drying black, bracts 1-4 mm 
long, lanceolate, caducous, pedicels ca. 10 mm 



long. Flowers with calyx 15-27 mm long, 6-14 
mm diam., glabrous or with few minute peltate 
hairs, usually split on 1 side (spathaceous) with 
the narrowed apex recurved; corolla 40-85 mm 
long, tubular-funnelform, yellow, glabrous exter- 
nally, 8-18 mm diam. at the mouth, lobes 7-17 
mm long; filaments ca. 23 and 18 mm long, the- 
cae 2.5 mm long. Fruits 16-30 cm long. 15-19 
mm wide, valves drying dark brown; seeds 12- 
14 mm long, 20-25 mm wide, transversely ob- 
long-rectangular with short lateral wings, dark 
brown throughout. 

Plants of swampy sites in evergreen lowland 
forest formations of both Caribbean and Pacific 
slopes, 1-110 m elevation. Probably flowering 
throughout the year. This species, often found in 
mangrove formations, ranges from Mexico to 
Brazil. 

Macfadyena uncata is recognized by its climb- 
ing habit with three-clawed tendrils, bifoliolate 
leaves with acuminate blades, few-flowered axil- 
lary inflorescences, spathe-like calyx, yellow co- 
rollas glabrous externally, and narrow fruits with 
short-winged seeds. It is also unusual in being re- 
stricted to swampy habitats at low elevations. 

Macfadyena unguis-cati (L.) A. Gentry, Britton- 
ia 25: 236. 1973. Bignonia unguis-cati L., Sp. 



BURGER: FLORA COSTARICENSIS 



131 



PI. 2: 623. 1753. Doxantha unguis-cati (L.) 
Miers emend. Rehder, Mitt. Deutsch, Dendrol. 
Gesel. 1913: 262. 1913. B. dasyonyx Blake, 
Contrib. Gray Herb. 52: 93. 1917. D. dasyonyx 
(Blake) Blake, J. Bot. 61: 192. 1923. Figure 17. 

Lianas to 30 m high, to 7 cm diam., tendrils 
2-5 cm long with 3 stiff curved terminal "claws" 
5-14 mm long, leafy stems 1.3-4 mm diam., gla- 
brescent and grayish, gland fields present or ab- 
sent at nodes; pseudostipules lanceolate to ovate, 
often with parallel longitudinal ridges (striate). 
Leaves dimorphic (young vines vs. mature 
stems), 2-foliolate, petioles 1 1-27(-47) mm long, 
0.8-1.5 mm diam., glabrous or sparsely and mi- 
nutely puberulent, petiolules 4-16(-25) mm long; 
leaf blades 5-16 cm long, 1-7 cm wide, narrowly 
ovate-elliptic to narrowly ovate or elliptic, apex 
acuminate with narrowed tip to 2 cm long, base 
rounded to acute, drying thinly chartaceous, most- 
ly glabrous except for minute puberulence on the 
midvein and with peltate hairs beneath, 2 veins 
4-8/side. Inflorescences terminal on short-shoots 
or axillary to leafless nodes, panicles of up to 15 
flowers, peduncles 8-28 mm long, usually gla- 
brous, pedicels 8-10 mm long. Flowers with ca- 
lyx 6-15 mm long, 7-12 mm wide, cupular to 
campanulate, thin-textured, distal margin irregu- 
larly split or with lobes 0.5-2 mm long, glabrous 
or with few peltate hairs; corolla 42-80 mm long, 
tubular campanulate or funnelform, deep yellow 
or yellow-orange, glabrous externally, tube 12-18 
mm wide at mouth, lobes 9-18 mm long; fila- 
ments ca. 20 and 14 mm long, thecae ca. 3 mm 
long, staminode rarely forming a fifth stamen. 
Fruits 24-145 cm long, 11-19 mm wide, linear, 
tapering at both ends, valves smooth, with minute 
peltate hairs, drying dark brown; seeds 9-14 mm 
long, 40-65 mm wide, mostly brown with lateral 
tips transparent. 

Deciduous lianas of seasonally dry deciduous, 
partly deciduous, and (less often) evergreen forest 
formations, 10-900(-1300) m elevation. Flower- 
ing primarily in February-June. This species 
ranges from Mexico and the West Indies to Ar- 
gentina. 

Macfadyena unguis-cati is recognized by its 
climbing habit with short distally three-clawed 
tendrils, opposite bifoliolate leaves, inflorescences 
usually flowering before the new leaves are fully 
flushed, thin calyx with irregular margin, deep 
yellow corollas glabrous externally, long-linear 
flattened fruits, and two-winged seeds. The un- 



usual dimorphism and climbing habit (see discus- 
sion under genus) are additional distinctions. The 
leaves on very young plants may be only 1 cm 
long and rounded (Standley, 1938). Gentry 
(1973b) reported that the seedlings of this species 
can persist for extended periods in drier areas. 
Una de gato. 



Mansoa DeCandolle 

Lianas climbing with distally trifid tendrils or 
simple tendrils with a terminal peltate disc, stems 
terete or somewhat quadrangular, with 4-8 phlo- 
em areas in cross-section, often with conspicuous 
glandular fields at the nodes; pseudostipules well 
developed to inconspicuous. Leaves opposite, 2- 
or 3-foliolate, glabrous or puberulent, petioles 
sometimes with a glandular field at the apex, mar- 
gin entire, venation pinnate or subpalmate. Inflo- 
rescences axillary to distal leaves or leafless 
nodes (terminal), panicles, racemes or small cor- 
ymbs, usually few-flowered, pedicels subtended 
by minute bracts or ebracteate. Flowers with cu- 
pulate or tubular-campanulate calyx, 5-lobed or 
the margin truncated, often with plate-shaped 
glands on the distal surface; corolla narrowly tu- 
bular at the base, distally tubular-funnelform to 
tubular-campanulate, white to reddish purple, pu- 
berulent externally (tube may be glabrous); sta- 
mens 4, filaments of 2 lengths, anthers glabrous 
or villous, thecae curved or straight, divaricate, a 
staminode present; disc annular-pulvinate; ovary 
cylindric, lepidote to papillate, 2-locular, ovules in 
2-4 series in each locule. Fruits linear-oblong 
capsules, valves parallel to the septum, woody, 
thin and flat to thick and terete, surface smooth 
or with elevated spines; seeds flat with 2 thin lat- 
eral distally membranaceous wings or corky and 
lacking lateral wings. 

Mansoa is a genus of 15 Neotropical species, 
mostly South American. This generic concept 
now includes a number of species formerly placed 
in Pachyptera and Pseudocalymma (Gentry, 
1973b; Standley & Williams, 1974). Five distinc- 
tive species are placed here: one with very small 
leaves (M. parvifolia), two whose vegetative parts 
smell like onions (M. hymenaea and M. standle- 
yi), one with verrucose-echinate fruits (M. verru- 
cifera), and one with large white corollas (M. ker- 
ere). Except for M. hymenaea, the species of this 
genus have rarely been collected in Costa Rica. 



132 



FIELDIANA: BOTANY 



Key to the Species of Mansoa 

la. Leaflets < 30 mm long; tendrils < 3 cm long, with flat disc-like pad at the apex 

M. parvifolia 

Ib. Leaflets > 40 mm long; tendrils usually > 5 cm long, usually ending in 3 slender tips (disc absent) 

2 

2a. Fruit surfaces covered with spine-like tubercles 1-5 mm long; gland fields absent at the nodes; 
calyx with well-defined lobes to 4 mm long [leaves without a garlic-like or onion-like odor) .... 

M. verrucifera 

2b. Fruits with flat smooth surfaces; gland fields usually present at the nodes; calyx with truncated or 

irregular margins or rarely with lobes to 2 mm long 3 

3a. Vegetative parts lacking the odor of onion; pseudostipules usually in a vertical series of 3 in 
each leaf axil, acute; leaves mostly 3-foliolate; inflorescences racemose; corolla pubcruleht on 

tube and lobes externally, white (in Central America) M. kerere 

3b. Vegetative parts with the odor of onion or garlic when crushed; pseudostipules usually incon- 
spicuous, obtuse; leaves 2-foliolate; inflorescences panicles or racemes, corolla puberulent only 

on the lobes externally, reddish purple 4 

4a. Calyx 4-7 mm long, green at maturity; fruits 15-25 mm wide, with midvein raised on surface of 

the valves; plants of deciduous and partly deciduous forests M. hymenaea 

4b. Calyx 9-21 mm long, green to purplish at maturity; fruits with the midvein obscure on the valve 
surface; plants mostly in evergreen forest formations M. siandleyi 



Mansoa hymenaea (DC.) A. Gentry, Ann. Mis- 
souri Bot. Card. 66: 782. 1979. Bignonia hy- 
menaea DC., Prodr. 9: 158. 1845. Adenocalym- 
ma macrocarpum J. D. Smith, Bot. Gaz., 40: 9. 
1915. A. ciliolatum Blake, Contrib. Gray Herb. 
52: 90. 1917. A. hosmeca Pittier, Contrib. U.S. 
Natl. Herb. 18: 256. 1917. Petastoma tondu- 
zianum Kra'nzl., Fedde Repert. 17: 56. 1921. 
Figure 22. 

Lianas to 5 cm diam., tendrils 10-15 cm long, 
leafy stems 0.8-7 mm diam., subtetragonal, gla- 
brous (new shoots sparsely minutely puberulent), 
becoming pale yellowish brown or gray; pseudo- 
stipules ca. 3 mm long, ovoid, striate. Leaves 2- 
foliolate, petioles 6-38 mm long, 0.7-2 mm 
diam., glabrous or very minutely (0.05 mm) pu- 
berulent, petiolules 8-26 mm long, thickened at 
the blade; leaflet blades 4-14 cm long, 2-8 cm 
wide, ovate to ovate-oblong or suborbicular, apex 
short-acuminate to bluntly obtuse, base obtuse to 
rounded-truncate (subcordate), drying stiffly char- 
taceous and grayish or green, glabrous on both 
surfaces, 2 veins 3-10/side. Inflorescences axil- 
lary or terminal, 4-10 cm long, panicles, pedun- 
cles 1-5 cm long, ca. 2 mm diam., pedicels 5-20 
mm long, sparsely to densely puberulent with thin 
straight hairs. Flowers with calyx 4-7 mm long, 
4-6 mm diam., cupulate, puberulent to subgla- 
brous, margin subentire or minutely 5-lobed, cil- 
iolate; corolla 37-65 mm long, tubular-campan- 



ulate (slightly 2-lipped), white to lavender or dark 
pink, glabrous or with the lobes puberulent exter- 
nally, tube 8-14 mm diam., lobes 9-14 mm long; 
filaments ca. 18 and 13 mm long, thecae ca. 3 
mm long. Fruits 15-34 cm long, 15-23 mm wide, 
linear-oblong, flat, midvein raised, surface 
smooth; seeds 11-15 mm long. 34-62 mm wide, 
center ca. 12 mm diam., not clearly differentiated, 
wings translucent. 

Common plants of seasonally dry deciduous 
and partly deciduous forests of the Pacific slope. 
10-1300 m elevation. Flowering primarily in De- 
cember-March. This species ranges from northern 
Mexico to southeastern Brazil. 

Mansoa hymenaea is recognized by its climb- 
ing habit with trifid tendrils, opposite bifoliolate 
broadly ovate leaves, short inflorescences, white 
to rose or lilac corollas puberulent distally. flat, 
narrow fruits tapering at both ends, and thin two- 
winged seeds. The strong odor of garlic or onions 
when vegetative parts are crushed is a very dis- 
tinctive feature. Specimens placed here were 
called Pseudocalymma sagotii (Bur. & K. 
Schum.) Sandwith by Williams in Flora of Gua- 
temala (Standley & Williams, 1974). Ajillo and 
pedo de padre are common names. 

Mansoa kerere (Aublet) A. Gentry, Ann. Mis- 
souri Bot. Gard. 66: 783. 1979. Bignonia kerere 
Aublet, Hist. PI. Guiane Fr. 2: 644, tab. 260. 
1775. Tanaecium zetekii Stand!., Contrib. Ar- 



BURGER: FLORA COSTARICENSIS 



133 



nold Arbor. 5: 140, tab. 19. 1933. Pachyptera 
kerere (Aublet) Sandw., Recueil Trav. Bot. 
Nteerl. 34: 219. 1937. Figure 22. 

Lianas to 5 cm diam., tendrils 6-16 cm long, 
leafy stems 1.8-6 mm diam., somewhat quadran- 
gular in cross-section, glabrous or minutely pu- 
berulent with thin straight hairs ca. 0.1 mm long, 
gland fields often conspicuous with sunken pits; 
pseudostipules 2-6 mm long, often 3/axil and the 
more abaxial progressively shorter, lanceolate. 
Leaves 2- or 3-foliolate, petioles 10-85 mm long, 
0.8-2.2 mm diam., minutely puberulent, longitu- 
dinally striate, with glandular pits at the apex, pet- 
iolules 12-61 mm long; leaflet blades 8-18(-24) 
cm long, 3-9(-ll) cm wide, ovate to ovate-ob- 
long or narrowly ovate-elliptic, apex acuminate to 
acute (rarely rounded or emarginate), base obtuse 
to rounded and narrowly subcordate, drying thinly 
chartaceous and brown, surfaces subglabrous with 
minute hairs along the major veins on both sur- 
faces, 2 veins 5-9/side. Inflorescences terminal 
or axillary, 1 .5-4 cm long, racemes, axis 2-3 mm 
diam., stem-like with many nodes and reflexed 
bracts 1-3 mm long, pedicels 8-14 mm long. 
Flowers with calyx 9-1 1 mm long, 5-8 mm 
diam., deep-cupular, minutely papillate-puberu- 
lent to subglabrous with peltate hairs, rounded 
glands present on distal half, margin with few 
short (2 mm) broad (3 mm) lobes or irregular; 
corolla 4-7 cm long, narrowly tubular-funnel- 
form, white or dull white (yellowish within), gla- 
brous proximally or minutely puberulent exter- 
nally, tube 3-9 mm diam., lobes 5-15 mm long; 
filaments ca. 20 and 15 mm long, thecae 2-4 mm 
long. Fruits (10-)22-42 cm long, 17-40 mm 
wide, narrowly oblong with narrowed apex and 
base, flat or somewhat rounded, midvein elevated 
or obscure, surface smooth and with round glands; 
seeds 18-24 mm long, 28-38 mm wide, with 
corky texture, wings absent. 

Rarely collected plants of wet evergreen forests 
on both the Caribbean and Pacific lowlands, 0- 
150 m elevation. These plants are usually found 
along stream margins, lake shores, and coastal 
marshes (Gentry, 1973b). This species has been 
collected in the Rio Sixaola (Talamanca Valley) 
and Golfo Dulce areas in Costa Rica. Probably 
flowering throughout the year. This species ranges 
from Belize to Brazilian Amazonia. 

Mansoa kerere is distinguished by its climbing 
habit with three-tipped tendrils, opposite two- or 
three-foliolate leaves, short racemose inflores- 
cences, long white puberulent corollas, pubescent 



anthers, and narrowly oblong fruits. Deeply pitted 
gland fields at the apex of the petiole and the 
three-seriate pseudostipules in leaf axils are useful 
vegetative distinctions. The seeds may be corky 
(the typical variety dispersed by water) or thin and 
winged, e.g., var. incarnata (Aublet) A. Gentry. 
The Central American collections all have white 
corollas, but some South American populations 
have red flowers (var. erythraea Dugand). 

Mansoa parvifolia (A. Gentry) A. Gentry, Ann. 
Missouri Bot. Gard. 66: 783. 1979. Pachyptera 
parvifolia A. Gentry, Phytologia 26: 448. 1973. 
Figure 16. 

Woody or herbaceous vines with stems to 1 cm 
diam., climbing with short (2-5 mm) tendrils 
bearing a flat rounded terminal disc 1-5 mm wide, 
leafy stems 0.2-0.7 mm diam., glabrous or sparse- 
ly puberulent with hairs ca. 0.1 mm long; pseu- 
dostipules ca. 0.3 mm long, conical. Leaves 2- 
foliolate and often with a tendril, petioles l-3(-8) 
mm long, glabrous abaxially and sparsely to 
densely minutely puberulent on the adaxial sur- 
face, petiolules 0.5-2(-7) mm long; leaflet blades 
(3-)6-18(-30) mm long, 2-9(-17) mm wide, el- 
liptic-oblong to oblong or oblong-obovate, apex 
obtuse (rounded), base asymmetric with cuneate 
and slightly rounded sides, drying chartaceous 
and grayish beneath, surfaces with short (0. 1 mm) 
thin straight hairs, 2 veins 2-4/side, often ob- 
scure, loop-connected. Inflorescences axillary at 
leafless nodes, short, 2- or 3-flowered, bracts ab- 
sent, peduncle 5-6 mm long, pedicels 1-1.5 mm 
long, subglabrous or with minute peltate hairs. 
Flowers with calyx 4-6 mm long, 5-8 mm wide, 
campanulate, truncated or slightly 2-lipped, sub- 
glabrous; corolla 35-50 mm long, tubular-funnel- 
form, white or pale purple, minutely puberulent 
externally, tube 8-12 mm diam. at the mouth, 
lobes 11-13 mm long; filaments 16-25 and 11- 
18 mm long, thecae 2-4 mm long. Fruits 23-43 
cm long, 20-25 mm wide, linear-oblong, surface 
smooth (midvein obscure), pale brown; seeds not 
seen. 

Rarely collected plants of evergreen lowland 
rain forest formations, 10-700 m elevation. Only 
sterile trunk-climbing specimens with very small 
leaves have been collected in Central America. 
This species ranges from Honduras to Bolivia. 

Mansoa parvifolia is one of the most distinctive 
species of Bignoniaceae. The slender stems cling- 
ing to tree trunks by means of short tendrils end- 
ing in a peltate disc and the very small opposite 



134 



FIELDIANA: BOTANY 



bifoliolate leaves are unique among the species of 
the family. The few-flowered inflorescences and 
puberulent white or rose corollas are additional 
characteristics but are rarely seen. The leaves be- 
come 20-30 mm long on distal flowering stems, 
but these stems have not been collected in Central 
America. The slender climbing stems with four 
small leaflets at each node are reminiscent of 
some Peperomia species with whorled leaves. 

Mansoa standleyi (Steyermark) A. Gentry, Ann. 
Missouri Hot. Card. 66: 783. 1979. Pseudoca- 
lymma standleyi Steyermark, Publ. Field Mus. 
Nat. Hist., Bot. Sen 23: 235. 1947. Pachyptera 
standleyi (Steyermark) A. Gentry, Brittonia 25: 
236. 1973. Ps. alliaceum var. macrocalyx 
Sandw., Kew Bull. 1953: 468. 1954. 

Lianas to 8 cm diam., tendrils 10-14 cm long 
with 3 distal arms to 2 cm long, vegetative parts 
smelling like garlic, leafy stems 2.5-8 mm diam., 
glabrous, longitudinally striate, nodes with a few 
punctate glands; pseudostipules small, conical. 
Leaves 2-foliolate, petioles 9-48 mm long, 1.7- 
2.7 mm diam., glabrous, petiolules 7-25 mm long, 
thickened at the apex; leaflet blades 10-20(-24) 
cm long, 6- 11 (-14) cm wide, elliptic-ovate to 
broadly ovate, apex acute to short-acuminate, base 
obtuse or rounded, drying green to olive green 
and chartaceous (subcoriaceous in life), surfaces 
glabrous, with gland fields in the axils of basal 
veins beneath, 2 veins 6-9/side. Inflorescences 
axillary or terminal, to 45 cm long, pyramidal 
panicles or reduced to racemose, usually with 
short (1-3 cm) opposite branches, peduncles to 18 
cm long, glabrous, bracts 8-19 mm long, 6-11 
mm wide, bright red, caducous, pedicels 8-20 mm 
long. Flowers with cupulate calyx (9-) 17-21 mm 
long, (8-) 15-1 8 mm wide, rounded at the base, 
glabrous, margin with 5 undulate lobes or sub- 
entire, thin-papery in life and green to red-violet; 
corolla (38-)66-85 mm long, tubular-campanu- 
late, rose-red to violet on lobes and upper tube, 
base of tube yellow, glabrous externally, lobes 6- 
27 mm long, apically obtuse; filaments 1 1-22 and 
22-30 mm long, thecae 3-4 mm long, linear. 
Fruits 35-75 cm long, 26-38 mm wide, elongate- 
linear, acute at both ends, flat (midvein obscure), 
pale brown; seeds 20-28 mm long, 75-110 mm 
wide, central area poorly differentiated, wings 
whitish, thin. 

Uncommon plants of evergreen forest forma- 
tions on both the Caribbean and Pacific slopes, 
50-1100 m elevation (1300-1400, m elevation in 



Guatemala). Probably flowering throughout the 
year. This species ranges from Mexico to Peru and 
Brazil. 

Mansoa standleyi is recognized by its climbing 
habit with distally trifid tendrils, mostly glabrous 
parts, opposite bifoliolate leaves with large leaf- 
lets, large cupulate calyces, large pinkish corollas, 
and long linear fruits with large winged seeds. 
The plants are also distinctive because of the on- 
ion-like odor of vegetative parts and flowers and 
the gland fields in basal vein axils of the lower 
leaf surface. This species was treated under Pa- 
chyptera in Flora of Panama and as Pseudoca- 
lymma sagotii var. macrocalyx (Sandw.) L. O. 
Williams in Flora of Guatemala (Standley & Wil- 
liams, 1974, p. 216). The type collection (Stey- 
ermark 33533 F) cites bejuco de ajo as a common 
name in Guatemala. 

Mansoa verrucifera (Schldl.) A. Gentry, Ann. 
Missouri Bot. Gard. 63: 62. 1976. Bignonia ver- 
rucifera Schldl., Linnaea 26: 655. 1853. Aden- 
ocalymma verruciferum (Schldl.) Miers, Ann. 
Mag. Nat. Hist., ser 3, 7: 393. 1861. A. fissum 
Loes., Verh. Bot. Vereins Prov. Brandenberg 
65: 102. 1923. A. seleri Loes., Verh. Bot. Ver- 
eins Prov. Brandenberg 65: 101. 1923. Ono- 
hualcoa seleri (Loes.) Lundell, Contrib. Univ. 
Michigan Herb. 7: 52. 1942. O. fissa (Loes.) 
Sandw., Kew Bull. 1946: 88. 1947. A. perezii 
Standl. & L. O. Williams, Ceiba 3: 61. 1952. 
O. verrucifera (Schldl.) A. Gentry, Ann. Mis- 
souri Bot. Gard. 60: 885. 1973 (1974). Figure 
18. 

Lianas to 20 m high, to over 8 cm diam., ten- 
drils to 15 cm long, leafy stems 1.3-9 mm diam., 
terete or slightly quadrangular, subglabrous or 
sparsely and minutely puberulent with peltate or 
straight hairs ca. 0. 1 mm long, V-shaped interpe- 
tiolar ridges often present, gland fields absent; 
pseudostipules inconspicuous. Leaves 3- or 2-fo- 
liolate, petioles 2-9 cm long, 1.3-2.7 mm diam., 
subglabrous, longitudinally striate, lateral petio- 
lules 8-35 mm long, terminal petiolule 3-7 cm 
long; leaflet blades 6-14(-17) cm long, 2.5-9(-l 1) 
cm wide, ovate-elliptic to ovate-triangular or 
ovate-oblong, apex acute to acuminate, base ob- 
tuse to rounded, drying chartaceous, upper surface 
glabrous except for the midvein, lower surface 
subglabrous, often with gland fields in proximal 
vein axils beneath, 2 veins 5-7/side, basal sec- 
ondary veins strongly ascending. Inflorescences 
axillary to leaves or fallen leaves, 3-12 cm long, 



BURGER: FLORA COSTARICENSIS 



135 



racemose or narrowly paniculate (thyrse) with 3- 
15 flowers, peduncle puberulent and lenticellate, 
pedicels 4-7 mm long, ca. 1 mm diam., sparsely 
to densely puberulent with hairs 0.1-0.2 mm long. 
Flowers with calyx 8-12 mm long, 4-6 mm 
diam., narrowly cupuiar, surface sparsely to 
densely minutely puberulent, with 5 longitudinal 
ridges terminating in apical teeth 0.5-3 mm long; 
corolla 47-82 mm long, tubular-campanulate, li- 
lac to purple, puberulent with thin curved hairs 
ca. 0.2 mm long externally, lobes 13-35 mm long, 
rounded distally, throat white; filaments 23-24 
and 11-16 mm long, thecae ca. 4 mm long. Fruits 
13-30 cm long, 17-33 mm wide, linear-oblong, 
valves flat to rounded and covered with tubercles 
1-5 mm long from a base 1-4 mm wide; seeds 
12-17 mm long, 37-58(-80) mm wide, wings dis- 
tally translucent, central area poorly differentiat- 
ed. 

Rarely collected plants of evergreen and partly 
deciduous forest formations along the Pacific slope 
in Costa Rica, 10-1 100 m elevation. Flowering in 
January and April (March-June from Mexico to 
Nicaragua); fruiting in December. The species 
ranges from Mexico to Bolivia and Guyana. 

Mansoa verrucifera is recognized by its climb- 
ing habit with tendrils that are distally trifid, op- 
posite two- or three-foliolate leaves with gland 
fields in vein axils beneath, short axillary inflo- 
rescences, prominently five-dentate calyx lobes, 
large purple or rose corollas minutely puberulent 
externally, and unusual woody fruits with verru- 
cose-tuberculate surface. The longer petiolules of 
terminal leaflets (in three-foliolate leaves), strong- 
ly ascending basal secondary veins, and stems 
lacking well-developed interpetiolar gland fields 
or pseudostipules are additional distinctions. This 
species was treated as Adenocalymma fissum in 
Flora of Guatemala (Standley & Williams, 1974) 
and as Onohualcoa verrucifera in Flora of Pan- 
ama (Gentry, 1973b). 



