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FLORA OF GUATEMALA 



JOHNNIE L. GENTRY, JR. 



AND 



PAUL C. STANDLEY 



FIELDIANA: BOTANY 

 VOLUME 24, PART X, NUMBERS 1 and 2 



Published by 



FIELD MUSEUM OF NATURAL HISTORY 

 APRIL 11, 1974 





FLORA OF GUATEMALA 



PART X 





FLORA OF GUATEMALA 



JOHNNIE L. GENTRY, JR. 



Assistant Curator, Vascular Plants 

 Field Museum of Natural History 



AND 



PAUL C. STANDLEY 



Late Curator of the Herbarium 

 Field Museum of Natural History 



FIELDIANA: BOTANY 

 VOLUME 24, PART X, NUMBERS 1 and 2 



Published by 



FIELD MUSEUM OF NATURAL HISTORY 

 APRIL 11, 1974 




Library of Congress Catalog Card Number: 4.8-3076 



PRINTED IN THE UNITED STATES OF AMERICA 

 BY FIELD MUSEUM PRESS 




520.5 

 F6 



CONTENTS 

 Families Included in Part X, Numbers 1 and 2 



SOLANACEAE . 



PAGE 



1 





LIST OF ILLUSTRATIONS 



TEXT FIGURES 



PAGE 



1. Acnistus arborescens 5 



2. Bouchetia erecta and Browallia americana 8 



3. Brunfelsia nyctaginoides 11 



4. Capsicum frutescens 16 



5. Cestrum formosum 26 



6. Cestrum mortonianum 30 



7. Cyphomandra rojasiana 36 



8. Datura Candida 39 



9. Jaltomata procumbens 44 



10. Juanulloa mexicana 46 



11. Lycianthes barbatula 51 



12. Lycianthes connata 55 



13. Lycopersicon esculentum var. cerasiforme 67 



14. Markea neurantha 71 



15. Melanthus guatemalensis and Nicandra physaloides 74 



16. Nicotiana glauca 76 



17. Physalis lagascae var. glabrescens and Schwenckia americana 89 



18. Solandra grandifiora 96 



19. Solanum torvum 141 



20. Witheringia stramonifolia 150 





Flora of Guatemala - Part X, Numbers 1 and 2 



SOLANACEAE. Potato or Nightshade Family 

 JOHNNIE L. GENTRY, JR. AND PAUL C. STANDLEY 



Herbs, shrubs, or small trees, sometimes woody vines, occasionally epiphytic, 

 often armed with prickles, the pubescence of simple or often stellate or variously 

 branched hairs; leaves alternate, sometimes in pairs (subopposite), simple, entire 

 to deeply parted, or odd-pinnate, stipules absent; inflorescences cymose, often 

 racemose, subumbellately clustered or paniculately disposed or reduced to a single 

 flower, pseudoterminal, lateral and opposite the leaves or internodal, or axillary; 

 flowers perfect, actinomorphic or sometimes zygomorphic, typically 5-merous; 

 calyx gamosepalous, campanulate to tubular, 5-lobed or -parted, sometimes 4-6- 

 lobed, usually persistent and in some accrescent or much inflated in fruit; corolla 

 gamopetalous, subrotate, campanulate, funnelform, salverform, tubular, or urceo- 

 late, the limb 5-lobed, mostly plicate, valvate, or imbricate in bud; stamens 5, 

 sometimes 4, or only 2, epipetalous, alternate with the corolla lobes; anthers with 

 2 thecae, 1 theca sometimes reduced, dehiscent by longitudinal slits or by terminal 

 pores, sometimes connivent, the connective sometimes enlarged; pistil 1; ovary 

 superior, bilocular, sometimes tetralocular by false septae, or 3-5 locular, or rarely 

 unilocular, with axile placentation, the ovules several to many in each cell, rarely 

 few or one, the hypogynous disk usually present and apparent; style solitary and 

 terminal; stigma capitate and entire or shallowly bilobed, sometimes dilated or 

 somewhat expanded and flattened; fruit a berry or capsule; seeds several to many, 

 sometimes few, or rarely one, often strongly compressed, sometimes angular and 

 often with a pitted testa; embryo subperipheral, strongly curved, spiral, or straight; 

 endosperm carnose. 



A family of about 85 genera with some 2,300 species, cosmopoli- 

 tan, best developed in tropical America. 



All of the solanaceous genera in Central America are represented 

 in Guatemala except Margaranthus (included in this treatment). 

 Two other genera, Fabiana imbricata R. & P., a native of South 

 America, and Lycium halimifolium Miller, a native of Europe, are oc- 

 casionally grown in Guatemalan gardens. Valerioa originally placed 

 in the Solanaceae by Standley and Steyermark is synonymous with 

 Peltanthera in the Loganiaceae and Eutheta placed here by Standley 

 is a synonym of Melasma in the Scrophulariaceae. 



Solanaceae are of great economic importance, yielding foods, such 

 as Solanum tuberosum (potato), S. melongena (eggplant), Ly coper si- 




2 FIELDIANA: BOTANY, VOLUME 24 



con esculentum (tomato), Capsicum spp. (peppers), drugs, such as 

 Atropa belladonna (atropine or belladonna), Datura stramonium 

 (stramonium), Hyoscyamus niger (hyoscyamine and scopolamine) , 

 Nicotiana tabacum (tobacco), and ornamentals, such as Browallia, 

 Brunfelsia, Oestrum, Datura, Lycium, Nicotiana, Nierembergia, Pe- 

 tunia, Physalis, Salpiglossis, Schizanthus, Solandra, Solanum, and 

 Streptosolen. 



Corolla subrotate to campanulate or urceolate, rarely tubular-campanulate. 

 Anthers dehiscing by terminal pores or slits. 



Calyx truncate and often bearing 5 or 10 appendages just below the margin 



of the calyx Lycianthes. 



Calyx 5-lobed or 5-parted; calyx appendages absent. 



Anther connective broad and thickened adaxially; plants unarmed. 



Cyphomandra. 

 Anther connective slender, not prominently thickened adaxially; plants 



often armed with prickles Solanum. 



Anthers dehiscing by longitudinal slits. 



Leaves pinnate; anthers coherent and terminated by sterile tips. 



Ly coper sicon. 



Leaves simple; anthers free or sometimes coherent but not terminated by 

 sterile tips. 



Corolla urceolate Margaranthus. 



Corolla subrotate to campanulate, rarely tubular-campanulate. 



Calyx lobes sagittate-cordate at their bases; corolla blue Nicandra. 



Calyx lobes not sagittate-cordate at their bases; corolla white, greenish- 

 white, yellow or yellowish-brown. 



Fruiting calyx accrescent and spreading beneath the berry or bladdery- 

 inflated and enclosing the berry, the calyx contracted at the apex 

 and often invaginated basally. 



Inflorescences umbellate and long-pedunculate; fruiting calyx spread- 

 ing beneath the berry Jaltomata. 



Inflorescences consisting of a single pedicellate flower or sometimes 

 in axillary fascicles; fruiting calyx bladdery-inflated and en- 

 closing the berry, the calyx contracted at the apex . . . Physalis. 

 Fruiting calyx not at all to strongly accrescent, sometimes surpassing 

 the berry but not bladdery-inflated and enclosing the berry, the 

 calyx not contracted at the apex nor spreading beneath the berry. 

 Inflorescences consisting of axillary fascicles, with the calyx 5 mm. 

 long or less, truncate, 5-lobed or -dentate not more than one-half 

 way to the base or rarely with the inflorescences long-peduncu- 

 late, and the leaves in pairs, dimorphic, very unequal in size and 



different in shape Witherlngia. 



