Trees, shrubs, vines, or herbs, terrestrial or hemiepiphytic (Markea and Lycianthes); stems often prickly; stellate pubescence often present. Leaves alternate or suboppo- site, petiolate; blades simple, entire or lobed; venation pinnate; stipules lacking. Flowers bisexual, actinomor- phic (zygomorphic in Browallia), generally solitary or in cymose axillary clusters; calyx 5-lobed; corolla (4)5-lobed; stamens (4)5 or 4 and didynamous (Browal- lia), epipetalous; anthers 2-celled, dehiscing longitudi- nally or poricidally (Solatium); ovary superior, 1- or 2-locular, 2-carpellate; placentation axile; ovules many (few in Cestrum); style 1; stigma minutely bilobed. Fruits generally berries with many seeds, rarely septicidally dehiscent capsules (Browallia); seeds with fleshy endo- sperm.
Members of the family are most easily distinguished by their actinomorphic, gamopetalous, usually plicate corollas and berries of many seeds. They are closely related to the Scrophulariaceae (125), which generally have zygomorphic flowers.
All species are probably insect pollinated. Solarium and Cyphomandra (G. Frankie, pers. comm.), with api- cal pores, are mostly pollinated by drumming, pollen- collecting bees. I have seen Solatium subinerme visited in this way by Melipona, and G. Frankie (pers. comm.) has observed xylocopids drumming flowers of 5. tridy- namum Dun. (S. amazonicum Ker.) in Mexico. Cestrum species have flowers that open late in the afternoon and are mostly moth pollinated. The flowers fall before mid- morning of the following day.
The fruits are endozoochorous, except perhaps in the case of Browallia americana. Capsicum has been widely dispersed by man. Most species are dispersed at least in part by birds (Ridley, 1930), but probably especially those of Cestrum, Lycianthes, Physalis, and Witheringia solanacea, which have fruits so small as to be unattractive to larger animals. Smaller fruits may be dispersed in their entirety by birds. Because species of Solanaceae have fairly thin-walled fruits, birds probably pick open the larger fruits and take the seeds. On the other hand, spiny rats eat fruits of Physalis (Hladik & Hladik, 1969), and white-faced monkeys are reported to eat fruits of Cestrum (Oppenheimer, 1968). The fleshy part of the Cestrum fruit is sweet, but the seed is bitter and is swallowed whole. Bats may be important in the distribution of fruits
124.
SOLANACEAE/BROWALLIA739
KEY TO THE TAXA OF SOLANACEAE Plants herbs to ca 1 m tall:
Flowers blue, zygomorphic; fruits capsular Browallia americana L.
Flowers not blue, actinomorphic; fruits berries:
Flowers pale yellow or white (often with a dark "eye" at base), solitary in leaf axils; fruits globose berries surrounded by the inflated calyx Physalis Flowers greenish or yellow-green, lacking a dark throat, not solitary in leaf axils; fruits not
as above:
Leaf blades ± alike, not deeply lobed; flowers in dense axillary clusters; fruits ca 8 mm diam, orange-red; plants occasional in clearings Witheringia solanacea L'Her.
Leaf blades dimorphic, some entire, others deeply lobed; flowers few in pendent axillary racemes; fruits ca 2 cm diam with green stripes when young; plants probably not on island Cyphomandra allophylla (Miers) Hemsl.
Plants trees or shrubs more than 1 m tall:
Corolla narrowly tubular, the tube much longer than lobes; fruits obovoid or ellipsoid, less than 9 mm diam, purple or black at maturity; calyx lobes acute Cestrum Corolla not narrowly tubular, the tube usually as short as or much shorter than lobes (tube
longer than lobes in Markea but corolla campanulate):
Corolla lobes divided nearly to base:
Flowers in very long, pendent cymes in age; petals greenish; anthers to 10 mm long, the connective thickened dorsally; fruits 3-5 cm diam Cyphomandra hartwegii (Miers) Dun.