Marti nella Bail Ion 

Lianas climbing with distally trifid tendrils, 
stems terete, cross-sections of stems with large 
pith and 4 phloem areas, nodes with an interpe- 
tiolar ridge, without glandular fields; pseudo- 
stipules absent. Leaves opposite, 2-foliolate, pet- 
iolate, often bearing a distally trifid tendril, mar- 
gin entire, glabrous or with short glandular hairs, 
venation pinnate. Inflorescences axillary ra- 
cemes, pedunculate, bracts minute or absent, ped- 



icels prominent. Flowers with calyx closed and 
conical in bud, tubular-campanulate, opening ir- 
regularly with 4- or 5-lobed margin or splitting 
and 2-lipped; corolla tubular-campanulate or fun- 
nelform, dark wine-red, tube with a narrow base, 
glabrous or with minute peltate hairs externally; 
stamens 4, included, filaments of 2 lengths, an- 
thers glabrous, thecae straight, divaricate, a small 
staminode present; ovary linear-cylindric, 2-loc- 
ular, ovules in 4 (2) series in each locule. Fruits 
long linear capsules, valves parallel to the septum, 
thin and flat, smooth, midvein obscure, margins 
slightly thickened; seeds thin, transversely oblong 
with 2 poorly differentiated lateral membrana- 
ceous wings. 

Martinella is a genus of two species, one South 
American and one ranging from Mexico to Brazil 
and Bolivia. The very long fruits and unusual col- 
or of the corolla help distinguish our representa- 
tive of this genus. 

Martinella obovata (Kunth in H.B.K.) Bureau & 
K. Schum. in Mart., Fl. Bras. 8(2): 161, pi. 84. 
1896. Spathodea obovata Kunth in H.B.K. , 
Nov. Gen. Sp. 3: 147. 1819. Bignonia obovata 
(Kunth in H.B.K.) Spreng., Syst. Veg. 2: 830. 
1825. Figure 23. 

Lianas to over 25 m high, to 7 cm diam., ten- 
drils 5-25 cm long with tips 2-24 mm long, leafy 
stems 1 .7-9 mm diam., glabrous or minutely glan- 
dular puberulent, a raised interpetiolar ridge usu- 
ally present at nodes. Leaves 2-foliolate, petioles 
2-7 cm long, 1-2.3 mm diam., petiolules 12-32(- 
66) mm long, glabrous or very minutely (0.05 
mm) puberulent; leaflet blades 6-16(-21) cm 
long, 2.5-8(-15) cm wide, ovate-elliptic to ovate- 
oblong or broadly ovate, apex short-acuminate or 
acuminate with narrow tip, base obtuse to rounded 
(subcordate) and slightly asymmetric, drying 
chartaceous and often dark, glabrous on both sur- 
faces with very minute hairs along the major 
veins beneath, flat glands often present along the 
midvein beneath, 2 veins 3-7/side. Inflorescenc- 
es axillary, 5-15 cm long, racemes with 1-21 
flowers, peduncles 16-26 mm long, ca. 1.5 mm 
diam., glabrous or very minutely puberulent, 
bracts less than 1 mm long, pedicels 6-14 mm 
long. Flowers with calyx 11-16(-21) mm long, 
6-9(-12) mm diam., tubular or tubular-campanu- 
late, margin irregularly split or 2-4-lobed, surface 
glabrous with many parallel longitudinal veins or 
minutely puberulent; corolla 48-65 mm long, 
campanulate above a narrowed (3-5 mm diam.) 



136 



FIELDIANA: BOTANY 



tubular base, dark reddish purple or maroon, 8- 
21 mm wide at the mouth, lobes 6-15 mm long, 
glabrous or with minute glandular peltate hairs 
externally; filaments 15-17 and 11-14 mm long, 
thecae 2.5-3 mm long. Fruits 31-130 cm long, 
14-22 mm wide, 2-4 mm thick, valves flat with 
smooth surface, acute at the ends, midveins slight- 
ly elevated or obscure, margins thickened; seeds 
12-15 mm long, (30-H3-60 mm wide, central 
area ca. 17 mm wide, wings membranaceous, pale 
brownish. 

Uncommon plants of moist evergreen forest 
formations on both Caribbean and Pacific slopes, 
20-1600 m elevation. Flowering in February-July 
in Costa Rica. This species ranges from Mexico 
to Brazil and Bolivia. 

Martinella obovata is recognized by its climb- 
ing habit with distally trifid tendrils, opposite two- 
foliolate leaves often with long petioles, axillary 
few-flowered racemes, tubular calyx with irregu- 
lar margins, deep purple or maroon corollas, and 
long thin linear fruits with smooth flat valves. In 
addition, the petioles and petiolules are often 
twisted, and the leaflets are lustrous above in life. 
The fruits can be more than 1 m long. 



Melloa Bureau 

Lianas, tendrils with 3 terminal claw-like arms, 
stems terete, cross-section with irregular phloem 
areas, interpetiolar glandular fields present or ab- 
sent; pseudostipules ovate to subulate. Leaves op- 
posite, 2-foliolate, petiolate, often with a tendril, 
margins entire, glabrous, venation pinnate. Inflo- 
rescences terminal or axillary, panicles or few- 
flowered cymes, foliaceous bracts present and ear- 
ly caducous, pedicels prominent. Flowers with tu- 
bular-campanulate thin-textured calyx, distally ir- 
regularly lobed or spathaceous with recurved tip; 
corolla tubular-funnelform, yellow, glabrous ex- 
ternally; stamens 4, filaments of 2 lengths, anthers 
glabrous, thecae straight, divaricate, a staminode 
present; ovary flattened-ovoid, 2-locular, ovules 
multiseriate in each locule. Fruits capsules, ellip- 
soid-oblong and slightly flattened, valves parallel 
to the septum, thick and woody, splitting along 
the middle at maturity; seeds thin, transversely 
oblong with 2 lateral wings, seed body well dif- 
ferentiated from the membranaceous wings. 

Melloa has a single species, ranging from Mex- 
ico to Argentina. 



Melloa quadrivalvis (Jacq.) A. Gentry, Brittonia 
25: 237. 1973. Bignonia quadrivalvis Jacq., 
Fragm. Bot. 37, tab. 40, fig. 3. 1800-1809. Fig- 
ure 17. 

Lianas to over 20 m high, tendrils with 3 stiff- 
ened curved tips 11-18 mm long, leafy stems 1.3- 
6 mm diam., glabrous, longitudinally striate, in- 
terpetiolar ridge usually present; pseudostipules 
1.5-3.5 mm long, subulate. Leaves with petioles 
1 1-55 mm long, 1-2 mm diam., glabrous, striate, 
petiolules 6-20 mm long; leaflet blades 4-1 1 (-15) 
cm long, 2-5(-7) cm wide, ovate-elliptic to ovate- 
lanceolate or elliptic, apex acute to acuminate, 
base obtuse or slightly rounded and truncate, dry- 
ing chartaceous, glabrous above and below, 2 
veins 3-6/side. Inflorescences axillary or termi- 
nal (often on lateral short shoots), paniculate or 
dichotomous cymes, mostly 3-9 flowered, flowers 
usually in distal triads (rarely solitary), peduncles 
13-36 mm long, 0.8-1.3 mm diam.. glabrous and 
drying dark, with caducous bracts 7-21 mm long 
and 3-6 mm wide, pedicels 16-32 mm long. 
Flowers with tubular-campanulate calyx. 10-21 
mm long, 9-13 mm diam., surface glabrous (rare- 
ly puberulent), without glands, margin split and 
spathaceous or irregularly lobed; corolla 35-75 
mm long, tubular-funnelform, bright yellow, gla- 
brous or subglabrous externally (lobes ciliolate), 
tube 8-15 mm diam., lobes 10-18 mm long; fil- 
aments 21-28 and 14-18 mm long, thecae 4-5 
mm long; disc 1.5-2 mm high (subtended by a 
broad thin layer and appearing double). Fruits 8- 
15 cm long, 25-52 mm wide, 15-20 mm thick, 
ellipsoid-oblong with narrowed ends, valves 5-8 
mm thick, each splitting longitudinally into 2 
equal parts, surface dark brown with many pale 
lenticel-like spots; seeds 10-16 mm long, 25-52 
mm wide, central area 9-12 mm wide and clearly 
differentiated from the transparent wings. 

Uncommon lianas of seasonally dry deciduous 
or partly deciduous (less often evergreen) forest 
formations on both Caribbean and Pacific slopes, 
10-1000 m elevation. Flowering in February-Au- 
gust; fruiting throughout the year. This species 
ranges from northeastern Mexico to Argentina. 

Melloa quadrivalvis is recognized by its climb- 
ing habit with three-clawed tendrils, opposite two- 
foliolate glabrous leaves, few-flowered inflores- 
cences, bright yellow corollas, and thick woody 
ellipsoid-oblong fruits that split into four parts. 
The large floral bracts would be a useful feature 
for determination, but they are quickly caducous 
and rarely collected. These plants are similar to 



BURGER: FLORA COSTARICENSIS 



137 



those of Adenocalymma innundatum (q.v.), but 
their fruits are very different. 



Mussatia Bureau ex Baillon 

Lianas, climbing with distally coiled simple 
tendrils, branchlets quadrangular, with 8 (16) 
phloem areas in cross-section, nodes without in- 
terpetiolar gland fields; pseudostipules large and 
resembling small leaves. Leaves opposite, petio- 
late, 2-foliolate, often with a terminal tendril, ten- 
dril with 1 tip, leaflets petiolulate, margins entire, 
venation pinnate, domatia present. Inflorescences 
terminal, panicles with opposite branching, bracts 
inconspicuous, flowers pedicellate. Flowers with 
short calyx forming a shallow cup, margin entire 
to lobed or split; corolla funnelform and 2-lipped, 
yellow with red or brownish stripes within, mi- 
nutely glandular with peltate hairs externally; sta- 
mens 4, included, filaments of 2 lengths, anthers 
glabrous, thecae short, divergent or divaricate; 
disc cupular, ovary 2-locular, ovules in 4-6 series 
in each locule, stigma simple. Fruits oblong to 
linear-oblong capsules, valves parallel to the sep- 
tum, woody, flat or slightly convex, surface tu- 
berculate to muriculate; seeds thin, with 2 poorly 
differentiated lateral wings. 

Mussatia is a genus of two species; the other 
species is found in eastern Amazonia and Guiana. 

Mussatia hyacinthina (Standl.) Sandw., Recueil 
Trav. Bot. Neerl. 34: 218. 1937. Tynanthus hy- 
acinthinus Standl., Carnegie Inst. Wash. Publ. 
461: 87. 1935. Figure 19. 

Lianas to 30 m high, stems to 18 cm diam., 
tendrils 12-23 cm long, leafy stems 2.5-8 mm 
diam., strongly quadrangular with 4 raised longi- 
tudinal ridged edges, minutely (0.1 mm) puberu- 
lent but soon glabrescent and pale gray; pseudo- 
stipules usually 4/node, 3-18 mm long, ovate or 
rounded. Leaves with petioles 4-9(-12) cm long, 
1.5-2.5 mm diam., subglabrous or minutely pu- 
berulent, petiolules 1.4-4.5(-6) cm long; leaflet 
blades 7-20(-26) cm long, 4-13(-17) cm wide, 
broadly ovate to ovate-elliptic or ovate-suborbic- 
ular, apex acute to acuminate, base obtuse or 
rounded and truncate, drying chartaceous, gla- 
brous above, minutely papillate-puberulent on the 
veins or subglabrous beneath, with longer hairs in 
vein axils (domatia), 2 veins 4-7/side, minor ve- 
nation slightly raised above (dried). Inflorescenc- 
es terminal (often on short lateral shoots), 5-16 



cm long, peduncles 3-40 mm long, ca. 1.5 mm 
diam., with minute (0.1 mm) peltate hairs, dark 
brown, 2 peduncles ca. 1 cm long, bracts 0.5-1 
mm long, pedicels 2-5 mm long (above bract 
scars). Flowers with broadly campanulate calyx 
1.2-2.3 mm long, 3-3.5 mm wide, margin entire, 
glabrous; corolla 15-24 mm long, funnelform and 
bilabiate, white with maroon tips, becoming yel- 
lowish brown or reddish brown, minutely papil- 
late-puberulent, tube 2.5-7 mm diam., lobes 5-10 
mm long; filaments ca. 8-12 and 6-7 mm long; 
ovary 3 mm long, with minute peltate hairs. 
Fruits 17-37 cm long, 4-6 cm wide, ca. 2 cm 
thick, valves obtuse at both ends, surface muricate 
with rounded projections ca. 1 mm high, brown; 
seeds 20-43 mm long, 50-120 mm wide, central 
area 20-40 mm wide, wings membranaceous, 
pale brown. 

Uncommon plants of evergreen rain forest for- 
mations on both the Caribbean and Pacific slopes, 
50-900 m elevation. Flowering in February-May 
in Central America. This species ranges from 
eastern Mexico to Brazil. 

Mussatia hyacinthina is recognized by the 
woody climbing habit, two-foliolate leaves, small 
flowers, short calyx cups with entire margins, bi- 
labiate white or yellowish brown corollas, long 
thick muricate woody fruits, and large winged 
seeds. Leaves with hairs in vein axils beneath 
(domatia), square stems with four prominent ridg- 
es, rounded pseudostipules, and tendrils with sim- 
ple tips are useful vegetative distinctions. Com- 
pare this species with Cydista diversifolia, which 
has similar stems and pseudostipules. The fruits 
are similar to those of Callichlamys, where the 
valves have a smoother surface. 

The genera Onohualcoa and Pachyptera are 
now considered to be synonyms of Mansoa (q.v.). 



Parabignonia Bureau ex K. Schumann 

Lianas climbing with tendrils with 3 stiff 
curved distal "claws," stems terete, with 4 phlo- 
em areas in cross-section, interpetiolar gland 
fields present; pseudostipules small and ovate. 
Leaves opposite, 2-foliolate, petiolate, blades en- 
tire, usually glabrous, venation pinnate or subpal- 
mate. Inflorescences axillary, very short and few- 
flowered, usually solitary racemes, bracts short 
and narrow, pedicels subglabrous or minutely pu- 
berulent. Flowers with tubular-campanulate ca- 
lyx, 4- or 5-lobed, rounded or acute at the apex; 



138 



FIELDIANA: BOTANY 



corolla funnelform-campanulate and somewhat 2- 
lipped, glabrous externally or with minute pub- 
erulence on the lobes, lobes 5, rounded; stamens 
4, included, filaments of 2 lengths, anthers gla- 
brous, thecae straight, divergent, staminode pre- 
sent; ovary oblong, with minute peltate hairs, 2- 
locular, ovules 2-seriate in each locule. Fruits 
capsules, narrowly linear, valves parallel to the 
septum but dehiscing perpendicular to it; seeds 
thin, transversely oblong, with 2 membranaceous 
lateral wings. 

A genus of two species (more if Paradolichan- 
dra is included), one ranging from southern Costa 
Rica to Venezuela and the other in southern Bra- 
zil. Recently collected material (1992-1993) is the 
first record of the genus in Central America. 

Parabignonia steyermarkii Sandw., Bol. Soc. 
Venez. Cienc. Nat. 26: 446. 1966. Figure 17. 

Lianas climbing to over 25 m high, stems to 
19 cm diam., tendrils 5-12 cm long, with 3 
curved tips, leafy stems 2-6 mm diam., glabrous 
(in ours) or with short thin hairs, terete, becoming 
brown with elliptic lenticels ca. 1 mm long; in- 
terpetiolar gland fields inconspicuous. Leaves 2- 
foliolate, petioles 10-28 mm long, 1.2-1.7 mm 
diam., petiolules 4-1 1 mm long, glabrous, drying 
dark; leaflet blades 4-11 cm long, 2-6.5 cm 
wide, ovate-elliptic, ovate, ovate-oblong, or ellip- 
tic-oblong, apex acute or short-acuminate, base 
obtuse to slightly rounded, drying stiffly charta- 
ceous and dark, glabrous above and below, 2 
veins 4-7/side, basal pair strongly ascending 
(subtripli veined). Inflorescences axillary, 1-5- 
flowered racemes or corymbs or flowers solitary, 
1-3 cm long, peduncle 1-5 mm long, glabrous, 
drying dark, bracts 1.3-2.5 mm long, linear-lan- 
ceolate or subulate, pedicels 6-18 mm long, gla- 
brous or very minutely (0.03 mm) puberulent. 
Flowers with calyx 6-12(-22) mm long, 4-7(-14) 
mm wide, cupulate or campanulate, subglabrous, 
drying dark, lobes 2-4(-7) mm long, rounded to 
obtuse; corolla 35-62 mm long, funnelform-cam- 
panulate, deep magenta (in ours) to rose-purple or 
pale lavender, 10-15 mm wide at the mouth, tube 
glabrous but the lobes minutely papillate puber- 
ulent externally, lobes 10-17 mm long; filaments 
17-24 and 12-16 mm long, thecae 2.5-3.5 mm 
long. Fruits not seen at maturity, young fruits ca. 
10 cm long, 4-5 mm diam., valves glabrous; 
seeds unknown. 

Rarely collected climbers of lowland evergreen 
rain forest formations in the Golfo Dulce area at 



300-450 m elevation (300-1200 m in Venezuela). 
Collected with flowers in January (Aguilar 817, 
Gentry el al. 78682) at Rancho Quemada in the 
Reserva Forestal Golfo Dulce. This species is 
known only from southwestern Costa Rica and 
Venezuela. 

Parabignonia steyermarkii is recognized by its 
climbing habit with distally 3-clawed tendrils, op- 
posite 2-foliolate leaves, short few-flowered axil- 
lary inflorescences, 4- or 5-lobed calyx, and ma- 
genta corollas with externally glabrous tube and 
minutely puberulent lobes. Costa Rican collec- 
tions seem to have broader thinner leaves than 
those from Venezuela. In addition, the calyx is 
smaller in Costa Rican material (Venezuelan mea- 
surements are in parenthesis in the description 
above). 



Paragonia Bureau 

REFERENCE W. D. Hauk. A review of the ge- 
nus Paragonia (Bignoniaceae). Ann. Missouri 
Bot. Card. 85: 460. 1998. 

Lianas climbing with the aid of minutely sim- 
ple, bifid or trifid tendrils, stems terete, with 4 
phloem areas in cross-section, becoming lenticel- 
late, glabrate to lepidote or puberulent, nodes 
lacking gland fields; pseudostipules conical with 
broad base and narrow tip, usually curved in to- 
ward the stem. Leaves opposite, 2-foliolate, pet- 
ioles usually with gland fields near the apex, 
blades entire, glabrate to densely puberulent, pin- 
nately veined. Inflorescences terminal or axillary, 
paniculate, often large and many-flowered, mi- 
nutely bracteate, pedicellate. Flowers with cupu- 
late calyx, with minute peltate hairs, distally trun- 
cate or irregularly lobed or split, ciliolate; corolla 
tubular-campanulate, rose red to lilac or magenta, 
densely and minutely puberulent with crooked 
moniliform hairs; stamens 4, filaments of 2 
lengths, anthers glabrous, thecae straight, divari- 
cate; disc prominent, ovary narrowly cylindric, 
locules 2, ovules in 2 series in each locule. Fruits 
woody capsules, linear to linear-oblong, valves 
flattened parallel to the septum, surface smooth to 
tuberculatc (in ours), midvein slightly elevated or 
not apparent; seeds thin, transversely oblong with 
2 lateral wings, brown, the central area not dif- 
ferentiated from the wings. 

Paragonia is a genus of two species, one re- 
stricted to eastern Brazil. 



BURGER: FLORA COSTARICENSIS 



139 



Paragonia pyramidata (L. C. Rich.) Bureau, Vi- 
densk. Meddel. Dansk. Naturhist. Foren. Kjob- 
enhavn 1893: 104. \S94.Bignoniapyramidata 
L. C. Rich., Actes Soc. Hist. Nat. Paris 1: 110. 
1792. B. sinclairii Benth., Bot. Voy. Sulphur 
129. 1844. Tabebuia pyramidata (L. C. Rich.) 
DC., Prodr. 9: 214. 1845. Arrabidaea dichasia 
J. D. Smith, Bot. Gaz. 20: 6. 1895. Figure 23. 

Lianas, stems to 5 cm diam., often with thick- 
ened nodes, tendrils 5-17 cm long, distally simple 
or bifid (rarely trifid), leafy stems 2-6 mm diam., 
glabrous or less often minutely puberulent, inter- 
petiolar ridge often present; pseudostipules 3-7 
mm long, sharply acute. Leaves 2-foliolate, peti- 
oles 9-45 mm long, 1.4-3 mm diam., subgla- 
brous to minutely puberulent, often with glands 
near the apex, petiolules 7-35(-46) mm long, sul- 
cate above; leaflet blades 6-19(-25 cm) long, 3- 
9(-13) cm wide, elliptic-oblong to elliptic or 
broadly ovate-oblong, apex acute to short-acumi- 
nate or rounded, base obtuse to slightly rounded, 
drying stiffly chartaceous, glabrous above, gla- 
brous beneath or minutely puberulent on the mid- 
vein beneath, with minute (0.05 mm) punctate de- 
pressions beneath, 2 veins 4-8/side. Inflores- 
cences terminal, 10-30 cm long, peduncles 3-11 
cm long, ca. 2 mm diam., minutely puberulent, 
pedicels 4-9 mm long. Flowers with cupular ca- 
lyx 4-7 mm long, 4-8 mm diam., subglabrous or 
minutely puberulent, margin entire to split or with 
minute (0.3 mm) teeth; corolla 35-65 mm long, 
tubular-campanulate, pink to lavender or magenta 
(white at the base), densely puberulent with 
crooked hairs 0.1-0.3 mm long externally, 12-22 
mm wide at the mouth; filaments ca. 18 and 13- 
16 mm long, thecae 2-2.5 mm long. Fruits (10- 
)32-61 cm long, 10-17 mm wide, 5-8 mm thick, 
narrowed at base and apex, valves slightly rough 
to the touch with muricate surface of small pro- 
jections; seeds 8-12 mm long, 21-43 mm wide, 
oblong with blunt tips, uniformly grayish brown 
to dark brown. Common plants of evergreen low- 
land rain forest formations on both Caribbean and 
Pacific slopes, 5-800 m elevation. They are found 
on well-drained slopes and in poorly drained 
swamps. Flowering occasionally throughout the 
year but with the majority of flowering in Janu- 
ary-May. The species ranges from Mexico to 
Brazil and Bolivia. 

Paragonia pyramidata is recognized by its 
climbing habit with simple or bifid tendrils, op- 
posite two-foliolate leaves with gland field at pet- 
iole apex, many-flowered terminal panicles, cu- 



pular calyx with irregular margin, pink to magenta 
corollas densely puberulent on the exterior, long 
linear fruits with flat muricate surfaces, and thin, 
uniformly brown, blunt-ended, two-winged seeds. 
The pseudostipules and sweet smell when leaves 
are crushed are useful vegetative features. The im- 
mature calyx has a conical apex that sometimes 
splits off in a calyptrate fashion as in Lundia 
(q.v.). Compare also the vegetatively similar Cer- 
atophytum tetragonolobium, which differs in hav- 
ing trifid tendrils and interpetiolar gland fields. 

Parmentiera DeCandolle 

REFERENCE A. Gentry, Parmentiera in Big- 
noniaceae part I (Crescentiae and Tourrettieae). 
Flora Neotropica Monogr. 25(1): 1-130. 1980. 



Trees or shrubs, stems terete, leaves subtended 
by simple spines or hard smooth shelf-like tissue, 
an interpetiolar line or ridge absent; pseudostip- 
ules absent. Leaves opposite or subopposite (oc- 
casionally alternate), sometimes fasciculate, usu- 
ally 3- or 5-foliolate (rarely simple or 2- or 4- 
foliolate), petioles sometimes with narrow lateral 
wings, leaflets mostly glabrous, entire or with few 
distal teeth, venation pinnate, domatia usually 
present in vein axils beneath. Inflorescences ter- 
minal or often on leafless short-shoots or cauliflo- 
rous on trunk and older stems, with 1-3 flowers, 
bracts minute, pedicels drying black. Flowers 
drying black, calyx tubular, entire distally but split 
abaxially and spathe-like, subglabrous; corolla 
white to greenish white, tubular-campanulate and 
often bent in the middle (with a transverse ridge 
across the lower side) of the throat, 2-lipped with 
2 upper lobes partly united, thick-textured, gla- 
brous externally; stamens 4, subexserted near the 
mouth, filaments subequal, thecae thick, straight, 
somewhat divergent, a staminode present; disc an- 
nular; ovary cylindric, 2-locular, ovules multiser- 
iate. Fruits fleshy and indehiscent, elongate ob- 
long to linear, terete or with prominent longitu- 
dinal ridges, surface smooth, outer cortex firm and 
fleshy; seeds borne on a fibrous-fleshy central 
core, small, flattened, with or without a narrow 
circumferential wing. 

Parmentiera is a genus of ten species ranging 
from Mexico to Panama, with one in Colombia. 
The genus is unusual because of its glabrous flow- 
ers that dry black, tubular calyx split down one 
side, large white or greenish white two-lipped co- 
rollas with thick texture that usually have a trans- 



140 



FIELDIANA: BOTANY 



verse fold at the bottom of the tube, and the elon- 
gate fleshy fruits. The usually opposite three- or 
five-foliolate leaves, blades with domatia beneath, 
and hard spines or hard tissue at leaf bases are 
additional distinguishing features. All of the spe- 



cies are probably pollinated by bats, with seeds 
adapted to dispersal by mammals. Our introduced 
species may resemble Crescentia alata, but that 
species has leaves in alternate fascicles and the 
petioles are broadly winged. 



Key to the Species of Parmentiera 

la. Native trees found in evergreen forest formations; leaves usually 3-foliolate or 5-foliolate, petioles 
lacking narrow green margins, larger leaf blades 3-17 cm long 2 

Ib. Introduced trees planted in gardens and parks (rarely collected); leaves mostly 3-foliolate. petioles 

usually with narrow green margins or wings, larger leaf blades 2-9 cm long 4 

2a. Leaves usually 5-foliolate, largest blades to 9 cm long; 600-1500 m elevation in the Cordillera 

de Guanacaste and near Tilaran P. valerii 

2b. Leaves usually 3-foliolate, largest blades to 17 cm long; wet forests of the Caribbean slope 10- 

750 m 3 

3a. Corollas ca. 25 mm long; fruits ca. 15 mm diam.; rarely collected P. dressleri 

3b. Corollas 50-75 mm long; fruits ca. 30-60 mm diam.; more commonly collected 

P. macrophylla 

4a. Stems usually with 2 small hard spines subtending each node; flowers terminal or in axils of distal 
leaves; fruits > 30 mm diam. in life; petioles with very narrow margins P. aculeata 

4b. Stems without spines; flowers borne on trunks and older branches; fruits < 25 mm diam. in life; 
petioles often with winged margins P. cereifera 



Parmentiera aculeata (Kunth in H.B.K.) Seem., 
Bot. Voy. Herald 183. 1854. Crescentia aculea- 
ta Kunth in H.B.K., Nov. Gen. Sp. 3: 158. 
1819. C. edulis Moc. ex DC., Prodr. 9: 244. 
1845, non C. edulis Desv. P. edulis DC., Prodr. 
9: 244. 1845. Figure 11. 