Inflorescences axillary, consisting of 1-2 (-3) pedicellate flowers or if 

 the flowers in axillary fascicles the calyx 12 mm. long or more 

 and 5-parted nearly to the base. 



Corolla throat pubescent; corolla limb parted one-half way or 

 nearly to the base, calyx appendages absent and the anthers 



yellowish; fruit not pungent to the taste Athenaea. 



Corolla throat glabrous; corolla limb scarcely lobed, with the 

 calyx truncate and bearing 5 linear or subulate appendages 

 just below the margin of the calyx or the anthers bluish to 

 bluish tinged; fruit often very pungent to the taste. 



Capsicum. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 3 



Corolla tubular, funnelform or salverform. 

 Fertile stamens 4 and didynamous, or the stamens 2, sometimes with 1-3 



staminodes; corolla lobes sometimes linear or clavate. 

 Corolla tubular, the flowers small and inconspicuous. 



Corolla very small, 3 mm. long or less; fruit tuberculate, tardily dehiscent, 



acute at the apex, one-seeded Melananthus. 



Corolla 10-13 mm. long; fruit smooth, readily dehiscent, obtuse at the 



apex, several-seeded Schwenckia. 



Corolla salverform, the flowers large and showy. 



Corolla tube less than 2 cm. long, the throat closed by filaments; thecae of 



the upper anthers dimorphic; seeds angulate; herbs Browallia. 



Corolla tube 3 cm. long or more, the throat not closed by filaments; thecae 

 of the upper anthers equal in size; seeds rounded; shrubs or sometimes 



suffrutescent herbs Brunfelsia. 



Fertile stamens 5, sometimes one reduced; staminodes absent. 

 Leaves in pairs, dimorphic, very unequal in size and different in shape. 



Witheringia. 

 Leaves solitary, rarely in pairs, but then not dimorphic. 



Inflorescences consisting of axillary fascicles Acnistus. 



Inflorescences racemose, cymose or paniculate, or consisting of a single 



pedicellate flower. 

 Corolla 16 cm. long or more. 



Corolla yellowish or yellowish-brown, the corolla lobes obtuse, imbri- 

 cate in bud; scandent shrubs Solandra. 



Corolla white, the corolla lobes acuminate to long-caudate, plicate in 



bud; erect shrubs or small trees Datura. 



Corolla 12 cm. long or less. 



Fruit a berry; plants shrubs or small trees. 



Corolla salverform or narrowly funnelform, 3 cm. long or less; fruit 

 much exceeding the calyx, and short-stipitate; seeds few; 



plants terrestrial Cestrum. 



Corolla broadly funnelform or tubular, 3.5 cm. long or more; fruit 

 surpassed by the calyx, if not then the leaves glandular- 

 punctate, the fruit not stipitate; seeds numerous; plants 

 nearly always epiphytic. 

 Pubescence of stellate and branched hairs; corolla tubular. 



Juanulloa. 

 Pubescence of simple hairs or hairs absent; corolla funnelform. 



Markea. 

 Fruit a capsule, septicidally dehiscent; plants nearly always herbaceous. 



Inflorescences racemose or paniculate Nicotiana. 



Inflorescences consisting of a single pedicellate flower. 



Plants viscid-pubescent; calyx parted nearly to the base. Petunia. 

 Plants pubescent to nearly glabrous, but not viscid-pubescent; 



calyx lobed not more than one-half way to the base. 

 Corolla tube 4.5 cm. long or more; calyx deciduous, circumscissle 

 near the base and leaving a persistent flaring collar under 

 the fruit; fruit armed with spines; leaves subentire to shal- 



lowly lobed, 3 cm. wide or more Datura. 



Corolla tube 1.5 cm. long or less; fruit unarmed, included in the 

 persistent calyx; leaves entire, 1 cm. wide or less. 



Bouchetia. 



ACNISTUS Schott 



Shrub or small trees; leaves alternate, entire, petiolate; inflorescences consist- 

 ing of axillary fascicles, the flowers long pedicellate; calyx campanulate, 4-5-lobed 




4 FIELDIANA: BOTANY, VOLUME 24 



or sometimes truncate, slightly or not at all accrescent in fruit; corolla funnelform, 

 white, the limb 5-lobed, the lobes valvate or induplicate-valvate in bud; stamens 5; 

 filaments slender, inserted just below the middle of the tube; anthers oblong or 

 ovate, longitudinally dehiscent, apiculate; ovary bilocular, the ovules numerous, 

 the disk prominent, carnose; style slender and exserted; stigma dilated and shortly 

 bilobed; fruit a berry; seeds numerous, laterally compressed; embryo strongly 

 curved. 



About 20 species, with only one native in Central America, rang- 

 ing from southern Mexico to Costa Rica, West Indies, and South 

 America. A few other species have been described from Mexico. 

 Some authors treat Acnistus Schott as a synonym of Dunalia H.B.K., 

 see Sleumer, Lilloa 23: 117-142. 1950, but also see A. T. Hunziker, 

 Bol. Acad. Nac. Ci. 41 : 211-244. 1960. The generic limits of Acnistus 

 and Witheringia need to be critically reviewed. 



Acnistus arborescens (L.) Schlecht. Linnaea 7: 67. 1832. 

 Atropa arborescens L. Cent. PL 2: 10. 1756. Palo de gallina. 



Damp or wet thickets, 1,360 m.; Santa Rosa; Jalapa. Southern 

 Mexico. Nicaragua; Costa Rica. West Indies. South America. 



A shrub or a small tree with corky-like bark, commonly 3-6 m. tall; leaves 

 elliptic to oblong, mostly 5-15 cm. long, 3-5.5 cm. wide, acute or obtuse, attenuate 

 at the base, glabrous or nearly so above, densely tomentose-floccose with branched 

 or simple multicellular hairs below, sometimes glabrate; petioles 1-1.5 cm. long; 

 flowers numerous, in axillary fascicles; pedicels 1.5-2.5 cm. long; calyx chartaceous, 

 light brown, 2.5-4 mm. long, glabrous, the lobes short and rounded, about 1 mm. 

 long, sometimes truncate; corolla white, 8-11 mm. long, glabrous within, glabrous 

 or puberulent externally, the tube 4.5-6.5 mm. long, the lobes 3-5 mm. long, 

 spreading and acute; stamens exserted; filaments 4-5.5 mm. long; anthers 2-2.5 

 mm. long, apiculate; style 9-11 mm. long; berry orange or yellow, 5-6 mm. in 

 diameter, globose, pulpy; seeds reticulate-foveate. 



Acnistus arborescens is not very common in Guatemala, at least 

 not often collected. It is more abundant in some parts of Costa Rica. 



ATHENAEA Sendtner, nom. cons. 



Herbs or shrubs; leaves alternate, sometimes in pairs, entire or sinuate- 

 angulate, petiolate; inflorescences consisting of axillary fascicles or of 1 or some- 

 times 2 pedicellate flowers; calyx campanulate, 5-lobed, slightly to much accrescent 

 in fruit; corolla subrotate, white or yellow, often with 5 darker contrasting spots 

 near the mouth of the corolla tube, the mouth villosulous, the limb 5-lobed, the 

 lobes valvate in bud; stamens 5; filaments filiform or dilated at the base, exserted; 

 anthers oblong, longitudinally dehiscent; ovary bilocular, the ovules numerous, the 

 disk inconspicuous; style slender, exserted; stigma entire or emarginate, slightly 

 dilated; fruit a berry, surpassed by or included in the accrescent calyx, the fruiting 

 calyx sometimes appressed to the fruit, but not bladdery-inflated and enclosing the 




FIG. 1. Acnistus arborescens. A, habit, X }/%', B, corolla dissected, X 4; C, 

 calyx and style, X 5. 