Flowers not in long, pendent cymes; petals white or lavender; anthers with the connective not thickened:
Calyx truncate, sometimes with submarginal teeth reduced to bumps; plants unarmed shrubs Lycianthes Calyx 5-lobed (with 5 minute, marginal teeth and appearing truncate in S. subinerme);
plants shrubs, vines, or herbs, often prickly Solarium Corolla lobes not divided to near base:
Calyx distinctly lobed; plants hemiepiphytic; leaves ± coriaceous, glabrous beneath; fruits ovoid, yellowish, ca 1.5 cm long; plants in forest Markea ulei (Damm.) Cuatr.
Calyx truncate; plants not hemiepiphytic; leaves thin, pubescent beneath; fruits globose or elongate, red at maturity; plants common in clearings:
Flowers solitary in axils; anthers bluish, not apiculate; fruits narrowly ovoid, more than 3 cm long; pubescence of leaves ± restricted to vein axils below Capsicum annuum L. var. annuum Flowers clustered in axils; anthers not bluish, apiculate; fruits globular, ca 8 mm diam;
pubescence of leaves not restricted to vein axils below .... Witheringia solanacea L'Her.
as well, especially the long-pendent fruits of Cypho- mandra and the larger arborescent or lianous Solanum species such as 5. hayesii and S. lanciifolium. Markea ulei is taken by bats (Bonaccorso, 1975). Heithaus, Fleming and Opler (1975) reported species of Solanaceae to be taken by several species of bat. A species of Solanum is taken by the bat Artibeus jamaicensis Leach in Mexico (Yazquez-Yanes et al., 1975).
About 80 genera and over 3,000 species; concentrated in the New World.
BROWALLIA L.
Browallia americana L., Sp. PL 631. 1753 Chavelita de monte
Herb, usually less than 1 m tall, puberulent to sparsely villous. Leaves alternate, simple; petioles to 2 cm long;
blades ovate, acute to acuminate, acute at base, 1.5-7 cm long, 1-4 cm wide, entire. Flowers 5-parted, short- pedicellate, solitary in axils, 1.5-2 cm long; calyx 6-8(10) mm long, narrowly campanulate, striate, pubescent;
corolla tube greenish, slender, pubescent, the limb laven- der to blue, the throat 1-1.5 cm wide, minutely puberu- lent apically, with a prominent white or green spot near throat (nectar guide); stamens 4, the upper pair situated in throat near apex opposite nectar guide, the connective broadened, violet-purple, pubescent, their anthers dehisc- ing into cup-shaped cavities on one side of the much- thickened, elaborate style apex, the lower pair of stamens held slightly below the upper pair, their filaments hooked at apex; ovary 1-1.5 mm long, appressed-pubescent at apex; stigma bilobed. Fruits 4-valved, ellipsoid capsules contained within the accrescent calyx; seeds irregularly round, to 1.5 mm long, muricate. Croat 10258.
Uncommon; found in shady places at the Laboratory Clearing. Probably flowering and fruiting all year, espe- cially in the dry season.
American tropics. In Panama, ecologically variable;
most common at middle and upper elevations, occurring in most life zones in highland Chiriqui and mountainous regions of central Panama; less frequently from tropical dry forest in Panama (Taboga Island) and from tropical moist forest in the Canal Zone, Panama, and Darien.
740 DICOTYLEDONEAE CAPSICUM L.
Capsicum annuum L. var. annuum, Sp. PL 188-89. 1753
C. frutescens sensu Standl.
Red pepper, Chile, Aji, Aji picante
Shrub, less than 2 m tall. Leaves alternate or subopposite with a large leaf opposed by a smaller one; petioles to 3 cm long, densely pubescent; blades ovate to elliptic, falcate-acuminate, attenuate at base, 6-11(15) cm long, 2.5-5 cm wide, puberulent above and on veins below, tufted in veins axils below, ciliate. Flowers 5-parted, solitary (rarely paired) in axils; pedicels to more than 2 cm long; calyx truncate, ca 3 mm long, nearly glabrous, the teeth 5, blunt, submarginal, accrescent and persisting in fruit; corolla greenish-white, divided one-half to one- third of the way to base, to ca 17 mm diam, reflexed at anthesis, glabrous on outside, papillate-puberulent inside and on margins, the lobes acute; anthers bluish; filaments violet-purple near apex, fused to tube basally; style slightly exceeding anthers. Berries variable, often ovoid, ca 4.5 cm long, bright red at maturity. Croat 9007.