Shrubs to medium-size trees 2-6 m tall, leafy 
stems 1-10 mm diam., glabrous, terete, leaf bases 
subtended by a forward-pointing sharp-tipped 
conical spine 3-8 mm long, with smooth glossy 
surface. Leaves opposite or occasionally alter- 
nate, 3-foliolate or simple, 2-4/node, petioles 1 1- 
47 mm long, 0.6-1.8 mm wide, with narrow 
wings 0.1-0.5 mm wide, petiolules 0-15 mm 
long; leaflet blades 25-90 mm long (distal leaf- 
lets), 6-40 mm wide, elliptic to rhombic-elliptic, 
apex obtuse, base cuneate and decurrent on the 
petiole, drying chartaceous, glabrous above, with 
depressions and minute hairs in the vein axils be- 
neath (domatia), 2 veins 3-7/side. Inflorescences 
of 1-4 terminal flowers or 1 or 2 flowers in axils 
of near-terminal leaves or fasciculate on older 
stems, pedicels 12-21 mm long, 0.7-1 mm diam., 
with few minute peltate trichomes, drying dark. 
Flowers with calyx 25-35 mm long, spathe-like 
or split more than once, glabrous, drying dark; 
corolla 4-6.5 cm long, campanulate, white or 



greenish with purple lines, glabrous externally, 
tube 20-25 mm wide at the mouth; thecae 8-9 
mm. Fruits 9-17 cm long, to 3 cm diam. in life 
(12-20 mm diam. when dried), linear-cylindric 
with narrowed tips, usually slightly curved, with 
thick longitudinal ridges. 

Pannentiera aculeata is recognized by the tree 
habit (usually with a short trunk and dense 
crown), nodes with short spines, mostly opposite 
and trifoliolate leaves, spathe-like calyx, white or 
greenish corollas, and slightly curved succulent 
fruits that are edible. This species has not been 
collected in Costa Rica, but Standley (1938) re- 
ported it to be cultivated in the hot lowlands. Its 
natural range is Mexico to Honduras. Common 
names in northern Central America are cuajilote, 
pepino de arbol, and cow okra. 

Parmentiera cereifera Seem, in Hook., J. Bot. 
3: 302. 1841. 

Small trees to 7 mm tall, trunks to 20 cm diam., 
with strongly ascending main branches from near 
the ground and open crown, leafy stems 1.2-6 
mm diam., glabrous, terete, usually with hard 
shelf-like tissue below the leaf base. Leaves op- 
posite, often with a smaller leaf in the axil of a 
larger (4/node), 3-foliolate, petioles l-4(-6) cm 



BURGER: FLORA COSTARICENSIS 



141 



long, to 3 mm wide with narrow greenish wings, 
petiolules not clearly differentiated; leaflet blades 
0.5_7(-9.5) cm long, 0.5-3(-4) cm wide, elliptic 
to elliptic-rhombic, terminal leaflets larger than 
the laterals, apex acute to acuminate, base acute 
and decurrent on the petiole, glabrous above, with 
small hairs in vein axils beneath (domatia), 2 
veins 3-6/side. Inflorescences of solitary flowers 
or 2 or 3 flowers on short-shoots borne on trunks 
and larger branches, peduncles (to bracts) ca. 3 
mm long, pedicels 9-16 mm long, 0.7-1 mm 
diam., glabrous or with minute peltate trichomes, 
drying black. Flowers with calyx 2-5 cm long, 
9-25 mm diam., split and spathe-like, glabrous; 
corolla 37-64 mm long, tubular-campanulate and 
with a transverse fold on the lower side, white, 
glabrous externally, tube 18-29 mm wide at the 
mouth, lobes ca. 10 mm long; filaments 26-35 
mm long. Fruits 30-54 cm long, 1-2.5 cm diam., 
linear-cylindric and slightly curved, yellow, gla- 
brous or with few minute peltate trichomes; seeds 
3-4 mm long, 3-4 mm wide, with a narrow mu- 
cilaginous wing. 

Parmentiera cereifera, originally described 
from the Caribbean slope of central Panama, is 
not known to occur in Costa Rica, but it is often 
planted as a botanical curiosity. The short trunk 
with spreading branches, opposite or fasciculate 
trifoliolate leaves, cauliflorous flowers, white co- 
rolla, and pendulous yellow linear fleshy fruits 
make the species quite distinctive. 

Parmentiera dressier! A. Gentry, Wrightia 7: 85. 
1982. 

Small trees 2-3 m tall, stems terete to suban- 
gulate, without spines or prominent lenticels. 
Leaves opposite or fasciculate, 3-foliolate, lateral 
petiolules poorly distinguished from the base of 
the blade, petioles 3-8 cm long, glabrous, petio- 
lules of terminal leaflet ca. 1-2 cm long; leaflet 
blades 5-13 cm long, 3.5-5.5 cm wide, elliptic to 
elliptic-ovate, apex acute to acuminate, base cu- 
neate, glabrous except for tufts of hairs in the vein 
axils below (domatia). Inflorescences cauliflorous 
or ramiflorous, usually of solitary flowers borne 
on pedicels less than 1 cm long, glabrous. Flow- 
ers with spathaceous calyx 16-17 mm long, lep- 
idote near the base, glabrous distally, bluntly 
acute; corolla ca. 25 mm long, broadly tubular- 
funnelform, white, 20 mm wide at the mouth, 
lobes ca. 8 mm long, glabrous; stamens and pistil 
not examined. Fruits 23-24 cm long, ca. 15 mm 
wide, cylindric, not ridged. 



Plants of rain forest formations in northern 
Costa Rica and the Caribbean lowlands. Flower- 
ing in August; fruiting in March (Panama). The 
species ranges from northern Costa Rica to central 
Panama. 

Parmentiera dressleri is recognized by its tree 
habit, trifoliolate leaves, solitary flowers with spa- 
thaceous calyx, and cylindric fruits. This species 
was at first thought to be a variant of P. macro- 
phylla, but the much smaller flowers and much 
narrower fruits indicate that the plants placed here 
are distinct. We have not seen Costa Rican ma- 
terial and base this report on the original descrip- 
tion and the Manual Flora manuscript of J. F. Mo- 
rales and Q. Jimenez (1997). 

Parmentiera macrophylla Standl., Publ. Field 
Columb. Mus. Hot. Ser. 4: 263. 1929. Figure 
11. 

Small or medium-size trees 3-12(-25) m tall, 
trunk unbranched for much of its length, to 30 cm 
diam., leafy stems 1.8-4 mm diam., glabrous, be- 
coming pale gray and terete, petioles often sub- 
tended by smooth thickened shelf-like tissue. 
Leaves opposite or subopposite, 3-foliolate (rare- 
ly 2-4-foliolate), petioles 2-10 cm long, 0.8-1.5 
mm diam., glabrous, sulcate above, lateral petio- 
lules 2-9 mm long and merging with base of 
blade, terminal petiolules 8-32 mm long; leaflet 
blades (2-)6-17 cm long, (l-)2-6 cm wide (lat- 
eral blades smaller than the terminal), elliptic to 
elliptic-obovate or elliptic-rhombic, apex acumi- 
nate or caudate-acuminate, base narrowly cuneate 
and decurrent on the petiole, drying thin-charta- 
ceous, subglabrous above, with minute sessile pel- 
tate trichomes beneath and tufts of hairs (or 
webbed tissue) in the vein axils (domatia), 2 
veins 4-7/side. Inflorescences cauliflorous, 1 or 
2 flowers borne on trunk and larger branches, ped- 
icels 1-2 cm long, glabrous or with minute (0.05 
mm) hairs, drying dark. Flowers with tubular ca- 
lyx 28-48 mm long, 8-13 mm diam., spathe-like 
(split abaxially), green or purplish, glabrous; co- 
rolla 50-75 mm long, trumpet-shaped to tubular 
campanulate, white or the tube greenish white, 
glabrous externally, with a saccate bulge in the 
floor of the throat, tube 10-14 mm diam., in the 
proximal half, mouth to 24 mm wide; filaments 
subequal, 32-38 mm long, thecae 5-6 mm long, 
1-2 mm wide. Fruits 20-47 cm long, 3-6 cm 
diam. in life, narrowly cylindric with longitudinal 
ribs (8-sided in cross-section), yellowish and 
fleshy (shrinking on drying); seeds ca. 6-9 mm 



142 



FIELDIANA: BOTANY 



long, 5-6 mm wide, flat, obovoid with rounded 
emarginate (obcordate) apex, brown. 

Trees of the shaded Caribbean rain forest inte- 
rior, 10-750 m elevation. Flowering irregularly 
throughout the year (more often in the wet sea- 
son). This species ranges from southeastern Nic- 
aragua along the Caribbean slope to central Pan- 
ama. 

Parmentiera macrophylla is recognized by its 
tree habit, usually opposite trifoliolate leaves, cau- 
liflorous flowers, spathe-like calyx, thick white 
corolla, and narrow fleshy fruits. This species may 
flower when only 3 m tall with a stem only 2.5 
cm diam. (Gentry, 1973b). It is similar to P. trun- 
ciflora Standl. & L. O. Williams, but the latter 
species has flowers borne on prominent (1-2 cm) 
woody short-shoots and grows at higher eleva- 
tions in the mountains west of Jinotega, Nicara- 
gua. 

Parmentiera valerii Standl., J. Wash. Acad. Sci. 
17: 16. 1927. Figure 11. 

Small to medium-size trees 5-20 m tall, trunk 
to 1 m diam., leafy stems 1.2-5 mm diam., gla- 
brous, pale grayish with conspicuous lenticels. 
Leaves opposite or subopposite, usually 5-folio- 
late, less often 3-7-foliolate, petioles 18-70 mm 
long, 0.8-1.5 mm diam., glabrous, petiolules 0- 
1 2 mm long and intergrading into the base of the 
blade, glabrous; leaflet blades (1.2-)2-9 cm long, 
(6-)8-34 mm wide, elliptic to narrowly elliptic or 
elliptic-obovate, apex acute to short-acuminate, 
base acute to cuneate and decurrent on the petiole, 
drying thin chartaceous, glabrous above or with 
the midvein minutely puberulent, glabrous be- 
neath but with crooked hairs in depressions in the 
vein axils (domatia), 2 veins 3-6/side. Inflores- 
cences of few fasciculate cauliflorous flowers on 
lower part of the trunk, pedicels 1 5-25 mm long, 
glabrous, drying dark. Flowers with tubular calyx 
32-40 mm long, 12-17 mm diam.. glabrous, 
spathe-like; corolla 60-75 mm long, tubular-cam- 
panulate, white or greenish white, glabrous exter- 
nally. Fruits 12-35 cm long, 2-4 cm diam. in life 
(1.2-3.5 cm dried), glabrous (purplish to green- 
violet in life); seeds 5-7 mm long, 7-8 mm wide. 



thin-lenticular, obovate with obtuse base and cor- 
date apex. 

Plants of the moist evergreen lower montane 
forest understory, 600-1500 m elevation. Flow- 
ering throughout the year. This species has been 
collected only on the Cordillera de Guanacaste 
and near TilarSn in northwestern Costa Rica. 

Pannentiera valerii is recognized by its tree 
habit, opposite usually pentafoliolate leaves, few 
cauliflorous flowers, spathe-like calyx, thick white 
corrolla, and pendulous cylindric fleshy fruits. 
The limited geographical area and narrow altitu- 
dinal range are also noteworthy. Jicaro danto and 
pepino de danta are common names. 



Phryganocydia Martius ex Bureau 

Lianas and herbaceous vines, climbing with 
simple (distally unbranched) tendrils, stems most- 
ly terete, cross-section with 4 or 8 phloem areas, 
interpetiolar gland fields absent; pseudostipules 
absent or early caducous. Leaves opposite, simple 
or 2-foliolate, petiolate, blades glabrous or with 
few minute peltate scales, venation pinnate. Inflo- 
rescences terminal or axillary, panicles with few 
flowers on bifurcating axes, or of 1 or 2 flowers, 
subglabrous with few peltate scales. Flowers with 
tubular-funnelform calyx split more than 60% and 
spathe-like, tip often thickened and reflexed; co- 
rolla tubular-funnel form, lavender to magenta, 
subglabrous with few minute peltate scales exter- 
nally, 2-lipped and 5-lobed; stamens 4, filaments 
of 2 lengths, anthers glabrous, thecae straight, di- 
varicate, staminode present; disc absent; ovary 2- 
locular. ovules in 2 series in each locule. Fruits 
linear-oblong flattened capsules, valves parallel to 
the septum, surface smooth but with a dense cov- 
ering of minute lepidote trichomes; seeds 2- 
winged with thin lateral wings or corky-ovoid 
without wings. 

Phryganocydia is unusual because of its spa- 
thaceous calyx with thickend tip and the fruit sur- 
face with a dense covering of minute hairs. Our 
species are mostly glabrous or with minute (0.05- 
0.1 mm) sessile peltate scales. 



Key to the Species of Phryganocydia 

la. Fruits narrowly oblong to linear, < 28 mm wide, to 60 cm long, surface grayish; seeds thin with 
lateral membranaceous wings; leaves consistently 2-foliolate; plants of well-drained sites . 

P. corymbosa 



BURGER: FLORA COSTARICENSIS 



143 



Ib. Fruits broadly ovate, > 35 mm wide, to 8 cm long, surface yellowish to dark brown; seeds thick 

and corky, lacking thin wings; leaves simple or 2-foliolate; plants of mangroves and lowland swamps 

P. phellosperma 



Phryganocydia corymbosa (Vent.) Bureau ex K. 
Schum. in Engler & Prantl, Naturl. Pflanzen- 
fam. 4(3b): 224, fig. 89H. 1894. Spathodea cor- 
ymbosa Vent., Choix, tab. 40. 1807; Mem. 
Math. Phys. Inst. Natl. France 1: 1-20. 1807. 
5. laurifolia Kunth in H.B.K., Nov. Gen. Sp. 3: 
114. 1819. 5. orinocensis Kunth in H.B.K., 
Nov. Gen. Sp. 3: 147. 1819. Figure 22. 

Lianas or herbaceous vines, 1-25 m high, to 5 
cm diam., tendrils 7-16 cm long, leafy stems 2- 
6 mm diam., sparsely to densely covered with flat 
peltate scales 0.05-0.1 mm diam., terete, with 8 
phloem areas in cross-section, interpetiolar ridge 
usually present. Leaves 2-foliolate (simple on 
young stems), petioles 8-33(-70) mm long, 1.2- 
2 mm diam., vesture similar to the stem, petiolules 
6-40 mm long; leaf blades 5-15(-22) cm long, 
3-9(-ll) cm wide, ovate-elliptic to ovate-oblong 
or oblong-rounded, apex acute to short-acuminate 
or rounded, base obtuse to rounded, stiffly char- 
taceous to subcoriaceous, glabrous on both sur- 
faces, 2 veins 3-7/side. Inflorescences terminal 
or axillary, 3-15 cm long, flowers usually cy- 
mose, peduncles to 5 cm long, pedicels 2-3 mm 
long (node to calyx base) and subtended by an 
internode 10-30 mm long, drying black. Flowers 
with calyx 26-37(-48) mm long, 3-14 mm diam., 
gradually expanding in width, with a straight or 
curved terminal projection 3-9 mm long, split 
down 1 side with entire margins, surface with few 
peltate scales; corolla 55-85(-97) mm long, tu- 
bular-funnelform, rose-lilac to purplish (often 
with a magenta area beneath the upper 2 lobes), 
subglabrous externally, 9-24 mm wide at the 
mouth, throat white with purple lines, lobes 1-3 
cm long; filaments 18-21 and 10-13 mm long, 
thecae 3.5-4.5 mm long. Fruits 13-53 cm long, 
16-26 mm wide, narrowly oblong to linear-ob- 
long, flat with smooth surface, grayish with dense 
covering of minute peltate scales, midvein slightly 
elevated; seeds 13-21 mm long, 27-73 mm wide, 
pale brown throughout, wings membranaceous. 

Plants of lowland evergreen or partly deciduous 
forest formations, 10-400 m elevation. Flowering 
in May-December in the Golfo Dulce area (not 
known elsewhere in Costa Rica); flowering 
throughout the year in Panama. The species is 
common in central and eastern Panama but has 



only been collected on the Burica Peninsula in 
western Panama. The species ranges from south- 
western Costa Rica to Brazil. 

Phryganocydia corymbosa is recognized by its 
climbing habit with simple tendrils, opposite bi- 
foliolate leaves, few-flowered dichotomously 
branched inflorescences, spathe-like calyx with 
terminal appendage, large pinkish corollas, and 
elongate flattened fruits differing greatly in length 
and with grayish crystalline surface. The leaflet 
blades are sometimes nearly as broad as long. An 
interesting feature of this species is that some 
plants flower while only 1 m high in open road- 
side vegetation. Vegetatively, this species is very 
similar to Cydista aequinoctialis, but the fruit of 
that species lacks the surface trichomes found in 
P. corymbosa. 

Phryganocydia phellosperma (Hemsl.) Sandw., 
Kew Bull. 1940: 302. 1941. Macfadyena phel- 
losperma Hemsl., Biol. Central Am. Bot. 2: 
492. 1892. 

Lianas to 2.5 cm diam., tendrils 8-15 cm long, 
stem with 4 phloem areas in cross-section, leafy 
stems 2-4 mm diam., terete, reddish brown and 
with scattered minute peltate scales, lacking con- 
spicuous lenticels; pseudostipules absent. Leaves 
2-foliolate or simple, petioles 8-42 mm long, 1- 
2.4 mm diam., subglabrous, petiolules 8-32 mm 
long, sulcate above; leaf blades 5-13 cm long, 3 
8 cm wide, ovate to ovate-triangular, ovate-oblong 
or oblong-elliptic, apex short- to long-acuminate, 
base obtuse to rounded and often truncate, drying 
chartaceous and grayish, glabrous above, glabrous 
or with few minute (0.05 mm) peltate scales be- 
neath, 2 veins 4-8/side. Inflorescences terminal, 
usually of 2 (1) flowers, peduncles ca. 10 mm 
long, 2 peduncles to 4 cm long, subglabrous, ped- 
icel 2-3 mm long, often terminated by a pair of 
linear-elliptic bracts 6-8 mm long. Flowers with 
calyx 18-27 mm long, 4-10 mm diam., terminat- 
ed by a thick tip 2-3 mm long, split along side, 
subglabrous; corolla 44-65 mm long, tubular- 
funnelform, lavender with a white throat, glabrous 
or with few minute hairs externally, 8-20 mm 
wide at the mouth, lobes 12-25 mm long; fila- 
ments 13-15 and 8-1 1 mm long. Fruits 4-8 cm 
long, 35-67 mm wide, ca. 6 mm thick, broadly 



144 



FIELDIANA: BOTANY 



ovate to ovate-suborbicular, surface dark brown 
and minutely papillate puberulent (smooth to the 
touch); seeds 20-27 mm long, 22-35 mm wide, 
thick, corky, and wingless. 

Rarely collected plants of mangrove swamps 
and wet muddy sites along the Pacific coast, 0- 
20 m elevation. Flowering mostly in the wet sea- 
son. The species ranges from southwestern Costa 
Rica to Colombia. 

Phryganocydia phellosperma is recognized by 
its climbing habit with simple tendrils, opposite 
bifoliolate or simple leaves, glabrous leaflets 
rounded at the base, usually two-flowered termi- 
nal inflorescences, spathe-like calyx, pink or ma- 
genta corollas, flattened broadly ovate fruits, and 
corky seeds lacking thin lateral wings. The seeds 
of this species are probably dispersed by water, 
consistent with its restriction to mangroves and 
low wet sites. 



Pithecoctenium Martius ex Meisner 

Lianas, tendrils trifid with 0-5 further distal 
divisions (sometimes with adhesive pads), stems 
strongly 6-angled, with 4 phloem areas in cross- 
section, interpetiolar lines often present, gland 
fields absent; pseudostipules narrowly spatulate or 
lanceolate, caducous. Leaves opposite, 2- or 3- 
foliolate (occasionally simple), petiolate, margin 
entire, venation palmate to pinnate. Inflorescen- 
ces terminal (or terminal on axillary short-shoots), 
racemes or racemose panicles, bracteate, flowers 
pedicellate. Flowers with cupulate calyx, distally 
truncated or minutely 5-toothed, densely puberu- 
lent, without glands; corolla tubular-campanulate, 
relatively thick, puberulent externally, 2-lipped, 5- 
lobed; stamens 4, filaments of 2 lengths, anthers 
glabrous, thecae straight, divaricate, a small stam- 
inode present; ovary 2-locular, ovules multiseriate 
in each locule. Fruits thick woody capsules, 
valves parallel with the septum and somewhat 
compressed, covered with short woody spines; 
seeds thin, with membranaceous wing around 3 
sides, central body clearly differentiated from the 
wings. 

Pithecoctenium is a Neotropical genus of four 
species, one wide-ranging and the others found in 
Argentina and eastern Brazil. 

Pithecoctenium crucigerum (L.) A. Gentry, Tax- 
on 24: 121. 1975. Bignonia crucigera L., Sp. 
PI. 2: 624. 1753. B. echinata Jacq., Enum. PI. 
Carib. 25. 1760. B. tiliaefolia Kunth in H.B.K., 



Nov. Gen. Sp. 3:. 136. 1819. P. echinatnm 
(Jacq.) Baill., Hist. PI. 10: 8. 1888. Figure 18. 

Lianas to ca. 10 cm ilium., to over 20 m high, 
tendrils 5-20 cm long, with 3-15 slender divi- 
sions (sometimes terminating in a disc that be- 
comes attached to bark), leafy stems 1.5-5 mm 
ilium., with minute peltate scales and with or 
without slender hairs ca. 0.2 mm long, with 6 
prominent longitudinal ridges; pseudostipules 8- 
15 mm long, 1-2 mm diam. Leaves opposite, 2- 
or 3-foliolate (simple), petioles 14-72 mm long, 
1-2 mm diam., subglabrous or with thin hairs ca. 
0.2 mm long, petiolules 28-80 mm long, with lon- 
gitudinal ridges; leaf blades (4-)6-18 cm long, 
(2.5-)4-13 cm wide, ovate to broadly ovate or 
ovate-oblong, apex acuminate, base rounded and 
truncate to cordate, drying thin chartaceous, both 
surfaces with lustrous peltate scales 0.05-0. 1 mm 
diam., simple hairs 0.2 mm long present or absent, 
venation palmate or subpalmate, 2 veins 4-67 
side, strongly ascending, with disc-like glands at 
the base beneath. Inflorescences 3-19 cm long, 
3-15 flowered racemes or racemose panicles, pe- 
duncle 1.5-2.5 mm thick, resembling the stems, 
lateral branches 7-12 mm long, bracts 1-6 mm 
long, pedicels 1-2 mm long. Flowers with calyx 
7-11 mm long, 8-10 mm wide, densely puberu- 
lent and lepidote, margin entire or minutely 5- 
dentate, eglandular; corolla 32-62 mm long, tu- 
bular-campanulate but often bent at almost 90 
near the base, white or yellowish white, densely 
puberulent externally with minute hairs (drying 
yellowish), lobes 6-17 mm long; filaments 17-21 
and 12-17 mm long, thecae 3-4 mm long. Fruits 
12-23(-31) cm long, 50-65(-75) mm wide, 23- 
38 mm thick, ellipsoid-oblong with somewhat 
flattened sides, surface densely tuberculate with 
woody projections 1-3 mm high, 1.5-2.5 mm 
diam. at the base, yellowish brown; seeds 24- 
36(-41) mm long, 45-88(-95) mm wide, central 
area 18-28 mm wide, wings transparent. 

Common lianas of deciduous, partly deciduous, 
and wet evergreen forest formations on both Ca- 
ribbean and Pacific slopes, 1-1100 m elevation. 
Flowering primarily in May-June; fruiting mostly 
in December-February. This species ranges from 
Mexico to Argentina. 

Pithecoctenium crucigerum is recognized by its 
climbing habit with multifid tendrils, opposite bi- 
or trifoliolate leaves (occasionally simple), race- 
mose inflorescences, usually entire densely pu- 
berulent calyx cups, densely puberulent white co- 
rollas sharply bent near the middle, echinate-tu- 



BURGER: FLORA COSTARICENSIS 



145 



berculate woody fruits, and thin seeds with large 
thin transparent wings on three sides. The hex- 
agonal twigs, elongate pseudostipules, multi- 
branched tendrils (sometimes terminating in 
discs), subpalmate leaf venation, and minute shiny 
peltate scales on leaves are useful vegetative at- 
tributes. Vegetatively, these plants resemble Am- 
philophium, but those species have branched hairs 
and trifid tendrils and lack the elongate pseudos- 
tipules of P. crucigerum. The seeds are often 
called mariposas and palomitas in Central Amer- 
ica (Standley, 1938). Batefta, cucharilla, peine de 
mico, and peina de mono are common names. 



Pleonotoma Miers 

Lianas and vines, climbing with tendrils usu- 
ally 3-parted at the apex, stems hollow at the cen- 
ter, with 4 radiating phloem arms in cross-section 
and quadrangular, an interpetiolar line usually pre- 
sent at the node; pseudostipules leaf-like, tubular, 
or absent. Leaves opposite, bi- or triternate to tri- 
pinnate, the terminal petiolule of the first division 
often replaced by a tendril, margins entire, vena- 
tion pinnate, domatia sometimes present. Inflo- 
rescences axillary or terminal racemes, bracts 
small, lateral bracteoles sometimes present, pedi- 
cels well developed. Flowers with calyx cupular, 
truncated at the apex and subentire or with 5 mi- 
nute teeth; corolla tubular-funnelform, white to 
yellowish, usually glabrous along the tube and pu- 
berulent distally; stamens 4, of 2 lengths, includ- 
ed, anthers glabrous, thecae straight, divaricate, 
pollen grains 3-colpate, a staminode present; disc 
cupular-pulvinate; ovary oblong, puberulent or 
lepidote, ovules many in 2 series on the central 
placentae. Fruits capsule, linear to narrowly ob- 
long, valves flat, paralleling the septum, smooth; 
seeds thin and with 2 membranaceous lateral 
wings. 