6 FIELDIANA: BOTANY, VOLUME 24 



fruit, the base not invaginate; seeds numerous, laterally compressed; embryo 

 strongly curved. 



About 15 species in tropical America. One other Central Amer- 

 ican species occurs in Costa Rica. The generic limits of Athenaea 

 are not well defined, at least in Central America. It is treated as a 

 synonym of Withania by some workers, but that genus is apparently 

 found only in the Old World. A few species previously placed here 

 are currently treated in Witkeringia. 



A more thorough study of the generic limits of Athenaea and re- 

 lated genera will probably result in some modification from the pres- 

 ent treatment (see comments under the species) . 



Inflorescences consisting of 1 or sometimes 2 pedicellate flowers; leaves narrow, 2-4 

 cm. wide, the base obtuse; corolla limb 1 cm. wide; calyx 3.5-4.5 mm. long. 



A. cernua. 



Inflorescences consisting of axillary fascicles, the flowers 5-9; leaves broad, 10-18 

 cm. wide, the base cordate; corolla limb 2.5 cm. wide; calyx 14 mm. long. 



A. viscosa. 



Athenaea cernua Donn.-Sm. Bot. Gaz. 48: 297. 1909. Physalis 

 melanocystis (Robinson) Bitter var. cernua Waterfall, Rhodora 69: 

 99. 1967, syn. nov. 



Moist forests, 900 m.; Pete"n; Alta Verapaz (type from near 

 Sasis, Tuerckheim II 2245) . Mexico. 



A much branched herb, the stems with short glandular hairs, often with long 

 multicellular hairs in addition to the shorter ones; leaves solitary or sometimes in 

 pairs, then similar in size and shape, the leaves ciliate, pubescent with short multi- 

 cellular hairs above and below, the hairs often glandular, mostly confined to the 

 veins below, apex acuminate or acute, base obtuse, the larger leaves mostly 4-13.5 

 cm. long, 2-4 cm. wide, the petioles 1-3.5 cm. long; flowers solitary or sometimes 2, 

 the pedicels slender, 1.5-1.8 cm. long, cernuous in fruit, with short glandular hairs; 

 calyx 3.5-4.5 mm. long, glandular pubescent with multicellular hairs, the lobes 

 lanceolate to narrowly triangular, accrescent in fruit; corolla subrotate, yellow or 

 white with 5 darker contrasting (at least when dried) villosulous spots near the 

 mouth of the tube, the mouth of the corolla tube villosulous, the limb 1 cm. wide, 

 parted to the middle or just below, the lobes about 2.5-3 mm. long, ciliate; fila- 

 ments 1.5 mm. long; anthers 1.2 mm. long; style 3-3.5 mm. long; berry globose, 

 red, about 8 mm. in diameter, at maturity the accrescent calyx splits and becomes 

 reflexed, exposing the mature fruit; seeds 1-1.2 mm. long. 



Athenaea cernua has been treated as a variety of Physalis melan- 

 ocystis by Waterfall (1967). The fruiting calyx indicates stronger 

 affinities with Athenaea than with Physalis. 



Athenaea viscosa (Schrader) Fern. Proc. Am. Acad. Sci. 35: 

 567. 1900. Saracha viscosa Schrader, Index Sem. Hort. Acad. Goett. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 7 



5. 1832. A. macrocardia Standl. & Steyerm. Field Mus. Bot. 22: 375. 

 1940 (type from below Finca Alejandria, Sierra de las Minas, Za- 

 capa, Steyermark 30004). 



In pine forested canyons, 2,000 m.; Zacapa. Mexico. 



An herb 3-3.5 m. tall, the stems densely villosulous, the hairs glandular and 

 multicellular; leaves solitary, entire to sinuate-angulate, broadly ovate, 15-30 cm. 

 long, 10-18 cm. wide, pubescent above and below with short and long multicellular, 

 glandular hairs, the longer hairs more dense on the veins, apex acuminate or long- 

 acuminate, base oblique, deeply cordate; petioles long, 4-12 cm. long, densely 

 glandular- villosulous; flowers few to numerous, the slender pedicels, 2.5-5 cm. 

 long, densely glandular- villosulous; calyx 1.4 cm. long, densely glandular-villo- 

 sulous, deeply parted, the lobes ovate, 1.2 cm. long, apex narrowly long-acute, 

 slightly accrescent in fruit; corolla subrotate, densely glandular-villosulous 

 externally, pale yellow-cream, with 5 olive-green spots near the mouth of the tube, 

 the mouth of the corolla tube villosulous, the limb about 2.5 cm. wide, parted to 

 below the middle, the lobes 1 cm. long, ciliate; filaments 4.5 mm. long; anthers 4 

 mm. long; style 10 mm. long; berry globose, orange, 1.2-1.5 cm. in diameter; seeds 

 1.5 mm. long. 



Athenaea viscosa is morphologically similar to a group of species 

 in Physalis that have the corolla 5-lobed and the flowers mostly sev- 

 eral in the leaf axils, but it differs in characters of the fruiting calyx. 

 The species is retained in Athenaea until further study is conducted. 

 The correct name is Physalis schraderiana Bernh., when referred to 

 that genus. 



BOUCHETIA A. De Candolle 



Herbs, erect or decumbent, pubescent; leaves alternate, entire, short petiolate 

 or sessile; inflorescences reduced to a single flower, pseudoterminal and subopposite 

 the leaves, the flowers pedicellate; calyx tubular-campanulate, 5-lobed, slightly 

 accrescent in fruit; corolla funnelform, white or blue, the limb 5-lobed, the lobes 

 short, imbricate-plicate in bud, the posterior lobe outermost in bud, the tube 

 slender below, ampliate above; stamens 5, 4 didynamous, the fifth one shorter than 

 either pair or with 2 long and 3 shorter stamens; filaments filiform, subequally 

 inserted below the middle of the corolla tube; anthers connivent, the thecae 

 confluent at the apex, longitudinally dehiscent; ovary bilocular, the ovules 

 numerous, the disk cupular; style filiform, smooth and included; stigma trans- 

 versely dilated, oblong, reflexed and somewhat acute on both ends; fruit a capsule, 

 septicidal, the valves bifid; seeds numerous and angulate; embryo almost straight. 



A genus of one or possibly two species ranging from the south- 

 western United States to Guatemala and South America. 



Bouchetia erecta DC. ex Dunal in DC. Prodr. 13: 589. 1852. 



Open grassy places, 1,400-1,600 m.; only one collection is known 



from Guatemala (Huehuetenango) between Las Palmas and Chacula, 




FIG. 2. Bouchetia erecta. A, habit, X 1; B, corolla dissected, X 1 1 A, calyx 

 dissected, X 1 1 A- Browallia americana. C, habit, X 1 A; D, flower, X 2; E, style 

 and stigma, X 10; F, upper stamen, X 10; G, lower stamen, X 10. 



8 




GENTRY & STANDLEY: FLORA OF GUATEMALA 9 



Sierra de los Cuchumatanes, Steyermark 51761. Southwestern 

 United States. Mexico. South America. 