Cultivated at the laboratory. Flowering and fruiting all year, apparently with most fruits maturing during the rainy season.
Probably native to tropical America; cultivated throughout the world and throughout Panama.
CESTRUM L.
Cestrum latifolium Lam., Illustr. 2:5, no. 2275. 1793 Shrub or small tree, to 3.5(12) m tall; stems, especially on younger parts, petioles, and leaves, especially on veins below, sparsely to densely pubescent with weak multi- cellular trichomes. Leaves alternate; petioles 1-2 (4) cm long; blades ovate to elliptic, acuminate, acute to obtuse at base, 7-16(25) cm long, 3.5-8.5 cm wide. Racemes or panicles short, congested, axillary, the branches and calyces pubescent; flowers 5-parted, to 3.5 cm long; calyx 1-3 mm long, the lobes acute; corolla narrowly tubular, 13-18(20) mm long, whitish-yellow or greenish, the lobes 5 (6), narrowly acute, 2-2.5 mm long, pubescent outside along margins, glabrous inside; stamens 5, in- cluded; filaments fused to tube most of their length;
anthers about as broad as long, held just below rim;
stigma truncate; style held just above anthers. Berries
obovoid, pink to black at maturity, 5-9 mm long; seeds several. Croat 11996.
Uncommon, known from Rear # 8 Lighthouse Clear- ing and from the forest near the Laboratory Clearing.
Flowers from April to September, but mostly in the early rainy season, from May to July. The fruits mature from July to November, mostly from August to October.
Nicaragua to the Guianas and Brazil; the Antilles. In Panama, known from tropical moist forest in the Canal Zone, San Bias, Veraguas, Panama, and Darien, from tropical dry forest in Los Santos, from premontane moist forest in the Canal Zone, from premontane wet forest in Panama, and from tropical wet forest in Chiriqui.
Cestrum megalophyllum Dun. in DC, Prodr.
13(1):638. 1852 C. baenitzii Ling.
Shrub or small tree, to 8(10) m tall; outer bark thin;
young stems green, sparsely crisp-pubescent, glabrous in age. Leaves alternate; petioles to 1.3(2.5) cm long, gla- brous; blades narrowly elliptic to obovate-elliptic, acumi- nate, gradually tapered to an acute or rounded base, 12-25(35) cm long, 3.5-8(12) cm wide, ± glabrous.
Racemes short, axillary, fasciculate; peduncles crisp- pubescent, bracteate; flowers 5-parted, usually many, often on leafless stems; calyx cupular, 2-3.5(4) mm long;
corolla salverform, ca 1.5 cm long, the tube greenish, constricted just below lobes, the lobes white, spreading at anthesis, 4-5 mm long; stamens included; filaments fused to tube in basal two-thirds, pubescent near their point of fusion, turned inward below anthers; style slightly longer than stamens; stigma capitate, held just above anthers; nectary inconspicuous at base of ovary.
Berries ellipsoid, 6-10 mm long, green becoming whitish then light violet and finally dark violet when mature;
mesocarp thin, fleshy, white; seeds 1 or 2, minutely papil- late. Croat 14568.
Occasional, in the forest. Flowering from November to June, mostly in the dry season, from February to April.
The fruits mature in the early rainy season, from May to August.
Guatemala to Venezuela. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, Los Santos, and Darien, from premontane wet forest in Colon and Chiriqui, from tropical wet forest in Colon, and from lower montane wet forest in Chiriqui.
See Fig. 480.
KEY TO THE SPECIES OF CESTRUM
Mature leaves pubescent beneath C. latifolium Lam.