Pleonotoma is a genus of 14 species; all are 
South American except for one, which ranges 
northward into Central America. 

Pleonotoma variabilis (Jacq.) Miers, Proc. R. 
Hort. Soc. N.S. 3: 184. 1863. Bignonia varia- 
blilis Jacq., Hort. Schoenb. 2: 45, tab. 212. 
1797. Figure 16. 

Vines or lianas, climbing by tendrils (3-15 cm 
long) with 3 short distal arms 5-20 mm long, larg- 
er stems ca. 3 cm diam., leafy stems 2-7 mm 
diam., with 4 prominent longitudinal ridge, subgl- 



abrous; pseudostipules 2-5 mm long. Leaves op- 
posite, bicompound with 2- or 3-trifoliate parts, 
1 petioles 1-7.5 cm long, 1-2 mm diam., gla- 
brous, 2 petioles 2-6 cm long, longitudinally stri- 
ate, glabrous, petiolules 7-40 mm long with those 
of central leaflet often twice as long as laterals; 
leaflet blades 3-18 cm long, 1-9 cm wide, elliptic 
to elliptic-ovate or elliptic-obovate, apex acute to 
acuminate, base obtuse to rounded (lateral leaflets 
often asymmetric at the base), drying chartaceous, 
glabrous except for short (0.2-0.5 mm) whitish 
hairs along the midvein and vein axils beneath 
(domatia), 2 veins 4-7/side. Inflorescences usu- 
ally terminal racemes, peduncle and rachis 2-8 
cm long, with 3-12 flowers, pedicels 6-14 mm 
long, 0.6-1.4 mm diam., drying dark, with minute 
peltate glands or subglabrous. Flowers with cu- 
pular calyx 5-9 mm long, 5-7 mm diam., mostly 
glabrous, margin subentire with teeth ca. 0.2 mm 
long and a ciliolate or glabrous edge; corolla 50- 
90 mm long, tubular-funnelform, white or yellow- 
ish white, tube 10-20 mm wide at the mouth, gla- 
brous along the tube and minutely glandular lep- 
idote distally, lobes 14-22 mm long, rounded; 
longer stamens ca. 27 mm long, thecae ca. 4 mm 
long, divaricate; ovary ca. 4 X 1.5 mm. Fruits 
18-35 cm long, 1.5-2.7 cm wide, 3-7 mm thick, 
linear to linear-oblong, surface flat and slightly 
muricate, drying dark and lustrous; seeds 1-1.5 
cm long, 3-5 cm wide, central part ca. 12 mm 
wide, wings translucent at the tips. 

Uncommon evergreen lianas of deciduous, 
partly deciduous, and evergreen forest formations, 
10-800 m elevation. Known from the Caribbean 
lowlands of Nicaragua but as yet collected only 
on the Pacific slope in Costa Rica. Flowering in 
February-May; fruiting in December-April. The 
species ranges from Guatemala to Trinidad and 
the Amazon Basin. 

Pleonotoma variabilis is recognized by its vin- 
ing habit with trifid tendrils, the strongly four-an- 
gled hollow stems, the twice-compound opposite 
leaves (each of which have two or three trifoliate 
parts), short racemes, truncated calyx cup, and 
white or yellowish corolla. The tendrils are usu- 
ally borne from a 1 petiole that also bears two 
trifoliolate 2 petioles. This species and the her- 
baceous Tourrettia lappacea are the only Costa 
Rican climbing bignons having twice-compound 
leaves. 



Podranea Sprague 

Scandent shrubs or climbing vines, tendrils ab- 
sent, stems terete, gland fields absent at the nodes; 



146 



FIELDIANA: BOTANY 



pseudostipules not developed. Leaves opposite, 
imparipinnate with opposite lateral leaflets, blades 
entire or distally serrate, venation pinnate. Inflo- 
rescences terminal, solitary, paniculate with op- 
posite lateral branches, pedicels subtended by ca- 
ducous bracts. Flowers large, calyx tubular or 
campanulate (slightly inflated), glabrous, with 5 
prominent lobes shorter than the tube; corolla tu- 
bular-funnelform or tubular-campanulate, laven- 
der, slightly 2-lipped, glabrous externally, the 5 
lobes rounded; stamens 4, included, filaments of 
2 lengths, anthers glabrous, thecae straight, par- 
allel and later divergent, staminode small; disc 
saucer-shaped, thick; ovary ovoid, 2-locular, 
ovules ca. 6-seriate in each locule, stigma of 2 flat 
ovate lobes. Fruits elongated linear capsules, 
valves leathery, compressed parallel to the sep- 
tum, dehiscing perendicular to the septum; seeds 
flat, 2-winged. 

Podranea is a monotypic African genus of east- 
tropical and southern Africa, now a popular gar- 
den ornamental in the tropics. The name Podra- 
nea is an anagram of Pandorea, a similar and 
closely related genus of five species native to Ma- 
laysia, Australia, and the western Pacific (with 
two species widely cultivated ornamental vines). 
Pandorea differs in having pod-like oblong cap- 
sules with woody valves, and a noninflated calyx. 

Podranea ricasoliana (Tanfani) Sprague in Thi- 
selton-Dyer, Fl. Cap. 4 (2): 450. 1904. Tecoma 
ricasoliana Tanfani, Bull. Soc. Tosc. Ortic. 
1887: 17. 1887. Pandorea ricasoliana (Tanfani) 
Baill., Hist. PI. 10: 40. 1888. Figure 15. 

Scandent shrubs and vines, tendrils absent, 
leafy stems 1-5 mm diam., glabrous, becoming 
pale gray in age with rounded lenticels, interpe- 
tiolar lines usually present. Leaves 6-25 cm long, 
with usually 5-13 leaflets, petioles 16-60 mm 
long, 0.6-1.5 mm diam., glabrous, petiolules 2- 
12 mm long; leaflet blades (12-)20-55 mm long, 
8-25 mm wide, lanceolate to ovate-lanceolate or 
ovate-elliptic, apex acuminate, margin with 3-9 
teeth/side, base somewhat asymmetric (rounded 
and cuneate) on lateral leaflets, glabrous, 2 veins 
3-5/side. Inflorescences terminal, 6-18 cm long, 
paniculate with short opposite lateral branches, 
glabrous, bracts 2-4 mm long, subulate, pedicels 
4-14 mm long. Flowers with calyx 14-20 mm 
long, 6-9 mm diam., whitish to pale lavender in 
life, with few round glands on the distal surface, 
with 5 apiculate lobes 3-6 mm long; corolla 5-8 
cm long, tubular-campanulate, pale lavender to 



pink with purple areas near the 2 adaxial lobes 
(white inside), glabrous externally but with dense 
crooked hairs in the sinuses of the lobes; filaments 
18-20 and 14-16 mm long, thecae 3-4 mm long. 
Fruits 25-30 cm long, linear and terete (rarely 
produced in cultivation). 

Podranea ricasoliana, native of southern Afri- 
ca, is now widely cultivated as a garden orna- 
mental. It is recognized by its clambering habit 
(without tendrils), opposite pinnately compound 
leaves, narrow leaflets with serrate margins, 
slightly inflated calyx with prominent lobes, and 
large pink corollas, often with tufts of hairs in the 
sinuses of the lobes. It flowers throughout the year 
and does well from near sea level to 2000 m el- 
evation when grown on a wall or trellis in open 
sunny locations. Called bombilla, linda, and m(r- 
ame linda in Guatemala. 



Pyrostegia Presl 

Vines or lianas, climbing with the aid of coiling 
petiole-borne tendrils usually trifid near the tip, 
stems with longitudinal ribs, lacking interpetiolar 
gland fields; pseudostipules small or inconspicu- 
ous. Leaves opposite or subopposite, petiolate, 2- 
or 3-foliolate, petiolules present, margins entire, 
vention pinnate. Inflorescences terminal or axil- 
lary panicles, often much branched, pedicels sub- 
tended by small paired bracts. Flowers with cu- 
pular calyx, minutely glandular lepidote or subgl- 
abrous, distal margin truncated with 5 small teeth; 
corolla tubular or narrowly funnelform, orange or 
reddish orange, with 5 lobes, valvate in bud; sta- 
mens 4, exserted, anthers glabrous, thecae 
straight, parallel or slightly divergent; disc annu- 
lar; ovary narrowly tubular, stigma bifid. Fruits 
dry capsules, valves parallel to septum, smooth 
and leathery, with an indistinct median vein; seeds 
thin, 2-winged and transverse-oblong, wings 
smooth and with hyaline margins. 

A genus of four species, originally native to 
tropical South America. One species is now wide- 
ly planted for its showy orange flowers. 

Pyrostegia venusta (Ker-Gawler) Miers, Proc. R. 
Hort. Soc. N.S. 3: 188. 1863. Bignonia venusta 
Ker-Gawler, Bot. Reg. 3: 5. 1818. B. ignea 
Veil., Fl. Flumin. 244. 1825. P. ignea (Veil.) K. 
B. Presl., Bot. Bemerk. 93: 1843. Figure 15. 

Vines, climbing with tendrils to 15 cm long, 
leafy stems 2-5 mm diam., glabrous or with short 



BURGER: FLORA COSTARICENSIS 



147 



(0.2-0.5 mm) hairs near the nodes, with 6 or 8 
longitudinal ribs. Leaves 3-foliolate or 2-foliolate 
and often with a tendril, petioles 7-35 mm long, 
0.8-1.8 mm diam., with short hairs along the ad- 
axial side, petiolules 4-18 mm long (distal leaflet 
with longer petiolules); leaflet blades 3-9 cm 
long, 1.5-6 cm wide, ovate-elliptic to ovate-ob- 
long, apex acute to caudate-acuminate, base ob- 
tuse to slightly rounded and truncate or subcor- 
date, drying chartaceous, glabrous, minutely 
punctate beneath, 2 veins 3-5/side. Inflorescenc- 
es dense panicles usually terminal on short shoots, 
often pendulous with 8-25 flowers, rachis to 9 cm 
long, borne on woody peduncles 5-50 mm long, 
1-2.5 mm diam., glabrous or puberulent along 1 
side, pedicels 12-20 mm long, glabrous. Flowers 
with calyx 4-5 mm long, 4-6 mm wide, puber- 
ulent and glandular lepidote or subglabrous, distal 
margin minutely ciliolate, subentire or with mi- 
nute (0.3 mm) lobes (teeth); corolla 6-9 cm long, 
2-3 mm diam. near the base, 8-12 mm wide at 
the mouth, orange, tubular to narrowly tubular- 
funnelform, straight or slightly curved, lobes 6- 
18 mm long, oblong with blunt apex, becoming 
reflexed, whitish puberulent along the margin; sta- 
mens slightly exserted, filaments 3-5 cm long, 
thecae ca. 4 mm long. Fruits 20-30 cm long, 10- 
16 mm wide; seeds 1.2-1.6 mm long, 3-4 cm 
wide. 

Pyrostegia venusta is recognized by its climb- 
ing habit with distally trifid tendrils, opposite 2- 
or 3-foliolate leaves, long narrowly tubular orange 
corollas, and exserted stamens. These plants are 
often seen climbing over garden walls, with their 
clusters of brilliant orange flowers making a fine 
display. The corollas are usually slightly curved, 
held horizontally when in anthesis, and drooping 
afterward. Native to Brazil, Bolivia, and Para- 
guay, this species is now planted as an ornamental 
throughout the tropics. In Central America it is 
grown from near sea level to 1500 m elevation. 
This species has been called chorro, chorro de 
oro, San Carlos, and Triquitraque in Central 
America. 



Saritea Dugand 

Lianas climbing with simple tendrils, young 
stems terete and without interpetiolar gland fields; 
pseudostipules conspicuous and leaf like (ca. 5- 
25 mm long). Leaves opposite, short-petiolate, 2- 
foliolate, leaf blades 5-12 cm long, 3-7 cm wide, 
cuneate at the base, subglabrous, basal 2 veins 



strongly ascending, usually with a gland field in 
the axil of proximal veins beneath. Inflorescences 
usually terminal with 1-9 flowers, peduncles and 
pedicels drying dark, subglabrous with few mi- 
nute lepidote hairs, flowers sometimes in distal 
triads. Flowers with tubular subglabrous calyx 6- 
10 mm long, margin truncate; corolla 6-9 cm 
long, tubular-campanulate, magenta, glabrous ex- 
ternally; stamens 4, filaments ca. 2.8 and 2.1 mm 
long, anthers glabrous and straight, staminode 
small; disc small; ovules 2-seriate (or appearing 
1 -seriate) in each locule. Fruits linear capsules, 
valves flattened parallel to the septum; seeds thin, 
with 2 membranous lateral wings. 

Saritea magnifica (Sprague ex van Steenis) Du- 
gand, native to Colombia and Ecuador, is the only 
species of this genus and is widely planted as an 
ornamental. This taxon is closely related to Cy- 
dista and Phyrganocydia and also to Arrabidaea 
(Gentry, 1973b). Although not recorded from 
Costa Rica, this species may be expected in gar- 
dens. The bifoliolate leaves with cuneate-based 
leaflets, gland fields in vein axils beneath, con- 
spicuous pseudostipules, and large magenta co- 
rollas make it easy to identify. 

Schlegelia is now placed in the family Schle- 
geliaceae (q.v.). 

Scobinaria is now considered to be part of Ar- 
rabidaea. See Gentry's discussion (Selbyana 2: 
43-45. 1977). 



Spathodea Beauvois 

Trees with well-developed trunks, nodes lack- 
ing interpetiolar lines or glands; pseudostipules 
absent. Leaves opposite or 3/node, petiolate, im- 
paripinnately compound, lateral leaflets opposite, 
venation pinnate. Inflorescences terminal and 
erect, compact racemes and somewhat corymb- 
like with the lower pedicels longer than the distal. 
Flowers large, calyx spathe-like and curved with 
a narrowed apex, opening from beneath (abaxi- 
ally); corolla with a narrow cylindric base and 
large curved-campanulate throat, deep red to or- 
ange-red, glabrous on the exterior, 5-lobed; sta- 
mens 4, anthers with slender divaricate thecae; 
ovary 2-locular, narrowly oblong, ovules multis- 
eriate in each locule, stigma flat and bifid. Fruits 
narrowly oblong-ellipsoid capsules, dehiscence 
perpendicular to the septum along 1 side; seeds 



148 



FIELDIANA: BOTANY 



flat with broad thin hyaline wing around the entire 
circumference. 

The genus Spathodea, member of the tribe Te- 
comeae, contains a single species native to tropi- 
cal Africa. A densely puberulent variant of East 
Africa is sometimes segregated as 5. nilotica 
Seem. The compact terminal inflorescences with 
large red corollas and spathaceous calyx are dis- 
tinctive. 

Spathodea campanulata Beauvois, Fl. Oware 1: 
47, tab. 27. 1805. Figure 15. 

Trees to 25 m tall, trunks to 40 cm diam., bark 
somewhat rough and scaling near the base, leafy 
stems 3-12 mm diam., minutely appressed puber- 
ulent, lenticellate. Leaves 19-40 cm long, with 
7-13(-17) leaflets, petioles to 20 cm long, petio- 
lules 1-3 mm long (terminal leaflet with petiolule 
to 15 mm long), ca. 1 mm diam., puberulent; leaf 
blades 5-14 cm long, 2-6 cm wide, elliptic to 
ovate-elliptic or ovate-oblong, apex acute to acu- 
minate, base acute to obtuse or rounded on 1 side 
and asymmetric, puberulent on the veins beneath 
to densely puberulent throughout, with a gland at 
the base of the midvein abaxially, 2 veins 5- 107 
side. Inflorescences compact terminal corymb- 
like racemes with a closely placed spiral of 6-35 
flowers, ca. 7 X 15 cm (not including corollas), 
pedicels to 6 cm long, distal pedicels shorter than 
the proximal and giving a flat-topped effect. 
Flowers with calyx 2-6 cm long, split on 1 side, 
recurved and acuminate, densely sericeous, yel- 
lowish brown; corolla 6-15 cm long, 5-9 cm 
wide at the apex, widely campanulate and curved 
from a short-tubular base, deep red to orange-red, 
lobes 2-3 cm long, obtuse, distal edge of the co- 
rolla crinkled and yellowish. Fruits 17-27 cm 
long, 3-7 cm wide, 1-2 cm thick, narrowly ellip- 
soid-oblong; seeds ca. 15 X 20 mm, with thin 
membranaceous wing. 

Spathodea campanulata is recognized by its 
tree habit, opposite pinnately compound leaves, 
compact terminal inflorescences, curved spatha- 
ceous calyx, and very large curved-campanulate 
red corollas. This is a very striking ornamental 
tree when in full flower, with the huge corollas 
often in a whorl-like configuration. The distal un- 
opened flower buds often form a tight cluster on 
which birds can perch while taking nectar from 
the upwardly open corollas. Native to evergreen 



forests of tropical central Africa, the species is 
now cultivated throughout the tropics. In Costa 
Rica it has been grown successfully in evergreen 
and partly deciduous areas below 1500 m eleva- 
tion. African tulip tree, llama del bosque, and tn- 
lipdn are common names. 



Sti/(>ph\ limn Miers 

Lianas or vines, climbing with simple or trifid 
tendrils, stems with 4 phloem areas in cros's-sec- 
tion, terete, longitudinally striate, hollow in the 
center, nodes without gland fields; pseudostipules 
spatulate or inconspicuous, caducous. Leaves op- 
posite, 2- or 3-foliolate, petiolate, blades with 
conspicuous pellucid-lustrous round sessile glands 
on the lower surface, venation palmate or subpal- 
mate. Inflorescences axillary to distal (sometimes 
early developing) leaves, few-flowered racemes, 
peduncles short, bracts well developed, pedicels 
puberulent. Flowers with cupulate or campanulate 
calyx, often slightly inflated, 5-lobed or irregular- 
ly split (2-lipped), densely puberulent. thin-tex- 
tured; corolla campanulate-funnelform, slightly 
2-lipped, white or yellowish white, puberulent and 
with minute glandular hairs externally, the lobes 
often tinged with pink, lobes rounded; stamens 4. 
filaments of 2 lengths, anthers glabrous or puber- 
ulent along the lines of dehiscence, thecae 
straight, divaricate, staminode present; disc thick; 
ovary linear-tetragonal, 2-locular, ovules 2-seriate 
in each locule. Fruits linear capsules, valves flat- 
tened parallel to the septum, slightly convex, de- 
hiscence septicidal, surface puberulent; seeds 
transversely oblong, thin and flat, with 2 translu- 
cent lateral wings weakly differentiated from the 
central area. 

A Neotropical genus of three species, one of 
which ranges from Mexico to Amazonia. The ge- 
nus is unusual because of the pellucid dots on the 
lower leaf surfaces, the hollow stems, and the 
very slender fruits, which may resemble stems. 
Central American material of our two species is 
difficult to separate on the basis of herbarium ma- 
terial. Both species exhibit a wide range of vari- 
ability as regards density of pubescence, size of 
leaves, and size and shape of calyx and corolla. 
Study in the field is the basis for determing their 
status as separate species (Gentry, 1 973b, p. 936). 



BURGER: FLORA COSTARICENSIS 



149 



Key to the Species of Stizophyllum 

la. Branchlets and petioles yellowish brown to reddish brown or dark brown with hairs 0.5-2 mm long; 

tendrils with 3-branched tips; calyx 10-18 mm long; fruit surface with longer (to 1 mm) hairs . . . 

5. inaequilaterum 

Ib. Branchlets and petioles grayish or pale brownish with short (0.1-0.5 mm) hairs; tendrils simple 

with unbranched tip (rarely 2-branched); calyx 7-12 mm long, chartaceous; fruit surface with minute 

(0. 1-0.3 mm) hairs S. riparium 



Stizophyllum inaequilaterum Bureau & K. 
Schum. in Mart., Fl. Bras. 8(2): 221. 1896. Big- 
nonia inaequilatera Poeppig ex Bureau & K. 
Schum. in Mart., Fl. Bras. 8(2): 222, 1896, pro 
syn., non Poeppig ex Bureau, Adansonia 8: 289. 
1868. Figure 26. 

Lianas and vines, to 3 cm diam., tendrils 8-18 
cm, with 3 short (6-20 mm long) distal parts, 
leafy stems 2-7 mm diam., densely villous with 
yellowish brown hairs 0.5-2 mm long, older 
stems glabrescent; pseudostipules to 8 mm long. 
Leaves 2- or 3-foliolate, petioles 22-68(-150) 
mm long, densely villous, petiolules 9-28(-60) 
mm long; leaflet blades 7-13(-19) cm long, 4- 
7(-14) cm wide, ovate to oblong or elliptic-ob- 
long, apex acute to acuminate, margin entire or 
denticulate, base of lateral leaflets asymmetric 
with a rounded and a cuneate side, drying char- 
taceous, upper surface with slender curved hairs 
0.5-1 mm long, lower surface more densely vil- 
lous, venation pinnate or subpalmate, 2 veins 3- 
6/side. Inflorescences a few-flowered terminal 
fascicle or contracted raceme on a short axillary 
shoot, peduncles ca. 1 cm long, bracts conspicu- 
ous, linear, pedicels 8-24 mm long, densely vil- 
lous with reddish brown hairs. Flowers with calyx 
10-18 mm long, 7-10 mm diam., inflated-cam- 
panulate, thin-chartaceous, villous with hairs to 
1 .5 mm long, margin irregular or with lobes to 4 
mm long; corolla 5-7 cm long, tubular-funnel- 
form, greenish yellow with pink lobes (purple), 
tube puberulent externally, lobes 8-1 1 mm long; 
1 1 km ic ntx 20-21 and 15-16 mm long, thecae 2 
mm long. Fruits 24-35 cm long, 4-6 mm wide, 
3-4 mm thick, linear, densely brownish villous; 
seeds not seen. 

Rarely collected plants of lowland evergreen 
rain forest formations on the Caribbean slope, 0- 
500 m elevation. Flowering in March-May. The 
species ranges from eastern Nicaragua to Peru. 

Stizophyllum inaequilaterum is recognized by 
its climbing habit with trifid tendrils, dense pu- 
bescence of reddish brown hairs, opposite bi- or 



trifoliolate leaves, short few-flowered inflores- 
cences, thin villous calyx with irregular margins, 
puberulent corollas with pink lobes, and slender 
long-linear pubescent fruits. The dense pubes- 
cence of longer (0.5-2 mm) hairs and distally tri- 
fid tendrils (rarely well preserved in herbarium 
material) help distinguish this species from its 
more common congener. 

Stizophyllum riparium (Kunth in II BK 
Sandw., Lilloa 3: 462. 1938. Bignonia riparia 
Kunth in H.B.K., Nov. Gen. Sp. 3: 138. 1819. 
Adenocalymma flos-ardeae Pittier, Contrib. 
U.S. Natl. Herb. 18: 256. 1917. A. punctifolium 
Blake, Contrib. U.S. Natl. Herb. 24: 22. 1922. 
5. flos-ardeae (Pittier) Sandw., Recueil Trav. 
Bot. Neerl. 34: 212. 1937. 5. punctifolium 
(Blake) Sandw., Recueil Trav. Bot. Need. 34: 
212. 1937. Figure 23. 

Lianas and vines to 5 cm diam., tendrils 7-15 
cm long, apex simple or rarely with reduced sec- 
ond tip, leafy stems 1.3-6 mm diam., sparsely to 
densely puberulent with hairs to 0.7 mm long, te- 
rete; pseudostipules 3-8 mm long, strap-shaped. 
Leaves 2- or 3-foliolate, petioles 12-110 mm 
long, ca. 1 mm diam., puberulent, lateral petio- 
lules 7-37 mm long (terminal to 70 mm); leaflet 
blades 3.5-1 7(-20) cm long, 2-9.5(-12) cm wide, 
ovate-elliptic, ovate, or elliptic-oblong to narrow- 
ly ovate-oblong, apex acute to short-acuminate, 
margin entire or with 1-7 obtuse teeth/side in ear- 
ly stages, base rounded and truncate to subcor- 
date, drying thinly chartaceous, sparsely puberu- 
lent with slender hairs 0.2-0.6 mm long above, 
more densely puberulent beneath, with pellucid 
dots 0.1-0.2 mm diam. beneath, venation subpal- 
mate, 2 veins 4-6/side. Inflorescences terminal 
fascicles or axillary few-flowered racemes, pe- 
duncles 1-5 cm long, puberulent like the stems, 
pedicels 4-7 mm long. Flowers with calyx 7-12 
mm long, 4-9 mm diam., deeply cupular or tu- 
bular-funnelform, densely minutely puberulent, 
margin 5-lobed or irregular, glandular on lobes; 



150 



FIELDIANA: BOTANY 



corolla 32-53 mm long, tubular-funnelform to tu- 
bular-campanulate, white to pale yellowish white, 
minutely papillate puberulent, lobes 5-8 mm 
long; filaments 15-18 and 1 1-12 mm long, thecae 
2-2.5 mm long. Fruits 24-45 cm long, 3-8 mm 
wide, 3.5-5 mm thick, linear and narrowed at the 
ends, straight or more often curved, lenticular in 
cross-section, surface densely minutely puberu- 
lent, grayish; seeds 4-6 mm long, 14-26 mm 
wide, transversely oblong, wings translucent. 

Climbers in lowland evergreen rain forest for- 
mations, 0-300 m elevation. Flowering mostly in 
June-November (but rarely seen with flowers and 
rare in herbaria). This species ranges from eastern 
Mexico to the Amazon basin. 

Stizophyllum riparium is recognized by its 
climbing habit with simple tendrils, hollow pu- 
berulent stems, opposite bi- or trifoliolate leaves, 
blades with pellucid dots beneath, white or yel- 
lowish corollas puberulent externally, and the 
very narrow linear usually curved fruits. Leaflets 
on juvenile growth may have conspicuous broad- 
based teeth. The differences between this and the 
preceding species are often very difficult to dis- 
cern in herbarium material. 