Low perennial herbs, the stems slender, few to several, erect or decumbent, 2 

 dm. long or less, sparsely pubescent with inwardly curved hairs; leaves linear to 

 lance-ovate, 1.5-3.5 cm. long, 0.3-1 cm. wide, acute, the base cuneate, nearly 

 glabrous or puberulent, the lower leaves short petiolate, the upper ones sessile; 

 flowers solitary, the pedicels 1.5-3 cm. long; calyx pilose, 6-10 mm. long, the lobes 

 2.5-4 mm. long, linear, unequal or equal, acute or obtuse; corolla white or blue, 

 pilose externally, the limb about 1 cm. wide, the tube 1-1.5 cm. long, the lobes 

 short, obtuse; stamens with two long pairs, the fifth one shorter than either pair or 

 with two long and three shorter stamens; filaments 4.5-7.5 mm. long; anthers 

 about 1 mm. long, the thecae confluent at the apex; style 6-9.5 mm. long, included; 

 capsule 5.5-7 mm. long, included in the accrescent calyx; seeds angulate, about 1 

 mm. long, reticulate-foveate. 



BROWALLIA L. 



Herbs; leaves alternate and entire, petiolate; inflorescences racemose or the 

 flowers solitary; calyx tubular, mostly 5-lobed, slightly accrescent in fruit; corolla 

 salverform, slightly zygomorphic, blue or violet, occasionally white, the limb 

 5-lobed, the lobes imbricate-plicate in bud, the posterior lobes outermost in bud, 

 the tube ventricose just below the mouth of the tube; perfect stamens 4, didyna- 

 mous, included, the fifth stamen rudimentary or absent; filaments subequally 

 inserted in pairs above the middle of the corolla tube; anthers longitudinally 

 dehiscent, the thecae of the upper anthers dimorphic, the larger thecae pollen- 

 bearing, the smaller theca sterile; ovary bilocular, the ovules numerous, the disk 

 inconspicuous; style filiform, corrugated above, included; stigma expanded and 

 somewhat flattened, slightly bilobed; fruit a capsule, membranaceous or sub- 

 coriaceous, septicidal, the valves bifid; seeds numerous and angulate; embryo 

 straight or slightly curved. 



Probably only two species in the genus, although several others 

 have been described, ranging from southern Mexico to Costa Rica, 

 Panama, West Indies, and South America. The other species (Bro- 

 wallia speciosa Hook.) occurs in Central America in the mountains 

 of Costa Rica and Panama. 



Browallia americana L. Sp. PI. 631. 1753. B. demissa L. Syst. 

 ed. 10. 1118. 1759. B. melanotricha Brandegee, Univ. Calif. Publ. 

 Bot. 6: 61. 1914 (type from Cerro del Boqueron, Chiapas, Purpus 

 6662). Teresa; Teresita; Santa Teresita. 



Moist or wet thickets or forests, often a weed in cafetales or 

 other cultivated land, 600-1,800 m.; Alta Verapaz; Baja Verapaz; 

 Jalapa; Santa Rosa; Guatemala; Sacatepe"quez; Chimaltenango; Hue- 

 huetenango; Suchitepe"quez; Quezaltenango. Southern Mexico to 

 Costa Rica. Panama. West Indies and South America. 




10 FIELDIANA: BOTANY, VOLUME 24 



Slender annual plants, 2.5-6 dm. tall, the stems much-branched, often glandu- 

 lar-pubescent above; leaves ciliate, sparsely pilose or sometimes glabrous, ovate or 

 lanceolate, 4-10 cm. long, 1-5 cm. wide, acute or acuminate, the base attenuate; 

 petiole 0.5-3 cm. long; inflorescence racemose or flowers solitary; pedicels 5-15 mm. 

 long; calyx glandular-pubescent, 4-8 mm. long, (4-)5(-6)-lobed, the lobes 1.5-2 

 mm. long, shorter than the tube and unequal, acute; corolla blue or violet, occa- 

 sionally white, the throat yellow, the limb 1.2-1.5 cm. wide, the lobes retuse, short 

 and broad, the tube 1-1.8 cm. long; stamens 4, didynamous, inserted about 3 mm. 

 below the mouth of the tube; filaments about 2 mm. long, dilated upward and 

 ciliate, the upper ones curved and closing the throat of the corolla tube; style 

 included, 1.2-1.5 cm. long, abruptly curved at the apex; capsule ovoid, 5-8 mm. 

 long, surpassed by the accrescent calyx, the valves with appressed hairs; seeds 1 

 mm. long, foveolate-reticulate. 



This is a common ornamental plant throughout Central America. 

 It is doubtful whether the plant is native in Guatemala or only an 

 escape from cultivation, since it is usually found about dwellings or 

 in cultivated land. 



Called no-me-olvides and simpatica in El Salvador. 



BRUNFELSIA L. 



Shrubs or small trees, rarely suffrutescent; leaves alternate and entire, often 

 coriaceous, petiolate or subsessile; inflorescences cymose or consisting of an 

 axillary and solitary flower, the flowers large and showy; calyx campanulate- 

 tubular, 5-lobed; corolla salverform, slightly zygomorphic, white or scarlet or deep 

 purple, the limb broad and spreading, shallowly 5-lobed, the lobes imbricate in 

 bud, the posterior lobes outermost in bud; stamens 4, didynamous, included; fila- 

 ments subequally inserted in pairs above the middle of the corolla tube; anthers 

 longitudinally dehiscent, consisting of apparently 1 theca or 2 thecae and then 

 confluent, the thecae equal in size; ovary bilocular, the ovules numerous, the disk 

 inconspicuous; style included, smooth; stigma dilated or expanded and flattened, 

 bilobed; fruit a capsule, fleshy or coriaceous, apparently indehiscent or septicidal 

 and tardily dehiscent; seeds several and rounded; embryo slightly incurved. 



About 40 species in tropical America. Only one other species is 

 in Central America. Mr. T. Plowman is currently working on a 

 monograph of the genus. 



Corolla white, the tube 10-11 cm. long; leaves broadest above the middle; shrubs 

 or small trees B. nitida. 



Corolla scarlet or orange, the tube 3-4 cm. long; leaves broadest below the middle; 

 suffrutescent B. nyctaginoides. 



Brunfelsia nitida Benth. in DC. Prodr. 10: 201. 1846. Galdn. 



Cultivated for ornament in Pete"n. Native of the West Indies. 



We have seen only two collections from Guatemala. These collec- 




FIG. 3. Brunfelsia nydaginoides. A, habit, X 1 A; B, corolla dissected, X 2; 

 C, calyx dissected, X 2. 



11 




12 FIELDI ANA: BOTANY, VOLUME 24 



tions (Tun 609 and Lundell 24.17) probably represent escapes from 

 cultivation. 



Shrubs or small trees, glabrous throughout; leaves subcoriaceous and lustrous, 

 oblanceolate or obovate, 5-8.5 cm. long, 1.5-3 cm. wide, acute or acuminate, the 

 base attenuate, the petiole 3-4 mm. long; flowers axillary and solitary on short 

 bracteate branches shorter than the pedicels, or sometimes clustered; pedicels 5-10 

 mm. long; calyx 1-1.2 cm. long, parted to near the base, the lobes 7-8.5 mm. long, 

 lanceolate or elliptic, acute or acuminate; corolla white, the limb 3.3-5 cm. wide, 

 the lobes undulate, the tube slender, about 10 cm. long, inconspicuously ampliate 

 just below the mouth; filaments about 1 cm. long; anthers apparently one-celled; 

 style 9.5-10 cm. long; stigma dilated and bilobed; capsule about 1 cm. in diameter, 

 globose, smooth and glabrous; seeds about 5 mm. long, reticulate-foveate. 



A very attractive plant that should be more widely planted as an 

 ornamental. Called Golan de noche in Mexico (Chiapas). 



Brunfelsia nyctaginoides Standl. Field Mus. Bot. 22: 47. 1940 

 (type from Chiapas, Volcan de Tacana, Matuda 21+11}. 