Mature leaves glabrous beneath:
Largest leaves more than 15 cm long, gradually tapered to a usually acute base, often broadest above middle C. megalophyllum Dun.
Largest leaves less than 15 cm long, not gradually tapered to base, broadest at or below middle:
Filaments conspicuously dentate near base; leaf blades usually acute at base .... C. nocturnum L.
Filaments not dentate at base; leaf blades usually obtuse to rounded at base C. racemosum R. & P.
Cestrum nocturnum L., Sp. PI. 191. 1753 Dama de noche
Shrub or small tree, to 5 m tall. Leaves alternate; petioles ca 1 cm long; blades ovate, acute to acuminate, obtuse to rounded at base, to 11 cm long, to 5.5 cm wide, minutely puberulent when young, glabrous in age. Panicles axil- lary or terminal, congested; peduncles with scattered leaf- like bracts; pedicels to 4 mm long, with minute bracteoles;
flowers 5-parted; calyx campanulate, puberulent, 2 mm long; corolla greenish to yellowish-white, 14-17 mm long, glabrous outside; stamens equal; filaments ca 3 mm long, dentate near point of insertion; stigma slightly exserted. Berries ellipsoid, to 10 mm long, black at ma- turity; seeds not studied. Bangham 429.
Collected once on the shore. Seasonal behavior uncer- tain. According to our records, flowers mostly in the dry season, but also in August.
Native to the Antilles, but cultivated and escaped in many other places in tropical America. In Panama, known from tropical moist forest in the Canal Zone and Bocas del Toro and from tropical wet forest in Colon and Chiriqui.
Cestrum racemosum R. & P., Fl. Peruv. 2:29, pi.
154. 1799
C. panamense Standl.; C. racemosum R. & P. var. panamense (Standl.) Franc.
Yedi
Tree, to 12 m tall. Leaves alternate; petioles ca 1-2 cm long, glabrate; blades lanceolate-elliptic, long-acuminate, mostly obtuse to rounded at base, 7-15 cm long, 2.5-5 cm wide, ± glabrous. Cymes branched, axillary or ter- minal, 3-9 cm long, the branches crisp-villous; flowers 5-parted; calyx 2.3-3.3 mm long, the lobes acute, ciliate;
corolla to 15 mm long, slender, cream or greenish-white, the lobes villous inside; stamens included; filaments fused to tube most of their length; style flat on end, held slightly above anthers; nectaries golden-yellow. Berries obovoid or ellipsoid, ca 6 mm long, black, with thickened pedicels ca 2 mm long; seeds usually 3, wedge-shaped (like orange segments), densely papillate, ca 4 mm long, embedded in a juicy white matrix, very bitter before maturity, sweet at maturity. Croat 8899.
Uncommon, known from the edge of the Laboratory Clearing. Flowering from December to May, mostly in the late dry season and the early rainy season (April to May), rarely as late as September. Frequently flowering more than once per season. The fruits mature mostly from February to December.
Belize to Brazil. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, Panama, and Darien, from premontane wet forest in Chiriqui and
124. SOLANACEAE/CYPHOMANDRA 741 Code, and from tropical wet forest in Colon and Code.
Reported from premontane rain forest in Costa Rica (Holdridgeetal., 1971).
CYPHOMANDRA Mart, ex Sendt.
Cyphomandra allophylla (Miers) Hemsl., Biol. Centr.- Amer. Bot. 2:417. 1882
Solarium allophyttum (Miers) Standl.