Tabebuia Gomes ex De Candolle 

REFERENCES A. Gentry, A revision of Tabe- 
buia (Bignoniaceae) in Central America. Brittonia 
22: 246-264. 1970. A. Gentry, Bignoniaceae 
Part II (tribe Tecomeae). Flora Neotropica, Mon- 
ogr. 25(11). 1992. 

Small to large trees or shrubs, wood usually 
dense and fine-grained, branchlets terete, nodes 
lacking interpetiolar gland fields; pseudostipules 
absent. Leaves opposite, palmately 3-7(9-)-fo- 
liolate or less often simple or 1 -foliolate, petiolate, 



basal leaflets usually smaller and with shorter pet- 
iolules than the more terminal, venation pinnate. 
Inflorescences terminal, short, open or congested 
panicles, few-flowered racemes or fascicles of 1 
to few flowers, bracts subtending the pedicels; 
some species flowering when the tree is leafless. 
Flowers with tubular to cupular or campanulate 
calyx, margin 2-5-lobed, irregular or truncate, 
surface with flat rounded appressed, stellate or 
straight hairs; corolla tubular-funnelform to tu- 
bular-campanulate, white, yellow, rose, purple, or 
red, glabrous or puberulent externally, lobes 5. 
rounded; stamens 4, exserted or included, fila- 
ments of 2 lengths, anthers glabrous, thecae 
straight, divaricate, staminode present; disc cush- 
ion-shaped, ovary oblong, 2-locular, ovules in 2 
to many series in each locule. Fruits capsules, 
linear-cylindric to oblong-cylindric, subterete, 
valves dehiscing approximately perpendicular to 
the septum, surface smooth to verrucose-muricate, 
glabrous to variously puberulent; seeds thin, 
transversely oblong with 2 thin lateral wings 
(thicker, corky, and without wings in T. palustris 
et al.). 

Tabebuia, a member of tribe Tecomeae, is a 
Neotropical genus of 100 species, with many spe- 
cies in South America and the West Indies. In 
Central America most of the species can be dis- 
tinguished by the tree habit, opposite usually pal- 
mately compound leaves, compact but showy ter- 
minal inflorescences (often in flower when the 
leaves are absent), large corollas, and the linear 
to oblong fruits that are rounded in cross-section 
and split at right angles to the septum. In our spe- 
cies, the corolla may be white, yellow, or rose to 
purple. When in full flower, these are some of the 
most beautiful trees in Central America. The 
wood of this genus is a commercially important 
tropical hardwood, and the bark contains phar- 
macologically active compounds (Gentry, 1992). 



Key to the Species of Tabebuia 

la. Seeds corky, without thin transparent lateral wings; fruits to 1 1 cm long; corollas white with yel- 
lowish markings within; shrubs and small trees of mangroves and river deltas along the Pacific 
coast; leaves simple or 3-foliolate, leaflets mostly narrowly lanceolate T. palustris 

Ib. Seeds thin with transparent lateral wings; fruits usually > 15 cm long; corolla yellow to rose or 

purple (white with pink lobes in T. rosea); medium to large trees, not found in mangrove formations 

(occasionally in lowland riversides); leaves usually 3-, 5-, or 7-foliolate, leaflets rarely lanceolate 

2 

2a. Corollas rose, purple, or white, marked with pink; leaflets and stems without stellate hairs; fruits 
glabrous or with flat rounded appressed peltate (lepidote) hairs or minutely papillate, the surfaces 
smooth . 



BURGER: FLORA COSTARICENSIS 



151 



2b. Corollas bright yellow; leaflets with stellate hairs (sometimes only in vein axils beneath and difficult 
to see); fruits stellate-pubescent to minutely villous or with an undulate surface if with minute 

rounded appressed (lepidote) hairs 4 

3a. Surface of leaflets with minute flat rounded (lepidote) hairs; corolla glabrous externally; calyx 
with minute flat rounded hairs; capsule with minute papillate or appressed peltate hairs; native 
in dry deciduous to wet evergreen rain forest formations and planted as ornamental, 0-1500 m 

elevation T. rosea 

3b. Surface of leaflets with straight hairs, at least in the vein axils beneath; corolla puberulent 
externally; calyx with minute straight or scurfy hairs; capsule mostly glabrous; in deciduous 

and partly deciduous forest, 5-300 m elevation T. impetiginosa 

4a. Leaflets with minute stellate hairs only in the vein axils beneath, leaves often 7-foliolate; fruits 
with slightly undulate surface and appressed rounded flat (lepidote) hairs [trees of lowland rain 

forests] T. quayacan 

4b. Leaflets with stellate hairs scattered on the surface beneath or along the veins, leaves rarely 7- 

foliolate; fruit with few to many minute stellate or tomentulous hairs 5 

5a. Leaves puberulent mostly along veins and petioles; calyx without long simple hairs, lobes 
becoming reflexed; corolla lobes with dark interconnecting veins to the distal margin; fruits 

minutely stellate-puberulent; trees of evergreen rain forest formations (in Costa Rica) 

T. chrysantha 

5b. Leaves densely puberulent over the entire surface beneath; calyx yellowsh pubescent with thin 
straight hairs 2-7 mm long, lobes erect; corolla lobes with venation not so clearly reaching the 

margin; fruits tomentulous; trees of deciduous and partly deciduous forest formations 

. T. ochracea 



Tabebuia chrysantha (Jacq.) G. Nichols., 111. 
Diet. Gard. 4: 1. 1887. Bignonia chrysantha 
Jacq., Hort. Schoenb. 2: 45, tab. 211. 1797. Te- 
coma chrysantha (Jacq.) DC., Prodr. 9: 221. 
1845. Tec. evenia J. D. Smith, Bot. Gaz. 20: 8. 
1895, pro parte (fls.). Tec. palmeri Kranzl., Fed- 
de Repert. 17: 220. 1921. Tab. chrysantha ssp. 
pluvicola A. Gentry, Phytologia 35: 190. 1977. 
Figure 13. 

Trees to 30 m tall, trunks to 1 m diam., bark 
light gray and smooth with few widely spaced 
long vertical furrows, leafy stems 3-9 mm diam., 
terete, densely puberulent with stellate hairs 0.1- 
0.3 mm long, glabrescent, becoming pale gray. 
Leaves usually 5-foliolate (3-, 4-, 7-foliolate), 
petioles (5-)8-19(-28) cm long, 1.5-3.5 mm 
diam., densely stellate puberulent, petiolules 1-9 
cm long (basal laterals 8-20 mm), with node-like 
area below the blade; leaflet blades (4-)7-21(-26) 
cm long, (2-)3-12 cm wide (proximal lateral leaf- 
lets smaller than the distal), elliptic to broadly el- 
liptic or elliptic-obovate, apex acuminate to cau- 
date-acuminate, margin entire or sometimes blunt- 
ly serrate, base obtuse to rounded and slightly 
truncate, drying thinly to stiffly chartaceous, 
subglabrous to sparsely stellate puberulent above, 
more densely stellate-puberulent beneath, peltate 
trichomes 0.02-0.05 mm diam. often conspicuous 



beneath, 2 veins 6-12/side. Inflorescences ter- 
minal, condensed (almost fasciculate) panicles 
with 7-25 flowers, peduncles 4-20 mm long, 
densely stellate pubescent, pedicels 4-20 mm 
long, 1.2-2 mm diam. Flowers with calyx 8-14 
(-19) mm long, 7-10(-13) mm diam., cupular- 
funnelform or campanulate, densely pubescent 
with hairs 0.3-0.8 mm long, lobes 2-3 mm long, 
broadly triangular; corolla 52-95 mm long, tu- 
bular-funnelform, bright yellow with narrow red- 
dish lines within, glabrous externally, lobes 9-28 
mm long; filaments 16-22 and 10-15 mm long, 
thecae 2-3 mm long. Fruits 20-50(-90) cm long, 
14-24 mm wide, linear-cylindric, narrowed at the 
ends, surfaces smooth and minutely stellate pu- 
berulent; seeds 6-12 mm long, 14-38 mm wide, 
wings transparent, well differentiated. 

Trees of wet evergreen lowland rain forest for- 
mations on both the Caribbean and Pacific slopes 
in Costa Rica and Panama, 20-600 m elevation 
(also occurring in much drier habitats in its range 
from Mexico to Nicaragua and in Venezuela). 
Flowering primarily in January-March, occasion- 
ally at other times. The species ranges from Mex- 
ico, disjunctly, to Peru. 

Tabebuia chrysantha is recognized by its large 
tree habit, small stellate hairs on many parts, op- 
posite palmately 5-foliolate leaves, compact ter- 
minal inflorescences, large brilliant yellow exter- 



152 



FIELDIANA: BOTANY 



nally glabrous corollas, and long linear-cylindric 
fruits with transparent-winged seeds. The leaflets 
are sometimes serrate. The trees make a brilliant 
show when flowering, in part because they usually 
flower when all the leaves have fallen. Common 
names are corteza and cones amarilla. 

Our collections are placed in subspecies pluvi- 
cola A. Gentry, which ranges from northeastern 
Costa Rica to northern Venezuela and Ecuador. 
Subspecies chrysantha is found from Mexico to 
Nicaragua and disjunctly in Amazonian Peru. The 
two subspecies can be separated by the following 
key. 



la. Trees usually 10-20 m tall, usually found in 
seasonally dry deciduous or partly deciduous 
forests below 1 200 m elevation; calyx < 1 mm 
long, densely rufescent with stellate to curved- 
barbate hairs ca. 1 mm long; fruits 15-50 X 
0.8-2 cm, persistently stellate-tomentose, often 
striate or with rough surface. . . . subsp. chry- 
santha 

Ib. Trees to 30 m tall, usually found in lowland 
and lower montane evergreen forest formations, 
0-1500 m; calyx > 10 mm long, rufescent with 
short stellate hairs ca. 0.5 mm long; fruits 30- 
80 X 1 .5-2.4 cm, nearly glabrescent and smooth. 
subsp. pluvicola 



Tabebuia guayacan (Seem.) Hemsley, Biol. 
Centr. Am. Bot. 2: 495. 1882. Tecoma quaya- 
can Seem., Bot. Voy. Herald, 180. 1854. 

Trees to 50 m tall, trunks to 1.7 m diam., bark 
gray or brown and ridged with well-developed 
vertical furrows, leafy stems 2-8 mm diam., 
subglabrous or with few minute hairs, becoming 
pale grayish, terete. Leaves 5- or 7-foliolate, pet- 
ioles, 8-23 cm long, 1.5-3 mm diam., sparsely 
puberulent with scurfy hairs or glabrous, petio- 
lules 1-7 cm long (basal lateral petiolules 1-3 cm 
long), geniculate and drying dark below the blade; 
leaflet blades 5-20(-30) cm long, 3-1 1(- 15) cm 
wide (terminal larger than laterals), ovate-elliptic 
to elliptic or lanceolate, apex acute to acuminate, 
margin entire (serrate in juveniles), base rounded 
to obtuse (lateral leaflets usually asymmetric), 
drying thin-chartaceous, subglabrous or minutely 
puberulent on the veins beneath and with short 
stellate hairs in vein axils (domatia), lower surface 
with minute appressed peltate hairs, 2 veins 6-97 
side. Inflorescences compact panicles with flow- 



ers in diads or triads and opening at the same 
time, peduncles 6-14 mm long, pedicels 10-18 
mm long, 0.6-0.9 mm diam., sparsely puberulent 
with scurfy hairs 0.1-0.3 mm long. Flowers with 
calyx 9-15 mm long, 4-9 mm diam., campanu- 
late, with minute scurfy scattered hairs, margin 
split or with 2-5 rounded/obtuse lobes; corolla 
63-105 mm long, tubular- funnelform, yellow with 
brown lines within, glabrous externally, 13-22 
mm wide at the mouth, lobes 22-35 mm long; 
filaments 15-21 and 11-16 mm long, thecae 2-3 
mm long. Fruits 30-60 cm long, 12-28 mm 
wide, linear, surface subglabrous to thick stellate- 
puberulent; seeds 6-1 1 mm long, 23-40 mm 
wide, wings transparent and clearly differentiated. 

Rarely collected trees of lowland evergreen rain 
forest formations, 5-600 m elevation. Flowering 
in February-March (March-May elsewhere in 
Central America); seeds are released in May-July. 
The species ranges from Mexico to Peruvian 
Amazonia. 

Tabebuia quayacan is recognized by its tree 
habit, opposite palmately penta- or septafoliolate 
subglabrous leaves, usually narrow leaflets, com- 
pact terminal inflorescences, large yellow corol- 
las, and long linear cylindric fruits. The leaves are 
often almost glabrous, except for the minute 
round flat peltate trichomes and the stellate hairs 
in vein axils beneath. The hard wood has been 
used as uprights in home construction, as axles, 
and for tool handles (Englesing 181, Nicaragua). 
This species is very closely related to T. serrati- 
folia (Vahl) Nichols., a wide-ranging species of 
South America (Gentry, 1992). Common names 
are cartes or cortez. 

Tabebuia impetiginosa (Mart, ex DC.) Standl., 
Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 176. 
1936. Tecoma impetiginosa Mart, ex DC., 
Prodr. 9: 218. 1845. Tab. palmeri Rose, Con- 
trib. U.S. Natl. Herb. 1: 109. tab. 1 1. 1891. Tab. 
nicaraguensis Blake, Contrib. Gray Herb. 52: 
95. 1917. Tab. dugandii Standl., Trop. Woods 
36: 17. 1933. Figure 13. 

Trees to 30 m tall, trunks to 70 cm diam., bark 
slightly furrowed, leafy stems 3-8 mm diam., gla- 
brous or sparsely puberulent with minute (0.05 
mm) hairs, terete, longitudinally striate, becoming 
smooth and grayish. Leaves 3- or 5-foliolate (in 
Central America), petioles 4-14 cm long, 1.3-2.5 
mm diam., sparsely and minutely puberulent, lon- 
gitudinally striate, petiolules 7-50 mm long (basal 
laterals to 15 mm); leaflet blades (3-) 11 -19 cm 



BURGER: FLORA COSTARICENSIS 



153 



long, (2-)4-8 cm wide (basal leaflets often much 
smaller than distal), elliptic to elliptic-oblong or 
narrowly ovate-elliptic, apex short- to long-acu- 
minate, margin serrate in juvenile plants, base 
acute to obtuse or rounded, drying chartaceous, 
glabrous above, lower surface subglabrous or with 
short (0.4 mm) thin hairs along the midvein, and 
with tufts of hairs in the vein axils (domatia), 2 
veins 9-12/side. Inflorescences terminal, com- 
pact-paniculate (congested-fasciculate), peduncles 
2-12 mm long, 1-1.7 mm diam., densely yellow- 
ish pubescent with thick stellate hairs, pedicels 2- 
6 mm long. Flowers with calyx 5-8 mm long, 3- 
6 mm diam., tubular-campanulate, sparsely to 
densely yellowish stellate puberulent, margin 
truncate or slightly 5-lobed; corolla 35-75 mm 
long, tubular-funnelform, purple to red-purple 
(yellow within), sparsely to densely minutely pu- 
berulent externally, 15-25 mm wide at the mouth, 
lobes 10-17 mm long; filaments 16-23 and 10- 
16 mm long, thecae 2.5-3.5 mm long. Fruits 15- 
56 cm long, 12-25 mm wide, 10-13 mm thick, 
surface subglabrous and drying dark, with few 
longitudinal ridges; seeds 9-13 mm long, 30- 
48(-80) mm wide, central area 17-18 mm wide, 
wings translucent and clearly differentiated. 

Deciduous trees of seasonally very dry decid- 
uous or partly deciduous (rarely in lowland ev- 
ergreen) forest formations, 10-300 m elevation 
(to 1400 m elsewhere). Flowering in November- 
February; fruiting in January-February. This spe- 
cies ranges from Mexico to Argentina. 

Tabebuia impetiginosa is recognized by its tree 
habit, opposite palmately tri- or pentafoliolate 
leaves, compact terminal inflorescences, external- 
ly puberulent purple corollas, long linear fruits, 
and seeds with transparent wings. The leaflets are 
mostly glabrous. Gentry (1973b, 1992) discussed 
the regional differentiation and nomenclatural as- 
pects of this species. Common names are cortez 
negro and roble macho. Compare T. rosea 

Tabebuia ochracea (Cham.) Standl., Field Mus. 
Nat. Hist. Bot. Ser. 11: 176. 1936. Tecoma 
ochracea Cham., Linnaea 7: 653. 1832. Tab. 
chrysantha (Jacq.) Nichols., Diet. Gard. 4:1. 
1897, sensu Sandw., non Jacq. Tab. neochry- 
santha A. Gentry, Brittonia 22: 260. 1970. Tab. 
ochracea subsp. neochyrsantha (A. Gentry) A. 
Gentry, Ann. Missouri Bot. Gard. 60: 948. 
1973. Figure 13. 

Trees 8-25 m tall, bark with flat-surfaced gray 
ridges alternating with darker furrows, leafy stems 



3-7 mm diam., densely stellate pubescent with 
hairs 0.2-0.5 mm long, soon glabrescent and te- 
rete. Leaves 5-foliolate, petioles 4-18 cm long, 
1.3-2.7 mm diam., densely stellate pubescent, 
petiolules 5-58 mm long, lateral basal petiolules 
3-15 mm long; leaflet blades 4-22 cm long (bas- 
al laterals 2-11 cm), 2-14 cm wide, oblong-ob- 
ovate or elliptic-obovate to broadly elliptic, apex 
acuminate, margin serrate in juvenile plants, base 
obtuse to rounded and truncate, drying charta- 
ceous, densely stellate puberulent beneath with 
hairs ca. 0.5 mm wide, 2 veins 6-12/side. Inflo- 
rescences terminal, condensed subcapitate clus- 
ters of 2- and 3-flowered groups or contracted 
panicles, distal stems often terminating in a 
rounded pubescent inflorescence "bud" 1-2 cm 
diam., peduncles and pedicels not usually visible 
in the dense pubescence of yellowish (dried) hairs 
to 1.5 mm long. Flowers sweet-scented, calyx 8- 
13 mm long, 4-8 mm diam., campanulate, dense- 
ly yellowish (dried) tomentulous with straight 
hairs to 2(-7) mm long (stellate at base), margin 
with 5 lobes 1-2 mm long; corolla 36-75(-83) 
mm long, tubular-funnelform, yellow with red 
lines within, glabrous externally except near the 
lobes, lobes 10-25 mm long; filaments 15-20 and 
9-15 mm long, thecae 1.5-2.5 mm long. Fruits 
1 1-38 cm long, 10-17 mm wide, 5-10 mm thick, 
linear-cylindric, surface densely tomentulous with 
hairs 0.3-2 mm long (sometimes subglabrous and 
with prominent longitudinal ridges); seeds 6-12 
mm long, 17-88 mm wide, wings transparent, 
well differentiated. 

Common deciduous trees of seasonally dry de- 
ciduous and partly deciduous forest formations of 
the Pacific slope, 5-900 m elevation. Flowering 
in January-June; fruiting in February-June. This 
species is represented in our area by subspecies 
neochrysantha, which ranges from Guatemala to 
northwestern Venezuela; the other subspecies are 
South American. 

Tabebuia ochracea subspecies neochrysantha 
is recognized by its tree habit, opposite palmately 
pentafoliolate leaves, compact inflorescences (ear- 
ly stages form round pubescent "buds" at the tips 
of distal stems), bright yellow corollas mostly gla- 
brous externally, long cylindric fruits, and seeds 
with transparent wings. The leaflets are densely 
stellate tomentulose beneath. This species is sim- 
ilar to T. chrysantha of evergreen forests; both are 
called cones amarilla and cortez. 

Tabebuia palustris Hemsley, Biol. Centr. Am. 
Bot. 2: 495. 1882. Figure 12. 



154 



FIELDIANA: BOTANY 



Shrubs or small trees 1 .5-4 m tall, trunks to 5 
cm diam., often twisted and sometimes forming 
dense colonies, leafy stems 2-8 mm diam., terete, 
surface with flat rounded trichomes. Leaves sim- 
ple or 3-foliolate (2-foliolate), petioles 24-100 
mm long, 1.5-2.5 mm diam., with appressed pel- 
tate hairs but often appearing glabrous, often with 
gland fields along the proximal half, petiolules of 
distal leaves 12-35 mm long; leaflet blades 6-19 
cm long, 1.4-5 cm wide, narrowly lanceolate to 
narrowly elliptic or elliptic-oblong, apex acute, 
base acute to cuneate, drying yellowish and sub- 
coriaceous, glabrous above, densely lepidote with 
minute (0. 1 mm) appressed rounded trichomes be- 
neath, 2 veins 8-13/side. Inflorescences terminal 
on short shoots or terminal in a distal branch-di- 
chotomy, flowers 1-5, often in diads or triads, pe- 
duncle 8-25 mm long, bracts 2-8 mm long, linear, 
pedicels 9-11 mm long, with minute appressed 
rounded hairs. Flowers with calyx 11-17 mm 
long, 6-8 mm diam., cupulate-funnelform, drying 
dark, surface with flat rounded hairs ca. 0.1 mm 
diam., margin cleft into 2-4 lobes 1-5 mm long; 
corolla 40-75 mm long, tubular-funnelform, 
white with yellow lines within, glabrous external- 
ly, 9-18 mm wide at mouth, lobes 8-23 mm long; 
filaments 18-26 and 14-19 mm long, thecae 3 
mm long. Fruits 5-1 1 cm long, 14-26 mm diam., 
oblong-cylindric with a narrow tip, surface 
smooth, covered by minute peltate scales; seeds 
14-18 mm long, 18-22 mm wide, suborbicular 
with corky texture, wings absent. 

Shrubs of tidal mangrove formations and sea- 
side marshes along river estuaries, 0-10 m ele- 
vation. Flowering throughout the year. The north- 
ernmost collection (Croat 663 from Playa Coco, 
Guanacaste) may be an outlier. The estuary of the 
Rio Tempisque may be the northernmost area in 
which this species is common; it is common 
around Golfo Dulce. This species ranges along the 
Pacific Coast from Costa Rica to northwestern Ec- 
uador. 

Tabebuia palusiris is recognized by its Pacific 
coastal habitats, short shrubby (small tree) habit, 
opposite usually trifoliolate leaves with narrow 
leaflets, few white flowers, and cylindrical fruit 
with corky wingless seeds. The habitat, gland 
fields on petioles, narrow leaf blades, and corky 
seeds easily distinguish this species from all its 
congeners in Costa Rica. 

Tabebuia rosea (Bertol.) DC.. Prodr. 9: 215. 
1845. Tecoma rosea Bertol., Fl. Guatimal. 25. 
1840. Tec. mexicana Mart, ex DC., Prodr. 9: 



218. 1845. Sparattosperma rosea (Bertol.) 
Miers, Proc. R. Hort. Soc. 3: 99. 1863. Tub. 
mexicana (Mart, ex DC.) Hemsl., Biol. Centr. 
Am. Bot. 2: 495. 1882. Tab. pentaphylla (L.) 
Hemsl., Biol. Centr. Am. Bot. 2: 495. 1882, non 
Bignonia pentaphylla L. Couralia rosea (Ber- 
tol.) J. D. Smith, Bot. Gaz. 20: 9. 1895. Tec. 
evenia J. D. Smith, Bot. Gaz. 20: 8. 1895, pro 
parte. Figure 13. 

Trees 5-30 m tall, to 1 m diam., bark dark gray 
and rough with a distinct pattern of vertical ridg- 
es, leafy stems 3-12 mm diam., with minute pel- 
tate scales, glabrescent, becoming terete. Leaves 
5-foliolate, basal leaflets often smaller than distal, 
petioles 6-17(-32) cm long, 2-3.4 mm diam.. lep- 
idote, petiolules of distal leaflets 2-7(-ll) cm 
long; leaflet blades 5-18(-35) cm long, 3-10 
(-18) cm wide (distal leaflets larger), elliptic to 
elliptic-oblong, apex acute to acuminate, margin 
entire, base cuneate or rounded, drying stiffly 
chartaceous and grayish, both surfaces with flat 
appressed rounded hairs 0.05-0.1 mm diam. (lep- 
idote), gland fields sometimes present along the 
midvein near the base beneath, 2 veins 7-12/side. 
Inflorescences 2-20 cm long, short or condensed 
panicles often with dichotomous branching, pe- 
duncles 8-25 mm long, lepidote, bracts 1-2 mm 
long, pedicels 4-20 mm long, with minute peltate 
scales. Flowers with calyx 11-21 mm long, 6-1 1 
mm diam., cupular-funnelform, surface densely 
minutely papillate puberulent or lepidote, margin 
with 2 or 3 rounded or obtuse lobes separated by 
irregular sinuses 1-3 mm deep; corolla 53-97 
mm long, tubular-funnelform, pink to purple or 
white with pinkish lobes, often yellowish within, 
glabrous externally, lobes 18-26 mm long; fila- 
ments 14-20 and 10-15 mm long, thecae 2.5-3.5 
mm long. Fruits 21-38 cm long, 10-18 mm 
diam., linear-cylindric, densely covered with mi- 
nute peltate or papillate hairs, apex narrowly acu- 
minate; seeds 8-12 mm long, 28-45 mm wide, 
wings transparent and sharply differentiated. 

Trees of deciduous, partly deciduous, and low- 
land evergreen rain forest formations on both Ca- 
ribbean and Pacific coasts, 0-1100 m elevation 
(planted at elevations up to 1500 m). Flowering 
primarily in January-March; fruits usually dehisc- 
ing in April-May. This species ranges from Mex- 
ico to Venezuela and western Ecuador. 

Tabebuia rosea is recognized by its tree habit, 
opposite palmately pentafoliolate leaves, tubular 
and irregularly lobed calyx, large pink or white 
and pink corollas, linear-cylindric fruits, and seeds 



BURGER: FLORA COSTARICENSIS 



155 



with thin transparent wings. The leaf surfaces ap- 
pear glabrous but have a dense covering of minute 
appressed peltate hairs; some leaves have gland 
fields along the base of the midvein beneath. Un- 
like T. chrysantha, the leaflets of this species are 
always entire. This species is one of the most im- 
portant commercial sources of fine quality hard- 
wood; it is used in making furniture and interior 
trim and for other purposes. The tree flowers with 
or without its foliage and is one of Central Amer- 
ica's most beautiful native species, often planted 
for ornament. Common names are roble, roble de 
bianco, and roble de sabana. 