Mixed or fir forests, 1,800-3,800 m.; El Progreso; San Marcos. 

 Mexico (Chiapas). 



Plants suffrutescent, usually scandent, the young stems pilose; leaves mem- 

 branaceous, ciliate, pilose, sometimes sparsely so above, ovate or ovate-elliptic, 

 4-10 cm. long, 2-3.5 cm. wide, acute or acuminate, the base attenuate, the petiole 

 0.5-1 cm. long; inflorescences cymose, few-flowered; pedicels pilose, mostly 1-1.5 

 cm. long; calyx 1 cm. long, lobed to the middle or more deeply so, the lobes pilose, 

 5-6 mm. long, ovate or lanceolate, acute; corolla scarlet, with the throat orange or 

 yellowish, or the tube orange and the lobes scarlet, the limb 2-2.5 cm. wide, the 

 tube 3-4 cm. long, gradually ampliate above; filaments 4-6.5 mm. long; anthers 

 2-celled, the cells confluent at maturity; style about 3 cm. long; stigma expanded 

 and flattened, slightly bilobed at the apex; capsule about 7 mm. in diameter, 

 globose, smooth and glabrous; seeds about 5 mm. long, reticulate-foveate. 



There has been some doubt regarding the generic disposition of 

 this species. See remarks by Standley and Steyermark in Field Mus. 

 Bot. 22: 376-377. 1940. 



CAPSICUM L. 



Herbs or suffrutescent plants or sometimes shrubs; leaves alternate, often in 

 pairs, entire or repand, petiolate; inflorescences axillary, consisting of 1-3 flowers, 

 the flowers pedicellate; calyx campanulate, truncate or with 5 minute lobes, 

 sometimes bearing 5 linear or subulate appendages just below the margin of the 

 calyx, slightly accrescent in age; corolla subrotate (in ours) or campanulate, white, 

 greenish-white, yellowish-white, yellow, or purple, the limb scarcely to shallowly 

 5-lobed, the lobes plicate or valvate-plicate in bud; stamens 5; filaments slender, 

 glabrous; anthers longitudinally dehiscent; ovary bilocular or rarely trilocular, the 

 ovules numerous, the disk inconspicuous; style slender, nearly always exserted; 




GENTRY & STANDLEY: FLORA OF GUATEMALA 13 



stigma clavate or slightly dilated; fruit a berry, usually very pungent, very juicy 

 or rarely almost dry, globose to oblong, sometimes inflated and much enlarged; 

 seeds numerous, laterally compressed, reticulate or nearly smooth; embryo curved 

 to subspiral. 



A genus of about 30 species, ranging from the southwestern 

 United States to South America and the West Indies, introduced in 

 tropical climates throughout the Old World. Certain species are of 

 great economic importance and are cultivated in temperate and trop- 

 ical regions in both hemispheres for their use both as a food and as a 

 condiment. The generic limits of Capsicum have been greatly im- 

 proved by the recent study of Witheringia by Hunziker (1969). 



Calyx bearing 5 linear or subulate appendages just below the margin of the 



truncate calyx. 



Leaves ovate to elliptic, the apex short-acuminate, pubescent throughout below, 

 the leaves often in pairs, the smaller ones similar in shape; calyx incon- 

 spicuously 5-ribbed, the calyx appendages 1.5-2.5 mm. long. . .C. ciliatwm. 

 Leaves narrowly lanceolate to narrowly elliptic, the apex narrowly long-acumi- 

 nate, pubescent only on the veins below, the leaves in pairs, the smaller 

 ones very different in shape and size; calyx conspicuously 5-ribbed, the 

 calyx appendages 3-5.5 mm. long C. lanceolatum. 



Calyx truncate or with minute lobes; calyx appendages absent. 



After first flowering node, one flower per node, rarely more; fruits globose or 



ovoid C. annuum var. aviculare. 



After first flowering node, two or more flowers per node, very rarely less; fruits 



ellipsoid-lanceolate or lanceolate C. frutescens. 



See D'Arcy and Eshbaugh, Baileya in press, for a key to the 

 species of Capsicum north of Colombia. 



Capsicum annuum L. Sp. PI. 188. 1753. C. petenense Standl. 

 Carnegie Inst. Wash. Publ. 461: 84. 1935 (type from La Libertad, 

 Pete"n, Lundell 3754). Chi-ic; chile bolita; chile chocolate; chile chulin; 

 chile diente deperro (Alta Verapaz) ; chile chocolate (Huehuetenango) ; 

 chile huaque (Chimaltenango and Huehuetenango) ; chile largo (Juti- 

 apa) ; chile zambo (Huehuetenango) . Widely known as chile or chili. 



Commonly cultivated in Guatemala, except at higher elevations, 

 mostly in small quantities in gardens. A native of tropical America, 

 that is now widely cultivated in temperate and tropical regions of 

 the world. The original distribution may never be known because 

 of its wide use as a food plant. Capsicum annuum var. annuum is 

 unknown as an escape from cultivation. It includes most of the cul- 

 tivated forms in Mexico and Central America. 



Chile, either fresh or dried, is used very commonly for flavoring 

 all kinds of meat and vegetable dishes, but not in such strength or 

 quantity as in Mexico. That country seems to be the center of in- 




14 FIELDIANA: BOTANY, VOLUME 24 



tensity of chile as a condiment. Its use in food in Central America 

 gradually decreases as one travels southward. 



Green chile is a favorite dish when available, the fresh fruits be- 

 ing used in large amounts. They are cooked alone by frying or in 

 other ways or mixed with eggs or vegetables. The well-known town 

 of San Martin Chile Verde in Quezaltenango received its name from 

 the fact that its inhabitants were particularly fond of chile verde. 



A condiment called chilmol, consisting of chile, vinegar, onions, 

 and various condiments, is in popular use for flavoring soup and other 

 dishes after they are placed on the table. Sweet peppers, chile duke, 

 are not very common but are obtainable in the larger markets. 

 Chiles stuffed with meat constitute a popular and often intensely 

 "hot" dish. A saucer of small fresh chiles, usually chiltepe, often is 

 placed on the dining table and one to three of them are placed in a 

 plate of caldo or soup. Chile leaves and young shoots are sometimes 

 boiled and eaten as a pot herb. 



Chile, both dry and fresh, is offered in large quantities in all the 

 Guatemalan markets, sometimes brought from remote regions of the 

 country by cargadores. Several varieties are always on sale. Chile 

 huaque is a common hot variety with long, narrowly conical, gradu- 

 ally tapering fruits; chile zambo is similar but with relatively thicker 

 fruits; chile pasa (which should mean merely dry chile) is a very 

 large form; chile chocolate has red, not very hot fruits; chile diente de 

 perro is small, green, and very pungent; chile de relleno is the large 

 sweet pepper, also called chile dulce; chile de zope and chile lengua de 

 gallina are small forms similar to chile diente de perro; chile bolito 

 has small globose fruits 1.5 cm. in diameter or somewhat larger; 

 chile chulin of Alta Verapaz has ovoid fruits about 2 cm. long; chi-ic 

 is a similar form of Coban. Chamborote is a form with very pale 

 green or almost white fruits, very irregular in form; chile ululte of 

 Alta Verapaz has a long and thick, very pungent fruit that is much 

 used in soup. The name chile is of Nahuatl derivation. The name 

 aji is only rarely used in Guatemala, although used in some parts of 

 Central America and particularly in Panama. It is a word of Antil- 

 lean derivation. 