Hierba de gallinazo, Hierba gallota
Erect herb, to 1 m tall, glabrate. Leaves 2 or 3 suboppo- site at a node (often with 1 leaf lobed and the other 2 entire), dimorphic; petioles 1.5-5 cm long, narrowly winged; blades broadly ovate, acuminate, rounded at base, entire or 3-5-lobed, the entire blades 5-6 cm long and 3-4 cm wide, the lobed blades 9-11 cm long and 6-10 cm wide. Racemes short, at the dichotomies of stems; peduncles 1-2 cm long; pedicels ca 1 cm long, broadening at apex; flowers 1-6; calyx ca 2 mm long;
corolla white or yellowish, 10-12 mm long, lobed about halfway; filaments ca 1 mm long, attached to dorsal sur- face of a membranous ring inserted near base of corolla tube; anthers lanceolate, ca 6 mm long, with 2 terminal pores; ovary elongate, conical; stigma punctiform. Berries ellipsoid, ca 2 cm long, white at maturity, sometimes striped with green when young; seeds yellow, flattened, ca 1.5 mm long. Aviles 105, Zetek 5040.
Collected twice; not seen in recent years, but it could reoccur in clearings. Flowering and fruiting mostly in the rainy season, but probably to some extent through- out the year.
Known only from the Pacific slope of Panama, from tropical moist forest in the Canal Zone, Panama, and Darien and from tropical dry forest in the Canal Zone.
Cyphomandra hartwegii (Miers) Dun. in DC, Prodr.
13(1):401. 1852
C. heterophylla Donn. Sm., nomen nudum; C. costaricensis Donn. Sm.; C. mollicella Standl.
Monca prieto
Soft-wooded shrub or tree, to 5 m tall; most parts sparsely to densely puberulent. Leaves alternate or subtended by a smaller leaf; petioles mostly 3-6(10) cm long; blades ovate, acuminate and often downturned at apex, un- equally obtuse to cordate at base, 6-25 (30) cm long, 5-13 cm wide, the upper surface sparsely puberulent to glabrate, some of the trichomes usually glandular- tipped; juvenile leaves often deeply lobed, to 45 cm long and 25 cm wide. Cymes upper-axillary, indeterminate, to 50 cm or more long before flowers cease being pro- duced, pendent; flowers 5-parted, closely spaced on pe- duncle; pedicels to 3 cm long in flower, elongating and
KEY TO THE SPECIES OF CYPHOMANDRA
Plants herbs to 1 m tall; leaves 2 or 3 per node, subopposite C. allophylla (Miers) Hemsl.
Plants shrubs or trees to 5 m tall; leaves alternate or opposite with 1 leaf of each pair much smaller C. hartwegii (Miers) Dun.
'* '\J'
%^^,^^1::U/,
'*/"<•'
mJ f 'JJKA
•^^
,_ ••••• ... y -•••:• - . • .••;.;.•••: :f,p~';^ "- •• • ;
rSrKiSKK
.
' J.
Fig. 482. Markea ulei
Fig. 483. Marfea wfe;
thickening at apex in fruit; calyx ca 5 mm long, the lobes short, acute to obtuse; corolla green, 1.5-3.5 cm long, the lobes long-tapered, free to calyx; filaments thick, short, fused to corolla tube; anthers to 10 mm long, papil- late, the connective swollen, purple, the pores terminal;
stigma held just above anthers in bud, elongating after anthesis. Berries globose to ellipsoid, 3-5 cm long, green with lighter stripes, yellow at maturity; seeds numerous, disk-shaped, reniform to round, to 6 mm long. Croat 9016.
Uncommon, appearing in tree-fall areas of the forest and at the edges of clearings. The plant flowers unceas- ingly for long periods, with individual plants flowering for as much as a year or more.
Visited by male Eulaema bombiformis bees {fide label on Dressier 3070).
Honduras to Brazil and Bolivia. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, and Panama, from premontane wet forest in Chiri- qui and Panama, from tropical wet forest in Colon, and from premontane rain forest in Colon, Code, and Panama.
See Fig. 481.
LYCIANTHES (Dun.) Hassl.
Lycianthes maxonii Standl., J. Wash. Acad. Sci.