Tanaecium Swartz 

Lianas climbing with simple tendrils, stems te- 
rete, with 8 or 16 phloem areas in cross-section, 
interpetiolar gland fields present or absent; pseu- 
dostipules small or absent. Leaves opposite, 2- or 
3-foliolate, petiolate, blades often with a cyanide 
odor when crushed, venation pinnate. Inflores- 
cences axillary or terminal, panicles or racemose 
panicles, or solitary in distal leaf axils. Flowers 
with cupulate calyx, margin entire or with 5 mi- 
nute teeth, often with flat rounded glands on the 
distal surface; corolla with a very long tube and 
salverform distally, white, externally glabrous or 
puberulent, lobes 5; stamens 4, exserted or partly 
exserted, filaments of 2 lengths, anthers glabrous, 
thecae straight or curved, somewhat divergent, 
staminode present; ovary oblong, lepidote, 2-loc- 
ular, ovules multiseriate in each locule. Fruits 
capsules, oblong to ellipsoid-cylindric, valves 
parallel to the septum, thick, woody, convex, 
smooth; seeds lacking wings or with 2 lateral 
poorly differentiated membranaceous wings. 

Tanaecium is a genus of about six species rang- 
ing from Costa Rica and the West Indies to Brazil. 

Tanaecium jaroba Sw., Prodr. 92. 1788. T. al- 
biflora DC., Prodr. 9: 245. 1845. Figure 16. 

Lianas climbing with tendrils 5-22 cm long, 
leafy stems 2.5-7 mm diam., glabrous, terete, 
nodes with 10-15 round flat glands between the 
leaf bases; pseudostipules ca. 3 mm long or ab- 
sent. Leaves 2- or 3-foliolate, petioles 4-9 cm 
long, 1.5-2.3 mm diam., glabrous or puberulent, 
petiolules 7-37 mm long (shorter on lateral leaf- 
lets); leaflet blades 6.5-16 cm long, 3-9 cm wide, 
elliptic-oblong to elliptic or ovate-elliptic, apex 
acute to short-acuminate, base obtuse to rounded 
and somewhat truncate, glabrous above, glabrous 



or with curled thin hairs to 0.6 mm long beneath, 
2 veins 5-9/side. Inflorescences terminal or ax- 
illary, solitary flowers in distal leaf axils or short 
few-flowered racemes on leafless stem-tips, ped- 
icels 7-24 mm long, 1-2 mm diam., glabrous, 
drying black. Flowers with calyx 9-16 mm long, 
5-10 mm diam, deeply cupulate, subglabrous, 
with flat circular glands distally, margin subentire 
with minute (0.3 mm) teeth; corolla 1 1-24 cm 
long, long-tubular-salverform, white, minutely pu- 
berulent externally, tube 3.5-5 mm diam. for most 
of its length, ca. 10 mm wide at the throat, lobes 
17-27 mm long; filaments ca. 28 and 15 mm long, 
thecae 5-6 mm long. Fruits 9-22 cm long, 51 1 
cm wide, 4-8 cm thick, ellipsoid-oblong to ob- 
long, pale brown, subglabrous; seeds 25-33 mm 
long, 30-32 mm wide, woody, angular with thin 
edge on 3 sides. 

Rarely collected plants of evergreen lowland 
rain forest formations along the Caribbean coastal 
plain. Collected in flower in late May (Pittier 
3631) and early October (Gomez-Laurito 12913) 
in Costa Rica. This species ranges from south- 
eastern Nicaragua and Jamaica to Venezuela and 
Peru. 

Tanaecium jaroba is recognized by its climbing 
habit with simple tendrils, opposite bi- or trifoli- 
olate leaves, few-flowered inflorescences, very 
long slender white corollas, and rounded woody 
fruits with woody seeds. No other species of Big- 
noniaceae in Central America has such long and 
narrow corolla tubes. The flowers have a sweet 
odor (like Hedychium, Zingiberacae) and open at 
night. They are probably pollinated by sphingid 
moths with very long tongues. Little Central 
American material of this species has been seen; 
the description is based largely on South Ameri- 
can material and Gentry's (1973b) description. 



Tecoma Jussieu 

Shrubs or small trees, stems terete, interpetio- 
lar lines weakly differentiated at the node (glan- 
dular fields absent). Leaves opposite, simple, 3- 
foliolate or pinnately compound with opposite lat- 
eral leaflets and a terminal leaflet, petiolate, blades 
very variable in form, margins serrate, venation 
pinnate. Inflorescences terminal racemes or ra- 
cemose panicles, bracts small and inconspicuous, 
flowers pedicellate. Flowers showy, calyx cupu- 
late, glands often present on the distal surface, 
margin with 5 deltoid lobes; corolla tubular-cam- 
panulate to tubular-funnelform, radially symmet- 



156 



FIELDIANA: BOTANY 



ric, glabrous externally, yellow to orange, 5- 
lobed; stamens 4, exserted or included, filaments 
of 2 lengths, anthers glabrous or puberulent, the- 
cae divergent, pollen 3-colpate, a staminode pres- 
ent; disc cupular; ovary lepidote, 2-locular, ovules 
in 2 series in each locule, stigma flat. Fruits long 
linear capsules, compressed parallel to the septum 
but dehiscing perpendicular to the septum, valves 
smooth and glabrous; seeds thin, with 2 lateral 



membranaceous wings sharply differentiated from 
the central area of the seed. 

Tecoma is a genus of 14 species, with two in 
Africa and 12 in the New World (ranging from 
the southernmost United States to Argentina). 
Gentry (1992) placed Tecomaria under Tecoma. 
One species is both native and planted in gardens 
in Central America; the other is an introduced or- 
namental. 



Key to the Species of Tecoma 

la. Corollas yellow, tubular-campanulate; leaflet blades 3-14 cm long; fruits 9-25 cm long; wild and 
P lanted T. stans 

Ib. Corollas orange to red-orange, tubular; leaflet blades 1-4 cm long; fruits 5-12 cm long; planted 
ornamental T capensis 



Tecoma capensis (Thunb.) Lindley, Bot. Reg. 13: 
tab. 1117. 1827. Bignonia capensis Thunb., 
Prodr. 105. 1800. Tecomaria capensis (Thunb.) 
Spach. Hist. Nat. Veg. Phan. 9: 137. 1840. Fig- 
ure 15. 

Shrubs or scandent subshrubs 1-2 m tall, leafy 
stems 1.5-4 mm diam., usually terete, minutely 
(0.05-0.1 mm) puberulent. Leaves 4-14 cm long, 
pinnate with 7 or 9 (11) leaflets, petioles 10-18 
mm long, 0.5-1.2 mm diam., rachis sulcate above, 
minutely puberulent; leaflet blades 8-37 mm 
long, 8-16 mm wide, broadly ovate to ovate- 
rhombic or broadly ovate-elliptic, apex acute to 
obtuse, distal margin with 4-7 teeth/cm, base 
broadly cuneate or rounded, mostly glabrous but 
with thin white hairs in vein axils beneath (dom- 
atia), 2 veins 4-7/side. Inflorescences 7-15 cm 
long, racemes with 10-40 flowers, with short (4- 
9 mm) lateral branches, bracts 3-14 mm long, 
mostly linear, pedicels 6-12 mm long. Flowers 
with calyx 4-8 mm long, 3-4 mm diam., sub- 
glabrous, lobes 0.5-1 mm long, triangular with an 
apiculate apex; corolla 4-8 cm long, tubular, or- 
ange to orange-red, glabrous externally, tube 2-3 
mm diam. at base, 8-1 1 mm diam. at mouth, 
lobes 9-14 mm long; thecae 3 mm long. Fruits 
rarely produced in Central America, 5-12 cm 
long, 6-1 1 mm wide; seeds 5-6 mm long, 18-22 
mm wide. 

Tecoma capensis is planted as an ornamental in 
the tropics and subtropics; its vigorous growth 
and scandent branches make it useful in hedges. 
The opposite dark green leaves with 7-11 small 
serrate leaflets and brilliant curved-tubular or- 
ange-red corollas with exserted anthers make 



these plants easy to identify. Not often seen in 
Central America, these plants are likely to grow 
best at middle (1000-2000 m) elevations. The ge- 
nus Tecomaria with two African species is now 
considered part of Tecoma (Gentry, 1992). Julia 
and cape honeysuckle are common names. 

Tecoma stans (L.) Juss. ex Humboldt in H.B.K.. 
Nov. Gen. Sp. 3: 144. 1819. Bignonia stans L., 
Sp. PI. ed. 2, 2: 871. 1763. Figure 15. 

Shrubs or small trees l-6(-12) m tall, with 
many branches, trunk up to 25 cm diam. with 
ridged dark brown bark, leafy stems 2-5 mm 
diam., glabrous, drying pale yellowish gray. 
Leaves 8-22(-28) cm long, usually with 7, 9, or 
11 (1-5) leaflets, petioles 25-90 mm long, 0.8- 
1.7 mm diam., glabrous or puberulent at base of 
leaflets, sulcate above, petiolules of lateral leaflets 
0-2 mm long, terminal petiolules 6-18 mm long; 
leaflet blades 3-14 cm long, 1-6 cm wide (ter- 
minal leaflet larger than laterals), elliptic-ovate to 
narrowly elliptic-oblong or lanceolate, apex acu- 
minate, margin with 3 or 4 prominent teeth/cm, 
base cuneate and slightly decurrent on the petiole, 
lateral leaflets asymmetric at base, drying thin- 
chartaceous, glabrous (in ours), 2 veins 6-10/ 
side. Inflorescences 5-18 cm long, racemose or 
paniculate with 8-20 flowers, peduncles 1-3 cm 
long, glabrous, bracts ca. 1 mm long, triangular, 
pedicels 3-8 mm long, glabrous. Flowers with ca- 
lyx 4-7 mm long, 2.3-4 mm diam., cupulate, gla- 
brous, with submarginal glands, lobes 0.7-2 mm 
long, acute or apiculate; corolla 3.5-6 cm long, 
to 4 cm wide at the rotate lobes, tubular-campan- 
ulate from a narrow (2-3 mm diam.) base, yellow, 



BURGER: FLORA COSTARICENSIS 



157 



12-20 mm diam. at the mouth, lobes ca. 15 X 18 
mm, rounded; filaments ca. 16 and 21 mm long, 
thecae 3-4 mm long, pubescent. Fruits 9-22 
(-26) cm long, 5-8 mm wide, linear, cylindric or 
slightly flattened perpendicular to the septum, nar- 
rowed at base and apex, surfaces glabrous, pale 
brown; seeds 4-6.5 mm long, 17-28 mm wide, 
wings translucent, central area ca. 4 X 7 mm. 

Plants of deciduous and partly deciduous forest 
(or escaped from gardens in evergreen areas) of 
the Pacific slope and central highlands, 10- 
1300(-1800) m elevation. Flowering primarily in 
November-March; fruiting throughout the year. 
This species, originally ranging from southern- 
most Arizona and Florida to Argentina, is widely 
planted as an ornamental in the tropics and sub- 
tropics. 

Tecoma stans is recognized by its opposite im- 
paripinnately compound leaves with serrate leaf- 
lets, large bright yellow corollas that are only 
slightly bilaterally symmetric (obscurely 2- 
lipped), and long narrow seed pods with 2-winged 
seeds. This species is rather uniform in morphol- 
ogy in southern Central America. In Mexico and 
among cultivated forms there is extraordinary var- 
iation, with some plants having simple linear (25 
X 1 cm) leaves and others with the blades densely 
hirtellous beneath. This great range of variation 
has produced many specific and varietal names 
(see synonomy of Gentry, 1992). An often strag- 
gly growth habit prevents this species from being 
a more popular ornamental. Candelillo, carbon- 
cillo, and vainillo are common names. 



Tourrettia Fougeroux 

Herbaceous annual vines, climbing by means 
of tendrils with 3 coiled branches and dichoto- 
mous tips (transformed leaflets), stems tetragon- 
ous with 4 longitudinal ridges, an interpetiolar 
line or ridge usually present at the node, glandular 
fields absent; pseudostipules absent. Leaves op- 
posite, twice divided with 3 2 petioles each bear- 
ing 3-5 leaflets or with 2 2 petioles bearing 3-5 
leaflets and a distally branched tendril; blades 
with strongly serrate margin, lateral leaflets asym- 
metric and sometimes divided, thin, venation pin- 
nate. Inflorescences terminal spike-like racemes, 
sparsely to densely puberulent with thin gland- 
tipped hairs, bracts linear, pedicels short, the up- 
per flowers usually sterile and deciduous (with ca- 
lyx and corolla not clearly separated). Flowers of 
two kinds with the distal usually sterile and short- 
er than the lower (proximal) fertile flowers, calyx 



tubular near the base and separating into 2 lobes 
distally (upper and lower) in fertile flowers, de- 
ciduous during anthesis; corolla tubular and 2- 
lipped, slightly longer than the calyx, orange to 
red, puberulent distally; stamens 4, of 2 lengths, 
included, anthers glabrous, thecae divaricate, 
staminode absent; disc a thin cup; ovary ovoid, 4- 
locular, ovules many on uniseriate placentas, sur- 
face short-echinate, style slender, stigma simple. 
Fruits slightly woody capsules covered with 
straight and uncinate spines, dehiscing septicidal- 
ly into 2 valves united near the base; seeds small, 
oblong, with a thin narrow wing. 

Tourrettia contains a single species, ranging 
from the highlands of Guatemala to Argentina. 
The unusual fruits are reminiscent of the family 
Pedaliaceae, but the compound leaves, vining ten- 
drils, and winged seeds are characteristics of the 
Bignoniaceae. 

Tourrettia lappacea (L'Her.) Willd., Sp. PI. ed. 
4, 3: 263. 1801. Dombeya lappacea L'Her., 
Stirp. Nov. 33, tab 17. 1785. T. volubilis J. F. 
Gmel., Syst. Nat. 2: 940. 1791. Figure 16. 

Herbaceous vines to 3 m high, climbing with 
slender (0.3-0.7 mm) coiled tendrils 3-10 cm 
long, leafy sterns 0.8-5 mm diam., with 2 or 4 
prominent longitudinal ridges, subglabrous or 
sparsely puberulent with thin multicellular hairs 
0.2-0.5 mm long (especially at the nodes). Leaves 
opposite, to 25 cm long, 1 petiole 3-5 cm long, 
1.5-3.3 mm diam., usually glabrous, 2 petioles 
2-6 cm long bearing 3-5 leaflets, petiolules of 
central leaflet 8-18 mm long; leaflet blades 0.8- 
10 cm long, the central leaflet 2-8(-10) cm long, 
1.3-5 cm wide, ovate to ovate-elliptic, apex acute, 
margin strongly serrate with teeth 0.3-3 mm high 
(4-9 teeth/cm), drying thin and dull greenish, gla- 
brous above and below, 2 veins 6-13/side. Inflo- 
rescences to 20 cm long, with both sterile distal 
flowers (5-10 mm long) and fertile (to 2 cm long) 
usually proximal fertile flowers, peduncles 3-12 
cm long, subglabrous to densely pubescent with 
gland-tipped hairs, pedicels 0.8-3.5 mm long, ca. 
0.3 mm diam., with a linear bract to 4 mm long 
from near the base. Flowers dimorphic, distal ca- 
lyx 4-7 mm long and reddish, fertile calyx 11- 
15 mm long, greenish, lobes slightly longer than 
the tube, subglabrous to densely glandular puber- 
ulent; corolla of fertile flowers 12-24 mm long, 
3-4 mm diam. near the base, greenish to orange, 
rose red or deep scarlet red, puberulent except 
near the base, upper lobe 8-12 mm long, hooded 
(galeate); filaments 8-9 and 9-12 mm long, an- 



158 



FIELDLANA: BOTANY 



ther thecae 2-3 mm long; style to 16 mm long. 
Fruits 3-5 cm long, body of the fruit ca. 3 X 1 .4 
cm, ellipsoid, surface covered with longer (7-14 
mm) uncinate reddish spines and shorter (1-3 
mm) acute spines, drying dark brown; seeds 6-7 
mm long, 4-5 mm wide, flat. 

Plants of moist open sites and forest edges in 
evergreen lower montane forest formations, 
(600-)1100-2100 m elevation. Flowering and 
fruiting throughout the year. This species ranges 
from Mexico to Venezuela, Peru, and northwest- 
ern Argentina. 

Tourrettia lappacea is a very unusual herba- 
ceous vine with slender divided coiling tendrils, 
twice-compound opposite leaves, strongly serrate 
thin blades, and colorful spike-like inflorescences 
usually bearing flowers of two different forms. 
The sterile (usually terminal) flowers are shorter 
and have the calyx and corolla often united, while 
the larger fertile flowers have the calyx clearly 
differentiated and deciduous. In addition, the two- 
valved fruits covered with both shorter straight 
spines and longer hooked spines are distinctive. 
Because of the slender tendril-bearing stems and 
spiny fruits, these plants are sometimes mistaken 
for Cucurbitaceae (see Gentry, 1980). 



Tynanthus Miers 

Lianas climbing with the aid of simple or trifld 
tendrils (rarely small trees?), stems subterete to 
quadrangular with 4 phloem areas in cross-sec- 



tion, often with an interpetiolar ridge at the nodes 
but interpetiolar gland fields lacking; pseudostip- 
ules lacking or leaf-like. Leaves opposite. 2- or 
3-foliolate, often with the terminal leaflet replaced 
by a simple or distally trifid tendril, blades entire, 
venation pinnate. Inflorescences terminal or ax- 
illary, usually small (4-12 cm) open or condensed 
panicles or dichasia, pedicels densely puberulent. 
Flowers with small cupular or campanulate calyx, 
puberulent, apex subtruncate with 5 denticulate 
lobes or setae, persisting; corolla funnelform and 
somewhat curved, bilabiate and split to near the 
middle, white, puberulent on the exterior, with 4 
or 5 lobes (upper lobe emarginate or 2-lobed); sta- 
mens 4, of 2 lengths, included or slightly exserted. 
anthers glabrous, thecae divaricate, staminode 
present; disc very small; ovary conical, densely 
puberulent, 2-locular, ovules in 2-4 series on the 
placenta. Fruits long-linear capsules, slightly flat- 
tened with valves parallel to the septum, flat and 
smooth with a slightly or distinctly raised 
(winged) margin, dehiscing septicidally; seeds 
flat, with 2 lateral hyaline wings. 

Tynanthus is a genus of about 1 2 species rang- 
ing disjunctly from southern Mexico and the West 
Indies to Brazil and Bolivia. The genus has a 
number of species with unusually small flowers 
for Bignoniaceae; for example, the corollas of T. 
guatemalensis J. D. Smith are only 6-9 mm long. 
Our species, based on only two collections, is the 
only representative of the genus known from be- 
tween Guatemala and Central Panama; for com- 
parison we include a key to the three species of 
the genus known in Central America. 



Key to the Central American Species of Tynanthus 

la. Corollas 25-30 mm long; small trees (?) without tendrils; lower leaf surfaces grayish with a dense 
covering of minute scurfy hairs; known only from the Caribbean slope of Costa Rica at 100-800 
m elevation T. macron thus 

Ib. Corollas 6-20 mm long; lianas climbing with tendrils; lower leaf surfaces greenish, glabrous or 
with minute hairs along the veins; Panama or Mexico to Guatemala and not included in the de- 
scriptions 2 

2a. Corollas 6-9 mm long; leaves mostly elliptic to ovate-elliptic; ranging from central Mexico to Belize 
and Guatemala T. guatemalensis 

2b. Corollas 12-20 mm long; leaves mostly ovate to broadly ovate; central to eastern Panama 

T. croatianus 



Tynanthus macranthus L. O. Williams, Fieldi- 
ana Bot. 31: 250. 1967. Figure 12. 

Small trees 3 m tall (in the type) but also lia- 
nas, leafy stems 1.5-5 mm diam., densely grayish 



puberulent with minute (ca. 0.1 mm) rounded, 
peltate or scurfy hairs, terete, node with a slightly 
elevated interpetiolar ridge. Leaves 2-foliolatc, 
petioles 8-17 mm long, ca. 1 mm diam., densely 
puberulent like the stems, petiolules 6-1 1 mm 



BURGER: FLORA COSTARICENSIS 



159 



long, sulcate above; leaf blades 4-12 cm long, 2- 
6 cm wide, elliptic-oblong to oblong, apex acute 
to abruptly short-acuminate, base asymmetric 
with narrowed and slightly rounded sides, drying 
stiffly chartaceous and brown above, upper sur- 
face minutely papillate puberulent or subglabrous, 
lower surface grayish to brownish and densely to 
sparsely puberulent with whitish hairs 0.05-0.1 
mm long, 2 veins 3-6/side. Inflorescences axil- 
lary, 1-3 cm long, few-flowered racemes or di- 
chasia, peduncle 10-16 mm long, 1 mm diam., 
densely puberulent, bracts 1-2 mm long, cadu- 
cous, pedicels 3-6 mm long. Flowers with calyx 
6-9 mm long, 4-5 mm diam., tubular-campanu- 
late, densely pale brownish puberulent (dried), 
margin subentire with 5 minute (0.3 mm) teeth; 
corolla 22-23 mm long, tubular-funnelform, 2- 
lipped and split along the sides to within 5-8 mm 
of the base, white, densely pale yellowish brown 
puberulent on the exterior when dried, tube 2-4 
mm diam. near the base, 5-8 mm diam. at the 
mouth (lower lip becoming bent downward and 
recurved in Barringer et al. 2671), lobes 5-12 
mm long, obtuse at the apex; thecae ca. 2 mm 
long, the pair U-shaped when dried; ovary mi- 
nutely puberulent. Fruits unknown. 

Rarely collected plants of wet evergreen rain 
forest formations of the Caribbean slope, 100-800 
m elevation. Flowering in late April and early 
July. Endemic to Costa Rica and northwestern 
Panama. 

Tynanthus macranthus is recognized by its op- 
posite bifoliolate leaves with blades usually gray- 
ish puberulent beneath, the short axillary racemes, 
the densely puberulous calyx with minutely den- 
tate margin, and the densely puberulous white co- 
rollas deeply split along the two sides. Although 
the type (Lent 42) was said to be a tree, other 
collections appear to be lianas. The flowers of this 
species, while modest in size for the family, are 
large for the genus. No other bignon in our flora 
has a corolla tube so deeply incised. 



Xylophragma Sprague 

Lianas climbing with simple tendrils (undivid- 
ed distally), stems terete or subquadrangular, with 
4 phloem areas in cross-section, interpetiolar 
glandular fields present; pseudostipules short, 
acute. Leaves opposite, 3-foliolate or 2-foliolate 
with the terminal leaflet sometimes replaced by a 
tendril, pubescence of small simple or branched 
(dendritic) hairs, venation pinnate. Inflorescences 



racemes or panicles with racemose branches, usu- 
ally axillary on older branchlets, bracts present, 
pedicels often with paired bracteoles. Flowers 
with cupular calyx, margin with 5 short teeth, pu- 
bescent with stellate or branched hairs; corolla 
tubular-campanulate, slightly bilabiate, rose to 
lavender or purple, puberulent on the exterior; sta- 
mens 4, of 2 lengths, anthers glabrous, thecae 
straight and divaricate, a staminode present; disc 
cupular; ovary ovoid, 2-locular, ovules 6-8 series 
on each placenta, stigma simple. Fruits capsules, 
valves woody, flat, and parallel to the septum, me- 
dian vein not evident, smooth or with raised glan- 
dular areas; seeds thin with 2 lateral membrana- 
ceous wings clearly delineated from the central 
seed area. 

Xylophragma is a genus of four species cen- 
tered in South America, with one species ranging 
northward as far as Mexico. 

Xylophragma seemannianum (Kuntze) Sandw., 
Kew Bull. 1953: 469. 1954. Saldanhaea see- 
manniana Kuntze, Rev. Gen. PI. 2: 480. 1891. 
Distictis rovirosana J. D. Smith, Bot. Gaz. 20: 
7. 1895. Adenocalymma cocleense Pittier, Con- 
trib. U.S. Natl. Herb. 18: 255. 1917. Figure 21. 

Lianas stems to 10 cm diam., tendrils to 20 cm 
long, leafy stems 2-6 mm diam., at first puberu- 
lent with branched or stellate hairs ca. 0.2 mm 
long, glabrescent, terete, developing whitish len- 
ticels, gland fields often present at the nodes; 
pseudostipules 1-3 mm long, conical. Leaves 3- 
foliolate or 2-foliolate with a tendril, petioles 4- 
16(-20) cm long, 0.7-2.2 mm diam., minutely pu- 
berulent with straight or crooked hairs 0.1-0.3 
mm long, petiolules 6-40(-70) mm long, petio- 
lules of central leaf often twice the length of lat- 
eral petiolules; leaflet blades (4-)6-16(-23) cm 
long, (2.5-)4-10(-13) cm wide, ovate, broadly el- 
liptic, obovate or slightly rhombic, apex acute to 
acuminate, base obtuse to slightly rounded and 
truncated, drying chartaceous, minutely puberu- 
lent on the veins above, usually puberulent be- 
neath with branched hairs 0.2-0.5 mm long be- 
neath, 2 veins 5-7/side. Inflorescences terminal 
on axillary short-shoots, racemes or short pani- 
cles, peduncles 5-20 mm long, ca. 1 mm diam., 
densely stellate puberulent, bracts 3-6 mm long, 
linear to linear-lanceolate, pedicels 5-15 mm 
long. Flowers with calyx 4-8 mm long, 3-5 mm 
diam., densely papillate or stellate puberulent, 
teeth (lobes) 0.2-1 mm long, ca. 0.5 mm wide; 
corolla (2.5-)3-6 cm long, tubular-funnelform, 



160 



FIELDIANA: BOTANY 



lavender to rose and minutely puberulent on the 
exterior, yellow and white within, tube 1.5-2 mm 
diam. near the base, 10-18 mm wide at the 
mouth, lobes to 22 X 16 mm and acute; filaments 
ca. 9 and 14 mm long, thecae 2.3-3.5 mm long; 
ovary ca. 3 mm long, lepidote. Fruits 5-16 cm 
long, 3-5 cm wide, ca. 5 mm thick, oblong-round- 
ed, valves flat and smooth; seeds 14-22 
(-29) mm long, 28-52 mm wide, central dark area 
17-24 mm wide. 