Capsicum annuum L. var. aviculare (Dierb.) D'Arcy & Esh- 

 baugh, Phytologia 25: 350. 1973. C. indicum 5. microcarpon var. 

 aviculare Dierb. Handb. Med.-Pharm. Bot. 30. 1819. C. baccatum 

 auct., non L. 1753. C. minimum Miller, Gard. Diet. ed. 8. Capsi- 

 cum no. 10. 1768. C. annuum var. minimum Heiser, Ciencia y Nat. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 15 



7: 52. 1964. Chile de montana (Huehuetenango) ; chiltepe (Izabal; 

 Jutiapa; Santa Rosa; Zacapa). 



Wet, moist or dry thickets, sometimes in rocky forests, 1,200 m. 

 or less; Pete"n; Izabal; Zacapa; Chiquimula; Jalapa; Solola; Huehue- 

 tenango; Jutiapa; Santa Rosa; Escuintla; Retalhuleu; Suchitepe- 

 quez. British Honduras. Florida westward to Arizona and south 

 throughout Mexico and Central America to Colombia. West Indies. 



An herb or shrub 1-2.5 m. tall, densely branched, the branches sparsely 

 pubescent or glabrous; leaves slender petiolate, solitary or often in pairs, ovate to 

 lanceolate or elliptic, apex acuminate, base attenuate, the larger leaves mostly 

 3.5-8 cm. long, 1-3 cm. wide; flowers solitary or rarely 2-3; calyx truncate or with 

 minute lobes; corolla white, rarely greenish; berry red, globose or ovoid, apex 

 obtuse, 5-10 mm. in diameter, rarely exceeding 15 mm. in length; seeds cream to 

 yellow. 



The name C. annuum var. baccatum L., has been applied incor- 

 rectly to this taxon. See Eshbaugh, Taxon 17: 51-52. 1968, for ex- 

 planation concerning this matter. 



This is the common and widely distributed, spontaneous variety 

 of the cultivated pepper, Capsicum annuum. The fruits are intensely 

 pungent. They are so hot that a single small one will sometimes make 

 a bowl of soup quite unpalatable for many people. They are used 

 chiefly for making the condiment chiltepe. Large quantities of them 

 are sold in the markets, but it is more convenient to have a bush 

 just outside the kitchen. They are much sought after by birds. 

 The fruits undoubtedly have been used for flavoring food in Guate- 

 mala for a very long time. 



Commonly referred to as "bird pepper." The Maya names of 

 Yucatan are max and maaxic. Called chile; chile bravo; chile de per- 

 ro; chile silvestre, and chiltepe in Honduras. 



Capsicum cilia turn (H.B.K.) Kuntze, Rev. Gen. 2: 450. 1891. 

 Witheringia ciliata H.B.K. Nov. Gen. Sp. PL 3: 15. 1818. Brachistus 

 pringlei Wats. Proc. Am. Acad. Sci. 25: 159. 1890. Capsicum pring- 

 lei Macbr. & Standl. Field Mus. Bot. 11: 173. 1936. 



Moist or dry forests, 780-1,650 m.; Guatemala; Sacatepequez; 

 Huehuetenango; Santa Rosa; Escuintla. Mexico. Honduras. Nica- 

 ragua. South America. 



A slender shrub 2-4.5 m. tall, the branches pubescent with small incurved 

 hairs, sometimes rather densely pubescent; leaves solitary or often in pairs, then 

 unequal in size and similar in shape, pubescent above with slightly incurved hairs, 

 the cells of the hairs not noticeably swollen, pubescent throughout below, the hairs 




B 



FIG. 4. Capsicum frutescens. A, habit, X 1 A; B, flower, X 5; C, anther, abax- 

 ial view, X 5; D, mature fruit, X 2. 



16 




GENTRY & STANDLEY: FLORA OF GUATEMALA 17 



more dense, especially on the veins, the larger leaves ovate to elliptic, 3-12 cm. 

 long, 1-4 cm. wide, apex short-acuminate, base attenuate, often narrowly winged; 

 petioles 0.5-2.5 cm. long, pubescent with incurved hairs; smaller leaves 1.5-4 cm. 

 long, 0.7-2.5 cm. wide, apex acute or acuminate, short petiolate or subsessile; 

 flowers mostly solitary or sometimes 2, the pedicels very slender, 5-15 mm. long, 

 sparsely pubescent; calyx 1-1.5 mm. long, only slightly 5-ribbed, sparsely to 

 densely pubescent, with 5 linear appendages, these 1.5-2.5 mm. long; corolla yellow 

 5-6.5 mm. long, glabrous, shallowly lobed, the lobes about 1 mm. long; filaments 

 2-2.5 mm. long; anthers 2 mm. long; style 4-4.5 mm. long; berry red, globose, 

 juicy, 6-8 mm. in diameter; seeds 2-2.5 mm. long, yellow. 



We are following A. T. Hunziker, Kurtziana 6: 250-259. 1971, in 

 applying this name to Central American plants. He also reports 

 that this species never has pungent fruits. 



Capsicum frutescens L. Sp. PI. 189. 1753. Chiltepe. 



Known from only one collection in Guatemala, La Libertad, and 

 vicinity, Aguilar 199, which may possibly represent cultivated ma- 

 terial. The label does not specify the habitat. A widely distributed 

 weed. British Honduras. Southern United States. Mexico to South 

 America. West Indies. It is grown throughout the islands of Poly- 

 nesia and has been reported from India (Heiser and Smith, 1953). 



A densely branched herb or shrub, the branches sparsely pubescent or glabrous; 

 leaves slender petiolate, solitary or often in pairs, ovate to ovate-lanceolate, apex 

 long acuminate or acuminate, base attenuate, the larger leaves 4-12 cm. long, 

 1-4.5 cm. wide; flowers 2 or more at a node, rarely 1 ; calyx truncate or with minute 

 lobes, prominent constriction lacking between base of the calyx and pedicel; 

 corolla greenish-white or yellowish-white; berry red, ellipsoid-lanceolate or lance- 

 olate, apex acute, 1.5-3 cm. long, 0.5-1.2 cm. in diameter; seeds cream to yellow. 



Capsicum frutescens is apparently rare in Guatemala, at least as 

 an escape from cultivation. It is often seen growing around houses, 

 where it is planted for its intensely pungent fruits. 



The larger fruited plants are widely cultivated in tropical and 

 subtropical regions of the world. 



Capsicum lanceolatum (Greenm.) Morton & Standl. Field 

 Mus. Bot. 22: 272. 1940. Brachistus lanceolatus Greenm. ex Donn.- 

 Sm. Bot. Gaz. 37: 212. 1904 (type from Chucaneb, Alta Verapaz, 

 Donn.-Sm. 1837) . Pajarito del rio; yerba de pajarito (Quezaltenango) . 



Moist or wet forests sometimes in mixed forests, 500-2,000 m., 

 rarely higher; Alta Verapza; Baja Verapaz; Sacatepe"quez ; Chimal- 

 tenango; Solola; Quezaltenango; Huehuetenango; Suchitepe"quez ; 

 San Marcos. Southern Mexico (Chiapas). Honduras. 