17:14. 1927
L. maxonii Standl. var. appendiculata Standl.
Slender shrub, to 1.5 m tall; younger stems and petioles puberulent; nodes somewhat swollen. Leaves alternate;
petioles to 5 mm long; blades mostly oblong-obovate, acuminate, cuneate to obtuse at base, 7-15 cm long, 2-5 cm wide, glabrous, entire, minutely undulate; major veins few, arcuate-ascending. Fascicles very short, axillary, usually with only 1-3 (6) flowers or fruits at any time;
pedicels 1-2.5 cm long in flower, to ca 3.5 cm long in fruit; flowers 5-parted; calyx truncate, with 5 short, wart- like protuberances below rim; corolla violet, 5-10 mm long, divided to near base, the lobes spreading at anthesis;
stamens yellow, shorter than corolla, held together at apex in a ring; filaments united to tube in basal half, very short; anthers 4 mm long, with 2 terminal pores; ovary ovoid, glabrous; style straight, held well above anthers, nearly equaling length of corolla. Berries globose to obovoid, to 1.2 cm long, orange becoming red at matur- ity; seeds ca 4 mm long, longer than broad. Croat 6307.
Frequent, especially in the old forest. Flowers from March to December, mostly from April to September.
The fruits develop within about a month, maturing from April to January, mostly from July to October. Plants usually have flowers and fruits simultaneously.
124.
SOLANACEAE/MARKEA743
Usually growing about a meter tall, but becoming nearly twice as tall in areas of the older forest.
Nicaragua to Panama. In Panama, known from tropical moist forest in the Canal Zone, Panama, and Darien, from premontane wet forest in Panama, from tropical wet forest in Colon, from lower montane wet forest in Chiriqui, and from premontane rain forest in Panama (summit of Cerro Jefe).
Lycianthes synanthera (Sendt.) Bitter, Abh. Naturwiss.
Vereine Bremen 23 (2):499-500. 1919
Shrub or tree, usually hemiepiphytic, ca 2 (10) m tall,
± glabrous except for tufted axils on lower leaf surfaces.
Leaves alternate or subopposite with the pairs unequal;
petioles 5-30 mm long; blades ± elliptic, acuminate, acute to obtuse at base, 5-20 cm long, 3-8.5 cm wide.
Flowers solitary or in fascicles of few flowers; pedicels ca 1.5 mm long; calyx cyathiform, nearly truncate, 2-3 mm long, becoming woody and 5-6 mm long in fruit;
corolla 5-lobed to near base, ca 1 cm long, purplish;
stamens 5, ± equal; anthers 6-7 m long, yellow, united into an ellipsoid column ca 3 mm diam; style exserted.
Berries globose, ca 7 mm diam, fleshy; seeds discoid and margined, ca 1.5 mm long, foveate, yellow. Croat 11899.
Rare, known only from the area east of Wheeler Trail 1600. Flowers from May to September. The fruits mature from June to October.
Mexico to Panama. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, Colon, Veraguas, and Panama, from premontane wet forest in Chiriqui, from tropical wet forest in Code and Panama, and from premontane rain forest in Code.
MARKEA L. C. Rich.
Markea ulei (Damm.) Cuatr., Feddes Repert.
61:78. 1958 M. panamensis Standl.
Hemiepiphytic shrub, often associated with ant nests;
smaller stems fleshy, sometimes arising from a large, swollen, tuberous stem perched in the crotch of a tree, with only a slender root trailing to the ground; stems and leaves glabrous. Leaves alternate or subopposite with the pairs equal or unequal; petioles less than 1 cm long; blades obovate to broadly elliptic, acuminate, acute to rounded at base, 6-20 cm long, 3-8.5 cm wide, coria- ceous. Panicles short, axillary, cymose; peduncles stout, to ca 7 cm long; pedicels 10-14 mm long; flowers 5-parted, ca 1.3 cm long; calyx deeply divided, the lobes oblong- ovate, curved inward, to ca 7 mm long, persisting in fruit;
corolla campanulate, to 14 mm long, greenish-yellow, densely pubescent outside with very short glandular
KEY TO THE SPECIES OF LYCIANTHES Plants hemiepiphytic; corolla more than 1 cm long Plants terrestrial; corolla less than 1 cm long
L. synanthera (Sendt.) Bitter L. maxonii Standl.