Common deciduous lianas of seasonally dry 
deciduous and partly deciduous forest formations 
(rarely collected in evergreen forest formations), 
1-500 m elevation. Flowering in March-May 
when the plants are usually leafless; fruits mature 
and open in the following year's dry season. This 
species ranges from Veracruz, Mexico, to north- 
ern Brazil. 

Xylophragma seemannianum is recognized by 
its climbing habit with simple woody tendrils 
(rarely shrubs), the usually trifoliolate leaves, 
short inflorescences usually blooming when the 
leaves are absent, showy lavender or magenta co- 
rollas, and oblong flattened fruits with two- 
winged seeds. The gland fields at older nodes, 
larger leaves with symmetric lateral leaflets, and 
pubescence of small branched hairs (dendroid, of- 
ten appearing farinose) are also helpful in recog- 
nizing this species. 

List of Accepted Species of Bignoniaceae 

Key: END-CR = endemic to continental Costa 
Rica; END-CR&WP = endemic to Costa Rica 
and western Panama; END-WP = endemic to 
western Panama; INTRO = introduced weed; 
ORNAM = cultivated ornamental; ORNAM & 
NAT = cultivated and naturalized; ?? = not col- 
lected in Costa Rica but known from nearby ar- 
eas. Total number of species covered is 79; the 
number of documented native species is 69. One 
new species, Amphitecna gentryi, is described 
here. 

Adenocalymma inundatum 

Amphilophium paniculatum 

Amphilophium pannosum 

Amphitecna gentryi END-CR & SP. NOV. 

Amphitecna isthmica 

Amphitecna kennedyi 

Amphitecna latifolia 

Amphitecna sessilifolia END-CR & WP 

Anemopaegma chrysanthum 

Anemopaegma chrysoleucum 

Anemopaegma orbiculatum 

Anemopaegma puberulum 



Anemopaegma santaritense 
Arrabidaea candicans 
Arrabidaea chica 
Arrabidaea conjugata 
Arrabidaea corallina 
A rrabidaea costaricensis 
Arrabidaea florida 
A rrabidaea mollissimu 
Arrabidaea patellifera 
Arrabidaea pubescens ?? 
Arrabidaea verrucosa 

Callichlamyx latifolia 
Ceratophytum tetragonolobum 
Clytostoma binatum 
Crescentia alata 
Crescentia cujete 
Cydista aequinoctialis 
Cydista diversifolia 
Cydista heterophylla 
Cydista lilacina 
Cydista potosina 

Distictella magnoliifolia 
Godmania aesculifolia 

Jacaranda caucana 

Jacaranda copaia 

Jacaranda mimosifolia ORNAM 

Kigelia pinnata ORNAM 

Lundia corymbifera 
Lundia puberula 

Macfadyena uncata 
Macfadyena unguis-cati 
Mansoa hymenaea 
Mansoa kerere 
Mansoa pan'ifolia 
Mansoa standleyi 
Mansoa verrucifera 
Martinella obovata 
Melloa quadrivalvis 
Mussatia hyacinthina 

Parabignonia steyermarkii 
Paragonia pyramidata 
Parmentiera aculeata ORNAM 
Parmentiera cereifera ORNAM 
Parmentiera dressleri 
Parmentiera macropltylla 
Parmentiera valerii END-CR 
Phyrganocydia < -orymhoxa 
Phryganocydia phellosperma 
Pithecoctenium crucigerum 
Pleonotoma variabilis 
Podranea ricolaaiana ORNAM 
Pyrostegia venusta ORNAM 

Spathodea campanulata ORNAM 
Stizophyllum inaequilaterum 
Sti-ophyllum riparium 

Tabebuia chrysantha 
Tahebuia guayacan 
Tabebuia impetiginosa 
Tabebuia ochracea 
Tabebuia palustris 
Tabebuia rosea 



BURGER: FLORA COSTARICENSIS 



161 



Tanned urn jaroba 

Tecoma capensis ORNAM 

Tecomu stems ORNAM & native 

Tourretliei lappacea 

Tynunllnis meicreinthtis END-CR & WP 

Xyliiphreignui seemannianum 



PEDALIACEAE 

By William Burger 

Herbs or more rarely shrubs or small trees, 
stems erect, often with mucilaginous glands that 
become slimy when wet; stipules absent. Leaves 
opposite or the distal alternate, petiolate, simple, 
entire or lobed, pinnately or palmately veined. In- 
florescences of usually solitary axillary flowers, 
sometimes cymose, usually with glands at the 
base of the pedicel. Flowers bisexual, bilaterally 
symmetric, sepals 5, united at the base or free; 
corolla tubular to campanulate and slightly 2- 
lipped, 5-lobed, imbricate in bud; stamens 4, fil- 
aments of 2 lengths, anthers dorsifixed, parallel or 
separate, a staminode usually present; disc present 
on 1 side of the ovary; ovary superior (rarely in- 
ferior), 2- or 4-locular, placentation axile, ovules 
few to many, style slender, stigma 2-lobed. Fruits 
capsules or indehiscent, often woody and spiny or 
with projections; seeds few to many, smooth, en- 
dosperm thin. 

Pedaliaceae is an Old World family of about 1 5 
genera and 50 species. Closely related to Scro- 
phulariaceae, this family is distinguished by its 
slime cells and unusual fruits. Some authors have 
enlarged this family to include Martyniaceae 
(Cronquist, 1981), but that family has parietal pla- 
centation and is restricted to the New World. The 
only representative of this family likely to be seen 
in Central America is Sesamum orientate, the 
source of sesame seed. 



Sesamum Linnaeus 

Annual or perennial herbs, stems erect, usually 
with mucilaginous sap or glands. Leaves opposite 
or alternate, petiolate, blades with pinnate vena- 
tion. Inflorescences of usually solitary flowers in 
leaf axils or sometimes in few-flowered cymes, 
pedicels subtended by lateral glands. Flowers 
with 5-parted calyx; corolla tube oblique at the 
base and slightly expanded (gibbous), slightly 2- 



lipped with 5 spreading lobes; stamens 4 (rarely 
2), inserted near the base of the corolla tube, an- 
thers divergent (sagittate); ovary superior or part- 
ly inferior, 2-locular with many ovules. Fruits 
capsules, oblong, somewhat 4-angled in cross- 
section, loculicidally dehiscent. 

Sesamum is a tropical genus of about 12 spe- 
cies, native to Africa and Asia. The following spe- 
cies has been widely cultivated in tropical regions 
for its oil-containing seeds. 

Sesamum orientate L., Sp. PI. 634. 1753. S. in- 
dicum L., Sp. PI. 634. 1753. Figure 27. 

Erect annual herbs to ca. 1 .5 m tall, stems sim- 
ple or branched, leafy stems 1-5 mm diam., with 
thin whitish hairs 0.2-1.5 mm long; bracts may 
resemble stipules at flowering nodes. Leaves 
mostly alternate, petioles 6-35(-80) mm long, 
0.5-1. 3(-2) mm diam., puberulent with thin hairs 
ca. 0.5 mm long; leaf blades 3-15 cm long, 0.7- 
9 cm wide, varying from broadly ovate-triangular 
(or deeply 3-lobed in basal leaves) to lanceolate 
or narrowly oblong, apex acute to obtuse, margin 
entire (sometimes with rounded lobes), base acute 
to obtuse, drying chartaceous, sparsely puberulent 
with thin hairs on both surfaces, 2 veins 5-6/side 
and strongly ascending. Inflorescences of solitary 
axillary flowers, bracts to 4 mm long, linear, ped- 
icels 1-2 mm long (to 5 mm in fruit), with a pair 
of lateral sessile round glands (0.7 mm diam.) at 
the base, puberulent. Flowers with calyx 4.5-6.5 
mm long, sepals united only at the base, ca. 5 mm 
long, narrowly triangular, acute, with thin white 
hairs; corolla 20-30 mm long, white to pink or 
lavender, sparsely puberulent with thin hairs 0.3- 
1 .5 mm long, lobes 2-7 mm long, broadly round- 
ed. Fruits 17-30 mm long, 8-10 mm wide, ob- 
long-rectangular, base rounded, apex acute or 
apiculate, with a central longitudinal sulcus on the 
nondehiscent sides, pale brown, puberulent; seeds 
3 X 1 .8 X 1 mm, ovate-lenticular, lustrous white. 

Cultivated or occasionally escaped plants in 
seasonally dry deciduous areas, 100-800 m ele- 
vation in Central America. Flowering in June-De- 
cember; fruiting in July-January. Rarely encoun- 
tered in Costa Rica, this species is naturalized in 
parts of Guatemala and Honduras. Cultivated 
since ancient times in southern and western Asia, 
it has been introduced throughout the tropics. 

Sesamum orientale is recognized by its solitary 
axillary flowers subtended by glands at the base 
of the pedicels, bent campanulate white or pink 
corollas, and the distinctive oblong capsules. 



162 



FIELDIANA: BOTANY 




Redrawn 

froK 

S.Troyo 



Tetranema florlbundum 

FIG. 27. Unusual herbs of the Martyniaccae, Pedaliaccac, and Scrophulariaccae families. 



BURGER: FLORA COSTARICENSIS 



163 



Ajonjoli and sesame are common names. The oil 
expressed from the seeds is used much like olive 
oil, and the seeds are used on bread or in making 
sweets. 



MARTYNIACEAE 

By William Burger 

Herbs, usually with viscid gland-tipped multi- 
cellular hairs; stipules absent. Leaves alternate or 
opposite, simple, petiolate, leaf blades entire to 
undulate, dentate or somewhat lobed. Inflores- 
cences terminal or axillary, racemose, flowers 
pedicellate, often subtended by 1 or 2 large brac- 
teoles, sometimes becoming thick and fleshy in 
age. Flowers bisexual, calyx 5-parted or 5-lobed 
and split down 1 side, glandular puberulent exter- 
nally; corolla bilaterally symmetric and somewhat 
2-lipped, tubular at base and expanded to cam- 
panulate or funnelform distally, 5-lobed with the 
2 upper lobes exterior in bud; stamens 4 or 2, 
filaments equal or of 2 lengths, arising from the 
base of the expanded corolla tube, anthers of each 
stamen pair at first coherent, 2-thecous, stami- 
nodes 1-3; disc annular; ovary superior, unilocu- 
lar but with 2 intruding parietal placentas and ap- 
pearing 2- or 4-locular by intrusion of the placen- 
tas, ovules few to many, style slender, stigma 2- 
lobed. Fruits woody capsules, usually with distal 
recurved horns, exocarp fleshy and deciduous, en- 
docarp hard and woody; seeds few to many, 
sculptured, black, compressed or oblong. 

This family of three genera is restricted to the 
New World, but some authors place it in the Pe- 
daliaceae (Cronquist, 1981). The Martyniaceae 
differ from the Pedaliaceae in having intruding 
parietal placentae and lacking glands at the base 
of the pedicels, and they do not become slimy 
when wet. The monotypic genus Martynia is rare- 
ly collected in Costa Rica. A second genus and 
species, Proboscidia triloba (Cham. & Schldl.) 
Decne., is found in northern Central America. 
That genus is distinguished by having a spathe- 
like calyx, four fertile stamens, and distal horns 
of the capsule usually surpassing the body of the 
fruit in size. 



Martynia Linnaeus 

Erect annual herbs, puberulent to villous with 
thin multicellular gland-tipped viscid hairs on 



most parts. Leaves opposite or alternate distally, 
petiolate, blade broadly ovate with palmate ve- 
nation. Inflorescences terminal or axillary, short- 
racemose, viscid puberulent, flowers subtended by 
2 large bracts. Flowers with deeply 5-parted ca- 
lyx, sepals unequal, thin-textured; corolla cam- 
panulate from a short tubular base, somewhat 2- 
lipped and oblique, with 5 broadly rounded lobes; 
stamens 2, included, filaments borne from the 
apex of the short narrow basal tube, anthers 
strongly divaricate (180) and coherent on their 
sides, staminodes 2 or 3; ovary with 2 intruding 
parietal placentae, stigma 2-lobed. Fruits a 
horned capsule, exocarp glandular pubescent but 
deciduous and exposing the hard woody exocarp, 
adaxial and abaxial sides of the exocarp each with 
4 longitudinal ribs separated by deep sulci, the 
hard recurved distal horns shorter than the body 
of the fruit. 

Martynia annua L., Sp. PI. 618. 1753, non L. 
Syst. Nat. 1113. 1759. M. diandra Gloxin, Obs. 
Bot. 14. 1785. M. angulosa Lam., Encycl. 
Meth. Bot. 2: 112. 1786. Carpoceras angulata 
A. Rich., Bull. Sci. Nat. Geol. 21: 98. 1830. 
Disteira angulosa Raf., Fl. Telluriana: 68. 1881. 
Vatkea diandra O. Hoffm., Verh. Bot. Bran- 
denb. 73: 45. 1881; Linnaea 43: 554. 1882. Fig- 
ure 27. 

Annual erect herbs 0.3-1.5 m tall, stems 
sparsely to densely viscid pubescent with hairs ca. 
0.5 mm long. Leaves with petioles 4-14(-40) cm 
long, 1.4-5 mm diam., villous with gland-tipped 
hairs 0.3-1 mm long; leaf blades 7-24(-40) cm 
long, 5-21 (-32) cm wide, ovate to ovate-trian- 
gular (sometimes weakly 3- or 5-lobed), apex ob- 
tuse, margin with widely triangular or obtuse 
lobes and more numerous minute (0.5 mm) teeth, 
base cordate to subcordate, drying membrana- 
ceous, upper surface with scattered short hairs, 
lower surface with hairs and pellucid dots, vena- 
tion palmate, 2 veins 3-4/side. Inflorescences 3- 
6 cm long, terminal or axillary to distal leaves, 
with 5-15(-20) flowers bracts 12-23 mm long, 
broadly elliptic and somewhat similar to the se- 
pals, pedicles 12-28 mm long, slender, glandular 
pubescent. Flowers with 5 unequal sepals 16-20 
mm long, 4-7 mm wide, oblong to broadly ellip- 
tic or oblanceolate, obtuse or rounded at the apex, 
ciliolate on the edge; corolla 40-55 mm long, 
white or pinkish white with rounded purple spots 
on the sepal lobes within and yellow area on the 
floor of the throat, lobes broadly rounded distally, 



164 



FIELDIANA: BOTANY 



the 2 smaller (adaxial) lobes held erect, the broad 
abaxial lobe forming the lower lip; anthers ca. 8 
mm wide, filaments ca. 12 mm long; style ca. 30 
mm long. Fruits 3-4 cm long, 1.6-2 cm wide, 
slightly curved and boat-shaped (curved ellip- 
soid), the distal ends with hard recurved sharp- 
tipped horns ca. 1 cm long, woody endocarp with 
deep sulci and drying black. 

Plants of seasonally dry deciduous forest areas 
5-300 m elevation (to 1000 m in Honduras, to 
2400 m in Guatemala). Flowering in July-August; 
fruiting in August-February. Although common 
in central Honduras and along the shore of Lake 
Nicaragua near Granada, this species has been 
collected only a few times in Guanacaste Prov- 
ince. The species ranges from Mexico to north- 
eastern Costa Rica. 

Martynia annua is recognized by its short-lived 
herbaceous habit, viscid pubescence, broad pal- 
mately veined leaves, racemose inflorescences, 
five large subequal sepals, and whitish slightly 
two lipped corolla with purple spots. The mature 
woody fruits are especially distinctive with their 
two distal recurved sharp-tipped horns. The some- 
what coherent anthers are reminiscent of those 
seen in Gesneriaceae. Unas de diablo is a com- 
mon name. 



OROBANCHACEAE 

By Luis D. Gomez P. and William Burger 

REFERENCES L. D. G6mez, Notes on the bi- 
ology of Central American Orobanchaceae. Bre- 
nesia 17: 389-396. 1980. G. B. von Mannagetta, 
Orobanchaceae, in A. Engler, Pflanzenreich IV, 
261: 1-348. 1930. J. Thieret, The Genera of Oro- 
banchaceae in the Southeastern United States. J. 
Arnold Arbor. 52: 404-434. 1971. 



Annual or perennial herbs, all parts lacking 
chlorophyll, parasitic on the roots of other plants, 
arising from a thickened base, stems simple or 
few-branched, thin to thick, usually succulent, 
glabrous to pubescent, hairs simple or glandular; 
stipules absent. Leaves alternate and simple, re- 
duced to sessile scales, without chlorophyll, mar- 
gins entire. Inflorescences simple or less often 
branched, flowers in racemes or spikes (rarely sol- 
itary and terminal), flowers solitary in the axils of 
bracts, sessile or pedicellate, bracteoles usually 
present. Flowers bisexual, calyx radially or bilat- 
erally symmetric, tubular or campanulate with 2- 
5 unequal lobes or teeth, sometimes split on 1 or 
2 sides, open or valvate in bud; corolla tubular to 
campanulate and usually 2-lipped, 4- or 5-lobed. 
tube curved or straight, upper lip 2-lobed or en- 
tire; stamens 4, included or exserted, alternate 
with the corolla lobes, filaments 2 pairs of unequal 
length, borne on the proximal half of the corolla 
tube, free, anthers free, dorsifixed, 2 thecous with 
1 or 2 thecae fertile, free or coherent, opening by 
a longitudinal slit, a staminode present or absent; 
ovary superior, 1-locular with 4 (2-6) intruded pa- 
rietal placentae, ovules many and anatropous, 
style simple, stigma simple or 2-4-lobed. Fruits 
capsules, dehiscing loculicidally into 2 or 3 
valves; seeds many, small, with ornamented testa, 
embryo undifferentiated. 

A family of ca. .17 genera and 200 species 
(Mabberley, 1987), in temperate and subtropical 
climates throughout the northern hemisphere, but 
with the greatest number of species in the Old 
World. The lack of chlorophyll, usually simple or 
few-branched stems, tubular two-lipped corollas, 
unilocular ovary with intruding parietal placentae, 
and capsular fruits make these plants easy to iden- 
tify. The aboveground parts of these plants are 
short-lived and are easily missed by collectors. 
One genus is native to Central America and one 
may be found as an introduced weed. This family 
is closely related to Scrophulariaceae. 



Key to the Genera of Orobanchaceae 

la. Scales overlapping on the lower stem; corolla glabrous externally; anthers slightly exserted; parasites 
in high-elevation oak forests Conopholis 

Ib. Scales not overlapping on the lower stems; corolla puberulent externally; anthers included; rarely 
collected parasites at higher elevations in open fields Orobanche 



BURGER: FLORA COSTARICENSIS 



165 



Conopholis Wallroth 

REFERENCE R. Haynes, A monograph of the 
genus Conopholis (Orobanchaceae). Sida 4: 246- 
264. 1971. 

Herbs with short erect unbranched stems, par- 
asitic on oak (Quercus) roots, aerial parts of 1 or 
several annual flowering stems that are covered 
by imbricate scales near the base, stems arising 
from a gall-like perennial base, simple or rarely 
branched, glabrous to densely glandular-pubes- 
cent. Leaves represented by broad-based sessile 
scales that are imbricate at the base and alternate 
distally, yellowish white at first and becoming 
brown, intergrading with the very similar bracts. 
Inflorescences spike-like racemes with thick axis, 
flowers separate or crowded, bracts exceeding the 
calyx in length, sessile or short-pedicellate, 1 or 
2 small bracteoles arising from the base of the 
calyx. Flowers with tubular or spathe-like calyx, 
cleft on one side, irregularly 2-, 4-, or 5-lobed or 
dentate on the distal margin, divisions acute or 
rounded; corolla tubular and 2-lipped, yellowish 
white, the upper lip curved or straight, the lower 
lip with (1 or 2) 3 subequal spreading lobes; sta- 
mens 4, exserted, anthers with spurs, thecae 
slightly divergent, glabrous or sparsely pilose; 
ovary unilocular with 4 parietal placentas, stigma 
discoidal. Fruits 2-valved capsules, usually en- 
closed by the persisting perianth, dehiscing irreg- 
ularly; seeds oval to angular, testa reticulate. 

Conopholis is a North and Central American 
genus of two species, according to Haynes (1971, 
reference above). A brief overview, however, 
makes it appear that the collections of Conopholis 
could also be interpreted as representing either a 
single variable species or several species. The 
main characteristics used by Haynes to distinguish 
the two species, apart from geographic separation, 
are quite variable, even within a single popula- 
tion. Earlier, Standley (1938) placed the Costa Ri- 
can material under the name C. americana (L.) 
Wallroth. The genus is distinctive because of its 
lack of green pigmentation, parasitic association 
with oak trees, short thick succulent spike-like 
stems with imbricate bract-like scales, many dis- 
tally crowded flowers, and pale yellowish tubular 
two-lipped corollas. 

Conopholis alpina Liebmann, Forh. Skand. Na- 
turf. Mode 4: 184. 1847. C. sylvatica Lieb- 
mann, Forh. Skand. Naturf. Mode 4: 185. C. 
mexicana Gray ex Watson, Proc. Am. Acad. 



Arts Sci. 18: 131. 1883. C. panamensis Wood- 
son, Ann. Missouri Bot. Gard. 25: 835. 1935. 
C. alpina var. mexicana (Gray ex Watson) 
Haynes, Sida 4: 255. 1971. Figure 28. 

Parasitic herbs, stems 6-24(-30) cm tall, 4-12 
mm diam., glabrous, proximal scales imbricate, 
thickened at the base, glabrous. Leaves repre- 
sented by sessile scales 6-18(-21) mm long, 4- 
7(-l 1) mm wide, ovate-lanceolate to narrowly ob- 
long-lanceolate, widest just above the base, apex 
acute to acuminate, margins entire, drying sub- 
coriaceous, surfaces glabrous, often dark brown 
distally (in life), drying uniformly yellowish 
brown. Inflorescences 2-18 cm long, 2-4 cm 
wide, bracts 8-20 mm long, separate or overlap- 
ping, usually concealing the calyx, pedicels 0-3 
mm long, bracteoles 2-4 mm long, ca. 1.5 mm 
wide, ovate-elliptic, on the lateral sides of the ca- 
lyx or absent. Flowers with calyx 8-12 mm long, 
3-4 mm diam., ovoid-tubular, usually split down 
the anterior side, lobes 2-5, acute to rounded; co- 
rolla 7-15 mm long, curved downward, white to 
greenish white (yellowish white when dried), gla- 
brous externally; filaments 9-15 mm long, anthers 
1.5-2.5 mm long, with basal appendages ca. 0.4 
mm long, usually glabrous. Fruits 8-18 mm long, 
6-1 1 mm diam., ovoid-rounded, drying dark, 
style often persisting; seeds 0.6-1 mm long, with 
rounded edges, surface yellowish to dark brown 
and lustrous, minutely reticulate (20X). 

Plants of the forest floor in montane oak forests, 
2000-2800 m elevation. Flowering in December- 
April; fruiting in February-July. Costa Rican col- 
lections come from the Cordillera de Talamanca 
and Volcan Irazu. This species, as circumscribed 
by Haynes, ranges from the southwestern United 
States to northern Panama. 

Conopholis alpina is recognized by its lack of 
green pigment, restriction to high-elevation oak 
forests, short thick unbranched stems, curved tu- 
bular yellowish corollas, exserted stamens, and 
many-seeded capsules. The seeds germinate only 
near young oak roots, and the flowers are polli- 
nated by bumblebees (Gomez, 1980). Haynes 
placed our material in variety alpina, which rang- 
es from southern Mexico to Panama. However, 
the senior author has seen these plants in a num- 
ber of areas ranging from Chiapas to Panama and 
believes that there is a discontinuity of morphol- 
ogy in the Guatemala-Honduras region. Thus, it 
is possible that the plants of southern Central 
America are deserving of taxonomic recognition 
as C. panamensis. 



166 



FIELDIANA: BOTANY 




FIG. 28. Orobanchaccac: Conopholis alpina. A, Calyx in lateral view. B, Scpaloid hractcolc. C, A different calyx 
in side view. D, Ventral view of stamen. E, Lateral view of stamen. F, Placcntation. G, Inflorescence and base. 



BURGER: FLORA COSTARICENSIS 



167 



Orobanche Linnaeus 

Herbs parasitic on roots, lacking chlorophyll, 
stems simple or branched, usually succulent, usu- 
ally covered with minute gland-tipped hairs. 
Leaves reduced to alternate sessile scales, sepa- 
rate along the stem, yellowish to brown or purple, 
margins entire. Inflorescences terminal spikes or 
racemes, bracts conspicuous and similar to the 
scales, bracteoles present or absent. Flowers with 
calyx cupulate to campanulate, 2-5-lobed or di- 
vided to the base adaxially and abaxially, lobes 
equal or unequal; corolla tubular, often with mi- 
nute gland-tipped hairs, 2-lipped, 4- or 5-lobed, 
upper lip entire or 2-lobed; stamens 4, included, 
filaments borne near the base of the tube, anthers 
2-thecous, thecae parallel; disc not apparent; ova- 
ry 1-locular, ovules many on 4 parietal placentas, 
stigma 2-lobed. Fruits capsules, oblong, enclosed 
within the persisting corolla; seeds ellipsoid, 
coarsely reticulate. 

Orobanche is a genus of about 150 species, 
mostly north temperate, and is especially speciose 
in Europe. The following species has become a 
serious agricultural problem in some parts of the 
world. 



Orobanche minor J. E. Smith, English Bot. 6: 
422, tab. 422. 1797. 

Herbs with single or several clustered stems 8- 
80 cm tall, the stems rarely with distal branches, 
4-10 mm diam., pale rose-yellow to brownish, 
surface with glandular hairs 0.2-1 mm long. 
Leaves represented by alternate simple scales 1- 
3 cm long, sessile, broadly ovate, acute, venation 
parallel. Inflorescences spicate, bracts 9-12 mm 
long, ca. 4 mm wide, sessile, difficult to distin- 
guish from the sepals in dried material, pedicels 
0-2(-5) mm long, glandular puberulent, bracte- 
oles absent. Flowers with calyx usually split to 
the base abaxially and adaxially, each portion 
with 2 long narrow acuminate lobes; corolla 12- 
18 mm long, slightly purplish with purple veins 
but usually yellowish at the base and lower lip, 
sparsely puberulent externally, tube ca. 4 mm 
diam.; filaments ca. 6 and 7 mm long. 