18 FIELDIANA: BOTANY, VOLUME 24 



A slender shrub or sometimes a slender tree or suffrutescent, 1-5 m. tall, the 

 branches pubescent with small incurved hairs, sometimes nearly glabrous; leaves 

 in pairs, very unequal in size and different in shape, sparsely pubescent above 

 with multicellular hairs, the cells slightly swollen, these hairs readily discernible, 

 pubescent on the veins below, the hairs incurved or glabrous, the larger leaves 

 mostly inequilateral, narrowly lanceolate to narrowly elliptic, 5-15 cm. long, 

 1.5-3 (-4) cm. wide, apex narrowly long-acuminate, base attenuate; petioles 0.5-1 

 cm. long, pubescent with incurved hairs or glabrous; smaller leaves ovate to sub- 

 rotund, 1-5 cm. long, 0.8-2.5 cm. wide, apex obtuse, short petiolate or subsessile; 

 flowers mostly solitary, or sometimes 2, the pedicels very slender, 1.5-3.5 cm. long, 

 glabrous; calyx 1.5-2 mm. long, conspicuously 5-ribbed, sparsely pilosulous or 

 glabrous, with 5 subulate appendages, these 3-5.5 mm. long, subequal, rather thick; 

 corolla white to pale yellow, 6.5-10.5 mm. long, glabrous, shallowly lobed, the 

 lobes 1-2 mm. long; filaments 2-2.5 mm. long; anthers 1.5 mm. long; style 4.5-5 

 mm. long; berry orange-red to red, globose, juicy, 7-10 mm. in diameter; seeds 

 2-2.5 mm. long, yellow. 



Capsicum pubescens Ruiz & Pavon, Fl. Peruv. 2: 30. 1799. 

 C. guatemalense Bitter, Fedde Repert. Sp. Nov. 20: 377. 1924 (type 

 from Las Nubes, Suchitepe"quez, Bernoulli & Cario 2339). Chile 

 cuarto caldos; cuile de caballo; chile garrapata; chile siete caldos. 



Cultivated frequently for its very pungent fruit; grown in Alta 

 Verapaz, Chimaltenango, Quezaltenango, San Marcos, and Solola, 

 and probably other departments as well. Also cultivated in Hon- 

 duras and Costa Rica. It is a native of South America, probably 

 Peru. 



Capsicum pubescens is the most easily recognized species of the 

 cultivated peppers. It is readily separated by its purple corolla, 

 black seeds, and rugose and glabrous leaves. The herbage is often 

 densely pubescent. 



Called chile petenero in Honduras. 



We are following Bukasov, Bull. App. Bot. Gen. and PL, Suppl. 

 47: 261-273. 1930, and Eshbaugh (personal communication) in treat- 

 ing C. guatemalense as a synonym here. 



CESTRUM L. 



References: P. Francey, Monographic du genre Cestrum L., Can- 

 dollea 6: 46-398. 1935; 7: 1-132. 1936. C. V. Morton, The genus 

 Cestrum in Guatemala, J. Arnold Arbor. 17: 341-349. 1936. 



Shrubs or trees, rarely scandent; leaves alternate and entire, rarely in pairs, 

 petiolate, axillary leaves sometimes present and persistent; inflorescences axillary 

 or terminal, paniculate, racemose, or fasciculate, or the flowers rarely solitary, the 

 flowers pedicellate or sessile, the bracts of the inflorescences mostly small or some- 




GENTRY & STANDLEY: FLORA OF GUATEMALA 19 



times large and leaf-like; calyx campanulate to tubular, 5-lobed; corolla funnel- 

 form or salverform, variously colored, the limb 5-lobed, the lobes valvate in bud or 

 somewhat valvate-plicate at the base of the lobes, the tube ampliate above; 

 stamens 5, included; filaments equal, variously inserted on the corolla tube, often 

 with a small appendage at their bases; anthers longitudinally dehiscent; ovary 

 bilocular, short-stipitate, the ovules few, the disk inconspicuous or moderately 

 thick; style filiform, included or shortly exserted; stigma entire or bilobed; fruit a 

 berry, often juicy, short-stipitate; seeds few, compressed, rounded, or angulate; 

 embryo straight or slightly curved. 



A genus of some 250 species all in tropical America. Several 

 other species are known from southern Central America. A few 

 species are cultivated for their fragrant flowers. 



Pubescence of stellate and/or branched hairs. 



Filaments geniculate at their bases, appendages absent; corolla tube glabrous 



externally (in ours) C. lanatum. 



Filaments relatively straight, not geniculate at their bases, appendages present; 



corolla tube pubescent externally C. mortonianum. 



Pubescence of simple hairs or hairs absent. 



Inflorescence axillary, consisting of a single long-pedunculate flower. 



Cestrum sp. 



Inflorescence axillary and/or terminal, 2-many flowered, not consisting of a 

 single long-pedunculate flower. 



Corolla tube pubescent externally C. dasyanthum. 



Corolla tube glabrous externally. 



Filaments with appendages at their bases, these sometimes inconspicuous. 

 Corolla tube 9.5-11 mm. long; leaves with a small tuft of hairs in the 



axils of the mid-vein below C. dumetorum. 



Corolla tube 14-23 mm. long; leaves without a small tuft of hairs in the 



axils of the mid-vein below. 



Leaves narrow, lanceolate-linear or very narrowly lanceolate, 1-2.5 



cm. wide, rarely wider, the larger leaves 4 to 10 times as long 



as wide; flowers borne in fasciculate inflorescences. 



Calyx tube green, sparsely glandular externally, the lobes lanceolate, 



0.8-1.5 mm. long; corolla greenish or greenish-yellow; axillary 



leaves often present C. luteovirescens. 



Calyx tube dark purple (at least when dried), glabrous, the lobes 

 short-mucronate, 0.5 mm. long or less; corolla white; axillary 



leaves always absent C. skutchii. 



Leaves broad, not lanceolate-linear or very narrowly lanceolate, the 



larger leaves rarely less than 3 cm. wide, the larger leaves 2 to 4 



times as long as wide; flowers borne in racemose or paniculate 



inflorescences. 



Axillary leaves present; calyx and rachis of the inflorescences purple 



(at least when dried) C. elegantissimum, 



Axillary leaves absent; calyx and rachis of the inflorescences green. 

 Calyx 5-6.5 mm. long, the lobes prominent; filaments 4-6.5 mm. 



long; corolla orange C. aurantiacum. 



Calyx 1.5-3.5 (-4) mm. long, the lobes nearly always inconspicu- 

 ous; filaments 2-3.5 mm. long; corolla greenish or greenish- 

 yellow. 



Appendages of the filaments densely pubescent . . . . C. franceyi. 

 Appendages of the filaments glabrous or only rarely incon- 

 spicuously pubescent in C. nocturnum. 




20 FIELDIANA: BOTANY, VOLUME 24 



Mature stems and petioles puberulent; leaves puberulent on 

 the mid-vein below, at least near the base; petioles 2-4 



mm. long C. cobanense. 



Mature stems and petioles glabrous; leaves glabrous; petioles 



5-15 mm. long. 



Inflorescences lax, dichotomously branched, the rachis 



slender; pedicels long, 1-1.5 mm. long. . .C. evanidum. 



Inflorescences erect, not dichotomously branched, the rachis 



stout; pedicels short, 0.5 mm. long or less. 

 Rachis of the inflorescences shorter than the subtending 

 petiole; corolla tube 14 mm. long; fruit with one seed. 



C. glanduliferum. 



Rachis of the inflorescences greatly surpassing the petiole ; 

 corolla tube 17-20 mm. long; fruit with five seeds. 



C. nocturnum. 



Filaments without appendages at their bases, although sometimes swollen 

 and geniculate. 



Corolla lobes 7-9.5 mm. long; scandent shrubs C. scandens. 



Corolla lobes 1-5 mm. long; erect shrubs or trees. 

 Leaves pubescent below; inconspicuously so on the veins in C. 



fraternum. 



Calyx lobes long, 2-5 mm. long; flowers borne in fasciculate inflores- 

 cences; filaments 9.5-11 mm. long C. regelii. 



Calyx lobes short, 0.8 mm. long or less; flowers borne in racemose or 



paniculate inflorescences, sometimes in fasciculate inflorescences 



in C. alter ni folium; filaments 0.5-5 mm. long. 