Orobanche minor is not native to the Americas 
and is rarely collected in the American tropics. 
The senior author observed this species near Pru- 
sia, at 1500 m elevation on the slopes of Vocan 
Irazu, Cartago province, in November 1979 (G6- 
mez 7140 CR, usj), where it was parasitizing a 



species of clover (Trifolium) and two species of 
grasses. This adventive population probably came 
from introduced grass or forage (G6mez, 1980). 
These plants are distinctive because of their lack 
of green pigmentation, erect unbranched stems 
with gland-tipped hairs, scale-like leaves similar 
to the bracts, split calyx, and tubular purple and 
yellowish corollas. Although associated with a va- 
riety of herbs and small shrubs, this species is 
often found parasitizing plants of the family Fa- 
baceae. In tropical areas they are unlikely to be 
found below 1000 m elevation. 



Literature Cited 

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CANNE, J. 1980. Seed morphology in Aureolaria and 70- 
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CRONQUIST, A. 1981. An Integrated System of Classifi- 
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D'ARCY, W. G. 1979. Scrophulariaceae, in Flora of Pan- 
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173-272. 

GENTRY, A. H. 1973a. Generic delimitations of Central 
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MORALES, J. F, AND Q. JIMENEZ. 1997. Bignoniaceae. 
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168 



FffiLDIANA: BOTANY 



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REVEAL, J. L. 1995. Newly required supragcneric names 
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BURGER: FLORA COSTARICENSIS 



169 



Index 

The index includes all accepted names (in roman type), synonyms (italics), common English names 
(roman) and vernacular Spanish names (italics). The text includes only one new species (boldface), 
Amphitecna gentryi. 



Adenocalymma 100 
Adenocalymma apurcnsc 101 
Adenocalymma calderonii 100 
Adenocalymma ci liola turn \ 33 
Adenocalymma cocleense 160 
Adenocalymma fissum 135, 136 
Adenocalymma flos-ardeae 1 50 
Adenocalymma heterophyllum \ \ 1 
Adenocalymma hintonii 100 
Adenocalymma hosmeca 133 
Adenocalymma inundatum 100 
Adenocalymma macrocarpum 133 
Adenocalymma ocositensis 1 1 8 
Adenocalymma perezii 135 
Adenocalymma punctifolium 1 50 
Adenocalymma seleri 135 
Adenocalymma standleyanum 1 1 7 
Adenocalymma veruciferum 135 
Adenocalymma verrucosa 1 1 5 
African tulip tree 149 
Agalinis 5 
Agalinis albida 5 
Agalinis hispidula 17 
Agalinis peduncularis 5 
ajillo 133 
ajonjoli 164 
Alectra 14 
Alectra aspera 14 
Alectra brasiliensis 14 
Alectra fluminensis 14 
Alectra melatnpyroides 14 
Alonsoa 15 

Alonsoa meridionalis 15 
Alonsoa warscewiczii 19 
Amphilophium 100 
Amphilophium molle 101 
Amphilophium oxylophium 102 
Amphilophium paniculatum 101 

var. molle 101 

Amphilophium pannosum 102 
Amphilophium pilosum 102 
Amphitecna 102 
Amphitecna donnell-smithii 104 
Amphitecna gentryi 103 
Amphitecna haberi 106 
Amphitecna isthmica 104 
Amphitecna kennedyi 104 
Amphitecna latifolia 105 
Amphitecna molinae 104 
Amphitecna obovata 105 
Amphitecna sessilifolia 105 
Amphitecna silvicola 104 
Anemopaegma 106 
Anemopaegma chrysanthum 107, 

109 

Anemopaegma chrysoleucum 107 
Anemopaegma macrocarpa 107 
Anemopaegma orbiculatum 107 



Anemopaegma pubcrulum 108 
Anemopaegma puncticulatum 1 07 
Anemopaegma santaritense 109 
Anemopaegma tobagense 117 
Anemopaegma tonduzianum 121 
angelon 17 
Angelonia 16 
Angelonia angustifolia 16 
Angelonia ciliaris 16 
Anisantherina 5, 17 
Anisantherina hispidula 17 
Anonymous umbrosa 53 
Antirrhinum 18 
Antirrhinum majus 18 
Antirrhinum canadensis 45 
Antirrhinum cymbalaria 36 
Antirrhinum linaria 46 
Antirrhinum vulgaris 117 
Antirrhinum muralis 87 
Arrabidaea 109 
Arrabidaea candicans 1 10 
Arrabidaea chica 1 1 1 
Arrabidaea conjugata 1 1 1 
Arrabidaea corallina 1 12 
Arrabidaea costaricensis 1 1 2 
Arrabidaea dichasia 140 
Arrabidaea e recta 1 12 
Arrabidaea florida 1 1 3 
Arrabidaea guatemalensis 121 
Arrabidaea inaequalis 109 
Arrabidaea isthmica 113, 121 
Arrabidaea lundellii 1 14 
Arrabidaea mollicoma 113 
Arrabidaea mollissima 113 
Arrabidaea pacy calyx 1 10 
Arrabidaea panamensis 113 
Arrabidaea pattelifera 1 14 
Arrabidaea potosina 123 
Arrabidaea pseudochica 121 
Arrabidaea pubescens 1 1 5 
Arrabidaea rhodothyrsus \ 10 
Arrabidaea verrucosa 1 1 5 
Asarina erubescens 49 
Asarina scandens 50 



Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 



18 

auriculata 21 
axillaris 20 
bacopoides 20 
bracteolata 20 
curtipes 23 
decumbens 21 
cgcnsis 21 
humilis 23 
lacertosa 21 
laxiflora 21 
limosa 22 
monnieri 22 



Bacopa monnierioides 22 
Bacopa myriophylloides 25 
Bacopa naias 25 
Bacopa pan'iflora 22 
Bacopa procumbens 52 
Bacopa ranaria 22 
Bacopa reflexa 25 
Bacopa rcpens 23 
Bacopa salzmannii 23 
Bacopa sessiliflora 24 
Bacopa valerii 23 
Bacopa violacea 23 
bateita 146 
bees, Centris 17 
bejuco de ajo 135 
Benjaminia 24 
Benjaminia reflexa 25 
Benjaminia utriculariaeformis 24 
Bignonia aeqinoctialis 121 
Bignonia aesculifolia 125 
Bignonia binata 118 
Bignonia candicans 110 
Bignonia capensis 157 
Bignonia chica \ 1 1 
Bignonia chrysantha 152 
Bignonia chrysoleucum 107 
Bignonia conjugata 111 
Bignonia copaia 127 
Bignonia corallina 1 1 2 
Bignonia corymbifera 129 
Bignonia crucigera 145 
Bignonia dasyonyx 132 
Bignonia diversifolia 122 
Bignonia echinata 145 
Bignonia glabrata 112 
Bignonia hymenaea 133 
Bignonia ignea 147 
Bignonia inaequilatera 150 
Bignonia kerere 133 
Bignonia latifolia \ 16 
Bignonia lepidota 122 
Bignonia litoralis 113 
Bignonia magnoliaefolia 124 
Bignonia mollicoma 113 
Bignonia mollissima 113 
Bignonia obliqua 112 
Bignonia obovata 136 
Bignonia orbiculata 107 
Bignonia paniculata 101 
Bignonia pannosa 102 
Bignonia patellifera \ 1 4 
Bignonia pubescens \ 14 
Bignonia pyramidata 140 
Bignonia quadrivalvis 137 
Bignonia riparia 150 
Bignonia sarmentosa 121 
Bignonia sinclairii 140 
Bignonia stans 157 



170 



FIELDIANA: BOTANY 



Bignonia tetragonoloba \ \ 1 
Bignonia tiliaefolia 145 
Bignonia umbrosa 129 
Bignonia uncata 131 
Bignonia uncinata 131 
Bignonia ungiiis-cati 1 3 1 
Bignonia variabilis 146 
Bignonia venusta 147 
Bignonia verrucifera 135 
Bignoniaccac 77 

list of species 161 
boca de dragon 18 
boca de la vieja 17 
boca de ledn 1 8 
bombilla 147 
baton de oro 28 
Bourreria costaricensis 75 
Buchnera 25 
Buchnera grandiflora 97 
Buchnera leiantha 26 
Buchnera longifolia 26 
Buchnera major 26 
Buchnera mexicana 26 
Buchnera palustris 27 
Buchncra pusilla 26 
Buchnera tinctoria 26 
Buchnera weberbaueri 26 

Cabomba 25 
Caconapea auriculata 21 
Caconapea axillaris 34 
Caconapea conferta 24 
Caconapea paniflora 22 
Caconapea sessiliflora 24 
calabacero 106, 120 
calabash 106, 120 
calabash, swamp 105 
calabasillo de playa 1 05 
Calceolaria 27 
Calceolaria irazucnsis 28 
Calceolaria costaricensis 28 
Calceolaria heterophylla 30 
Calceolaria mexicana 28 
Calceolaria microbefaria 29 
Calceolaria perfoliata 30 
Calceolaria sciadephora 30 
Calceolaria storkii 29 
Calceolaria trachelifolia 28 
Calceolaria trilobata 30 
Calceolaria tripartita 30 
Calceolaria urticina 28 
Callichlamys 116 
Callichlamys garneri 1 16 
Callichlamys latifolia 1 16 
candelillo 158 
carhoncillo 158 
Capraria 31 
Capraria bi flora 3 1 

var. pilosa 3 1 
Capraria Crustacea 47 
Capraria durantifolia 61 
Capraria gratioloides 48 
Carpoceras angulata 164 
Castilleja 32 
Castilleja agrestis 33 
Castilleja arvensis 33 



Castilleja aurantiaca 34 
Castilleja hicolor 34 
Castilleja chiriquiensis 34 
Castilleja communis 33 
Castilleja irasuensis 33 
Castilleja lentil 34 
Castilleja quirosii 34 
Caxtilleja seibertii 34 
Castilleja talamancensis 35 
Castilleja tayloriorum 35 
Ccratophy turn 1 1 7 
Ceratophytum tciragonolobum 117 
Ceratophytunt tobagense I 1 7 
Chelone gentianoides 54 
Chodanthus puberulum 108 
chorro 148 
chorro de oro 148 
Clerodendrwn epiphyticum 72 
Clytostoma 1 1 7 
Clytostoma binatum 1 1 8 
Clytostoma callistegioidcs 1 1 8 
C 'lyiostoma islhmicum 1 1 8 
Clytostoma mayanum 123 
Coliseum Ivy 88 
coloring agents, dyes 39 
Conobea alata 43 
Conobea pusilla 57 
Conopholis 165 
Conopholis alpina 166 

var. mexicana 166 
Conopholis americana 166 
Conopholis mexicana 166 
Conopholis panamensis 1 66 
Conopholis sylvatica 166 
coral 56 
coralillo 56 
corrimiento 62 
cones 152 

carles amarilla 153, 154 
coneza 153, 154 
cortez negro 154 
Couralia rosea 155 
cow okra 141 
Crcscentia 1 19 
Crescentia aculeata 141 
Crescentia alata 1 19 
Crcscentia cujcte 120 
Crescentia edulis 141 
Crescentia latifolia 105 
Crescentia ohovata 105 
Crescentia pinnata 1 28 
Crcscentia ternata 119 
cucharilla 146 
culantrillo 58 
(\bistax macrocarpa 125 
Cydista 120 
Cydista aequinoctalis 1 2 1 

var. hirtella 121 
Cydista divcrsifolia 122 
Cydista heterophylla 122 
Cydista lilacina 123 
Cydista potosina 123 
Cvdista puhescens 1 2 1 
Cvdista sarmentosa 121 
Cymbalaria 36 



Cymbularia muralis 36 

Daixsa 36 

Darcya costaricensis 37 
Darcya reliquiarum 37 
Dtndrosicos i\ihiuicn\ l()4 
Dfiulrosicos kennedyi 104 
Dendrosicos latijoliu\ 105 
Dermatocalyx par\-ijioms 76 
i/i.U/M/ 39 
Digitalis 38 
Digitalis purpurea 38 
Distictella 124 
Distictclla inagnoliilblia 124 
Distictis laxiflora 101 
Distictis rovirasana 160 
Disteira angulo\a 164 
Dodartieae 100 
domatia 196 
Dombeya lappacea 158 
Doxantha dasyonyx 132 
Doxantha unguis-cati 132 

Enallagma latifolia 105 
Enallagma sexsil (folia 1 05 
Erinus procumbens 52 
Erinus verticillatus 62 
escoba amarga 58 
Escobedia 39 
Escobedia curialis 39 
Escobedia grandiflora 39 
Escobedia lacvis 39 
Escobedia long/flora 39 
Escobedia reticulata 39 
Escobedia scabrifolia 39 
escoheta 58 
escobilla amarga 58 

Fagelia perfoliata 30 
figure scales 5 
y/or </<' //u/fV> 35 
flowering phenology 120 
foxglove 39 

gallinazo 128 
gallito 34 
gallitos 28 
Gentry, A. H. v. 78 
Gerardia 5. 
Gerardia hispidula 17 
Gibsoniothamnus 70 
Gibsoniolhamnus alatus 74 
Gibsoniothamnus cornutus 72 
Gibsoniothamnus cpiphyticus 72 
Gibsoniothamnus grandiflorus 73 
Gibsoniothamnus mirilicus 73 
Gibsoniothamnus parvifolius 73 
Gibsoniothamnus pithccobius 72 
Gibsoniothamnus ptcrocalyx 73 
Gibsoniothamnus stcllatus 74 
Globifcra umbrostim 53 
Glossotylis aspcra 14 
GcxJmania 125 
Gcxlmannia acsculifolia 125 
Godmannia macrocarpa 1 25 
Gratiola anagallidea 48 



INDEX 



171 



Gratiola dubia 48 
Gratiola inaequalis 48 
Gratiola repens 23 
guacales 120 

Hebe 66 
Hcmichacna 40 
Hemichaena fruticosa 40 
Hemimcridae 18 
Herpestis auriculata 2 1 
Herpestis axillaris 34 
Herpestis bacopoides 20 
Herpestis caparioides 52 
Herpestis ciliata 23 
Herpestis laxiflora 21 
Herpestis peduncularis 52 
Herpestis ranaria 22 
Herpestis reflexa 24 
Herpestis repens 23 
Herpestis salzmannii 23 
Herpestis sessiliftora 24 
hierba santa 62 
Hydranthelium egense 21 

Ilysanthes dubia 48 
Ilysanthes gratioloides 48 

jacaranda, jacaranda \ 28 
Jacaranda 126 
Jacaranda caucana 127 

ssp. sandwithiana 127 
Jacaranda copaia 127 

ssp. spectabilis 127 
Jacaranda ficifolia 127 
Jacaranda mimosifolia 128 
Jacaranda ovalifolia 128 
Jacaranda spectabilis 127 
Jacaranda superba 127 
Jacaranda trianae 127 
jfcaras 120 
jicarita 105 
jf'/caro 106, 119, 120 
jicaro danto 143 
jicaro de playa 105 
Jovellana 60 
>//a 157 

Kenilworth Ivy 88 
Kigelia 128 
Kigelia africana 128 
Kigelia pinnata 128 

Lamourouxia 41 
Lamourouxia gutierrezii 42 
Lamourouxia lanceolata 42 
Lamourouxia longiflora 42 
Lamourouxia longifolia 

var. lanceolata 42 
Lamourouxia rhinanthifolia 43 
Lamourouxia smithii 43 
Lamourouxia veijensis 42 
Lamourouxia viscosa 42 
Lamourouxia scabra 42 
Leucocarpus 43 
Leucocarpus alatus 43 
Leucocarpus fruiticosus 40 



Leucocarpus pcrfoliatus 43 

Levya nicaraguensis 121 

Limnophila costaricensis 25 
forma aquatica 25 
forma semiterrestris 25 

Limosclla 44 

Limosclla acaulis 44 

Limosella americana 44 

Limosella australis 45 

Linaria 45 

Linaria canadensis 45 

Linaria texana 45 

Linaria vulgaris 46 

Linaria cymbalaria 36 

///M/fl 393 

Lindernia 46 

Lindernia anagallidea 48 

Lindernia Crustacea 47 

Lindernia dianthera 52 

Lindernia diffusa 47 

Lindernia dubia 48 

Lindernia gratioloides 48 

Lindernia inaequalis 48 

Lindernia japonica 5 1 

Lindernia microcalyx 48 

Lindernia thouarsii 65 

Lindernia verticillata 62 

llama del bosque 149 

lluvia de coral 56 

Lobelia pumila 5 1 

Lophospermum 48 

Lophospermum erubescens 49 

Lundia 129 

Lundia colombiana 130 

Lundia corymbifera 129 

Lundia dicheilocalyx 130 

Lundia puberula 130 

Lundia schumanniana 130 

Lundia valenzuelae 129 

Lysimachia monnieri 22 

Macfadyena 130 
Macfadyena guatemalensis 1 3 1 
Macfadyena phellosperma 144 
Macfadyena uncata 131 
Macfadyena uncinata 131 
Macfadyena unguis-cati 131 
Macuillamia limosa 23 
Macuillamia repens 23 
Manga de la Senora 39 
Mansoa 132 
Mansoa hymenaea 133 
Mansoa kerere 133 

var. erythraea 134 

var. incarnata 134 
Mansoa parvifolia 134 
Mansoa standleyi 135 
Mansoa verrucifera 135 
mariposas 146 
Marline! la 136 
Martinella obovata 136 
Martinella verrucosa 1 1 5 
Martyniaceae 164 
Martynia 164 
Martynia angulosa 164 
Martynia annua 164 



Martynia diandra 164 
Maurandya 49 
Maurandya barclaiana 50 
Maurandya erubescens 49 
Maurandya scandens 50 
Maurandya semperflorens 50 
Mazus 51 

Mazus japonicus 51 
Mazus pumilus 51 
Mazus rugosus 51 
Mecardonia 51 

Mecardonia montevidensis 52 
Mecardonia procumbens 52 
medicinal properties 32, 39, 151 
Melasma melampyroides 14 
Mella laxiflora 21 
Melloa 137 

Melloa quadrivalvis 137 
Micalia grandiflora 97 
Micranthemum 52 
Micranthemum pilosum 53 
Micranthemum standleyi 53 
Micranthemum umbrosum 53 
Mimulus 53 
Mimulus glabratus 53 
Mimulus perfoliatus 43 
mi'rame Undo 147 
monkey flowers 53 
Monniera axillaris 34 
Monniera reflexa 50 
Monocardia humilis 23 
Monocardia violacea 23 
morro 1 1 9 
Mussatia 138 
Mussatia hyacinthina 138 
Myoporaceae 31 

Naiadothrix longipes 25 
Neotuerckheimia gonoclada 105 
Nortenia thouarsii 65 

Onohualcoa 138 
Onohualcoa fissa 135 
Onohualcoa seleri 135 
Onohualcoa verrucifera 135, 136 
Orobanchaceae 5, 165 
Orobanche 168 
Orobanche minor 168 

Pachyptera 132, 138 
Pachyptera kerere 134 
Pachytera parvifolia \ 34 
Pachyptera standleyi 135 
palomitas 146 
Pandorea 146 
Pandorea ricasoliana 147 
Parabignonia 138 
Parabignonia steyermarkii 139 
Paradolichandra 139 
Paragonia 139 
Paragonia pyramidata 140 
Parmentiera 140 
Parmentiera aculeata 141 
Parmentiera alata 1 19 
Parmentiera cerifera 141 
Parmentiera dressleri 142 



172 



FIELDIANA: BOTANY 



Parmentiera edulis 141 
Parmentiera macrophylla 142 
Parmentiera trunciflora 143 
Parmentiera valerii 143 
Pedaliaceae 162 
Pedicularis meUonpyroides 14 
pedo de padre \ 33 
peina de mico 146 
peina de mono 146 
Peloria 118 
Pennywort 88 
Penstemon 54 
Penstcmon gentianoidcs 54 
Penstemon skutchii 54 
pepino de danto 143 
Petastoma breviflorum 1 14 
Petastoma ocositense 1 1 8 
Petastoma patellifera 1 14 
Petastoma pubescens 1 14 
Petastoma tonduzianum 133 
Phryganocydia 143 
Phryganocydia corymbosa 144 
Phryganocydia phellospcrma 144 
Pithecoctenium 145 
Pithecoctenium crucigerum 145 
Pithecoctenium echinatum 145 
Pithecoctenium panamensis 107 
Pleonotoma 146 
Pleonotoma diversifolia 122 
Pleonotoma variabilis 146 
Podranea 146 
Podranea ricolasiana 147 
pollinators 23 
Porodittia 60 
porto hello 17 
Proboscidia triloba 164 
Pseudocalymma alliaceum 

var. macrocalyx 135 
Pseudocalymma sagotii 133 

var. macrocalyx 135 
Pseudocalymma standleyi 135 
Pyrostegia 147 
Pyrostegia ignea 147 
Pyrostegia venusta 147 
Pyxidaria Crustacea 47 
Pyxidaria diffusa 47 

quacalillo 106 

Quinquelobulus utriculariaeoides 
25 

Ranaria monnierioides 22 
Reichardia scandens 50 
Rhodochiton 49 
roble 155 

roble de bianco 1 55 
roble de sabana 155 
roble macho 154 
Russelia 55 

Russelia colombiana 56 
Russelia equisetiformis 55 
Russelia flavoviridis 56 
Russelia juncea 55 
Russelia oxyphylla 56 
Russelia sarmentosa 56 
Russelia tabacensis 56 



Saldanhaea costaricensis 1 1 2 
Saldanhaea seemanniana 160 
San Carlos 148 
Saritea 148 
Saritca magnifica 148 
Schistophragma 56 
Schistophragma mcxicana 57 
Schistophragma pusilla 57 
Schlcgcliaceae 69 
Schlegclia 74 
Schlegclia brachyantha 75 
Schlegelia costaricensis 75 
Schlegclia fastigiata 75 
Schlegelia fuscata 76 
Schlegclia nicaraguacnsis 76 
Schlegelia parasitica 76 
Schlegclia parviflora 76 
Schlegelia silvicola 76 
Schlegclia sulfurea 76 
Scobinaria 148 
Scobinaria japurensis 115 
Scobinaria verrucosa \ \ 5 
Scoparia 57 
Scoparia annua 58 
Scoparia dulcis 58 
Scrophularia fluminensis 14 
Scrophularia meridionalis 15 
Scrophularia procumbens 23 
Scrophulariaceae 1, 2 

list of species 69 
sesame 162 
Sesamum 162 
Sesamum indicum 162 
Sesamum orientalc 162 
Sibthorpia 58 
Sibthorpia pichinensis 59 
Sibthorpia rcpens 59 
Sibthorpia triandra 59 
slipper flowers 27 
snapdragon 18 
Sparattosperma rosea 155 
Spathodea 148 
Spathodea campanulata 149 
Spathodea corymbosa 144 
Spathodea laurifolia 144 
Spathodea nilotica 148 
Spathodea obovata 136 
Spathodea orinocensis 144 
Stemodia 59 
Stemodia angulata 60 
Stemodia arenaria 62 
Stemodia costaricensis 37 
Stemodia durantifolia 61 
Stemodia jorullcnsis 61 
Stemodia jorullensis 

ssp. re plans 60 
Stemodia pan-ijiora 62 
Stemodia pcduncularis 61 
Stemodia reliquiarum 37 
Stemodia siliquosa 57 
Stemodia verticillata 62 
Stemodiacra angulata 60 
Stemodiacra peduncularis 6 1 
Stizophyllum 149 
Stizophyllum flos-ardeae 150 
Stizophyllum inacquilatcrum 150 



Stizophyllum punctifolium 150 
Stizophyllum riparium 150 
swamp calabash 105 
sweet broom 58 

Tabebuia 151 
Tabehuia calderonii 100 
Tabebuia chrysantha 152. 154 
ssp. chrysantha 152 
ssp. pluvicola 152. 153 
Tabebuia dugandii 153 
Tabebuia guayacan 153 
Tabebuia impctiginosa 153 
Tabebuia latifolia 1 1 6 
Tahebuia me.\icana 155 
Tabebuia neochrysantha 154 
Tabebuia nicaraguensis 153 
Tabebuia ochracca 154 

ssp. neochrysantha 154 
Tabebuia palmeri 153 
Tabebuia palustris 154 
Tabebuia pentaphylla 1 55 
Tahebuia pyramidata 140 
Tabebuia rosea 155 
Tabebuia scrratifolia 153 
Tabebuia sessilifolia 105 
Tabebuia sinclairii 139 
Tabebuia speciosa 1 16 
Tanaecium 156 
Tanaecium albiflora 156 
Tanaecium jaroba 1 56 
Tanaecium zetekii 133 
Tccoma 156 
Tccoma capensis 157 
Tecoma chrysantha 152 
Tecoma evenia 152, 155 
Tecoma fuscata 1 25 
Tecoma guayacan 153 
Tecoma impetiginosa 153 
Tecoma mexicana 155 
Tecoma ochracea 154 
Tecoma palmeri 1 52, 4 1 2 
Tecoma ricasoliana 147 
Tecoma rosea 155 
Tecoma stans 157 
Tecomaria 156 
Tecomaria capensis 157 
Tclrancma 62 
Tctranema floribundum 63 
Tctrancma gamboanum 63 
Tctranema mcgaphyllum 63 
Tctrancma roscum 172 
Torcnia 64 
Torcnia asiatica 65 
Torenia Crustacea 47 
Torcnia fournicri 64 
Torcnia thouarsii 65 
Tourrcttia 158 
Tourrcttia lappacca 158 
Tourrettia volubilis 158 
triquitraque 148 
tulipdn 149 
Tynanthus 159 
Tynanthus guatcmalcnsis 159 
Tynanthus hyacinthinus I3H 
Tynanlhus macranthus 159 



INDEX 



173 



una de gato \ 32 Veronica 65 Veronica serpyllifolia 68 

unas de diablo 165 Veronica arvensis 66 van humifusa 68 

Uroskinnera 63 Veronica crenulata 68 Veronica tenella 68 

Usteria scandens 50 Veronica didyma 68 Veronica xalapensis 67 

Utricularia 25 Veronica humifusa 68 

Veronica pcregrina 67 

. . . co t en Wil ichia repens 59 

vainillo 1 58 van xalapensis 67 

Vandelia Crustacea 47 Veronica persica 67 

Vandelia diffusa 47 Veronica polita 68 Xylophragma 160 

Vatkea diandra 164 Veronica rotundifolia 67 Xylophragma seemannianum 160 



174 FIELDIANA: BOTANY 



UNIVERSITY OF ILLINOIS-URBANA