Leaves inconspicuously pubescent below, the hairs only on the 



veins; filaments 4.5-5 mm. long, pilosulous at their bases; 



pedicels 3-5 mm. long C. fraternum. 



Leaves evidently pubescent below, at least when young, the hairs 



on the veins and lamina; filaments 0.5-3 (-3.5) mm. long, 



glabrous or pilosulous at their bases; pedicels 0.5-3 mm. long. 



Inflorescences fasciculate; corolla lobes 5-6 mm. long; calyx 2-3 



mm. long; fruit purplish; lowland species, below 500 m. 



C. alternifolium. 



Inflorescences racemose or paniculate; corolla lobes 3-4 mm. 

 long; calyx 3.5-4 mm. long; fruit white; ranging from 



2,000-3,800 m C. guatemalense. 



Leaves glabrous below. 



Corolla tube short, 10-12 mm. long. 



Pedicels long, 3-5.5 mm. long; corolla tube 2 mm. wide at the 



middle; above 2,500 meters C. formosum. 



Pedicels short, 1 mm. long or less; corolla tube 1 mm. wide at the 



middle; below 1,500 meters. 



Leaves broadest at the middle or above, cuneate at the base; 

 filaments 3.5-4 mm. long, pilosulous at their bases; rachis 

 of the inflorescences equalling or shorter than the petioles. 



C. megalophyllum. 



Leaves broadest at their bases, mostly rounded at the base; 

 filaments 1-2 mm. long, glabrous at their bases; rachis of 

 the inflorescences mostly greatly exceeding the petioles. 



C. racemosum. 

 Corolla tube long, 14-20 mm. long. 



Calyx 5-6.5 mm. long, the lobes 1-3 mm. long, long mucronate or 



subulate; corolla orange C. aurantiacum. 



Calyx 2.5-4 mm. long, the lobes 0.5 (-1.5) mm. long, if exceeding 

 0.5 mm. in length, the lobes ovate-triangular in shape; corolla 

 pale yellow or greenish-yellow. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 21 



Filaments 6-7 mm. long, geniculate at their bases; corolla tube 

 glabrous internally; pedicels 3.5-6.5 mm. long.C. anagyris. 



Filaments 2-4.5 mm. long, relatively straight at their bases, not 

 geniculate; corolla tube pubescent internally; pedicels 0.3- 

 3.5 mm. long. 



Pedicels short, 0.3 mm. long; corolla lobes 5 mm. long; calyx 

 minutely puberulent C. franceyi. 



Pedicels long, 1.5-3.5 mm. long; corolla lobes 2-3 mm. long; 

 calyx glabrous C. pacayense. 



Oestrum alternifolium (Jacq.) 0. E. Schulz in Urban, Symbol. 

 Antill. 6: 270 1909. Ixora alternifolia Jacq. Enum. Syst. Plant. 12. 

 1760. 



In thickets, 250-400 m.; Zacapa; Chiquimula. Southern Mexico. 

 Honduras; El Salvador; Costa Rica. Panama and West Indies. 

 Northern South America. 



A slender shrub, 0.5-3.5 m. tall, the young branches pilosulous; leaves ovate to 

 ovate-lanceolate, 4.5-9 cm. long, 2-3.5 cm. wide, glabrous above, pilosulous on 

 the veins or throughout below, apex acute, base obtuse; petioles 0.4-0.6 cm. long, 

 pilosulous; inflorescences axillary and terminal, fasciculate, few-flowered, the bracts 

 linear to lanceolate, pilosulous; pedicels 0.5 mm. long or less, glabrous; calyx 2-3 

 mm. long, glabrous except for the ciliolate lobes, the lobes deltoid to triangular, 

 0.8 mm. long or less; corolla greenish, the tube 17-21 mm. long, glabrous externally 

 and internally, ampliate just below the mouth of the corolla tube, the lobes 5-6 

 mm. long, ovate to ovate-lanceolate, marginally pubescent externally; filaments 

 0.5-2 mm. long, glabrous, appendages absent; style 18-22 mm. long; fruit 9-12 mm. 

 long, purplish; seeds about 8, 5-6 mm. long. 



Oestrum anagyris Dunal in DC. Prodr. 13., pt. 1: 608. 1852. 

 Oak or pine forests, 2,220-3,000 m.; Totonicapan; Quezaltenango. 

 Southern Mexico. 



A slender shrub 1-2 m. tall, the branches glabrous; leaves lanceolate to elliptic, 

 5-10 cm. long, 1.5-3.5 cm. wide, glabrous above and below, apex acuminate or 

 acute, acute or short attenuate at the base; petioles 1-1.5 cm. long; inflorescences 

 lax, axillary and terminal, paniculate, glabrous, many-flowered, the bracts linear, 

 caducous; pedicels 3.5-6.5 mm. long, glabrous; calyx 3-4 mm. long, cleft on two 

 sides one-fourth to one-third length of the tube, glabrous except for the ciliolate 

 lobes, the lobes 0.2-0.5 mm. long, deltoid and very short-mucronate; corolla 

 greenish-yellow, the tube 14-17 mm. long, ampliate above from just above middle 

 of the tube, the lobes 3-4 mm. long, ovate, marginally pubescent externally; fila- 

 ments 6-7 mm. long, swollen, canaliculate and slightly geniculate at their bases, 

 glabrous, appendages absent, the corolla tube glabrous internally; style 13.5-16 

 mm. long; fruit 10-12 mm. long, white; seeds 2-4, 3.5-5 mm. long. 



Oestrum aurantiacum Lindley, Bot. Reg. 30: 71, misc. no. 

 65. 1844; 31: t. 22. 1845. C. chaculanum Loes. Verh. Bot. Ver. Prov. 

 Brandenb. 65: 97. 1923. C. paucinervium Francey, Candollea 6: 101. 




22 FIELDIANA: BOTANY, VOLUME 24 



1935 (type from San Miquel Uspantan, Quiche, Heyde & Lux 3135}. 

 C. aurantiacum var. chaculanum Francey, 1. c. 104. Chipin (Gua- 

 temala); hediondillo, huele de noche, pay-te (Huehuetenango) ; huevo 

 ( Jalapa) . 



Moist thickets or forests, often in pine-oak forests, 1,000-2,600 

 m.; Alta Verapaz; Baja Verapaz; Zacapa (described from a culti- 

 vated plant grown in England from seeds sent by Skinner from Chi- 

 malapa); Chiquimula; Jalapa; Guatemala; Sacatepe"quez ; Chimal- 

 tenango; Solola; Quiche"; Quezaltenango; Huehuetenango; Jutiapa. 

 Southern Mexico to Nicaragua. 



A shrub 1.5-6.5 m. tall, sometimes a tree 8.5 m. tall, the branches glabrous 

 or sparsely pilosulous; leaves ovate to elliptic, 7-17 cm. long, 2.5-5.5 cm. wide, 

 glabrous above and below, apex acute or short-acuminate, acute to obtuse or some- 

 times short-attenuate at the base; petioles 1-3 cm. long, glabrous; inflorescences 

 terminal or axillary, paniculate or racemose, several- to few-flowered, the rachis 

 pilosulous or glabrous, the bracts linear, caducous, the flowers sessile or subsessile 

 or the pedicels as much as 1.5 mm. long; calyx tubular, 5-6.5 (-9) mm. long, 

 glabrous except for the ciliolate lobes, the lobes subulate, long-mucronate or 

 rounded and long-mucronate, (0. 7-) 1-2 (-3) mm. long, the lobes extending down- 

 ward on the calyx tube as five nerves; corolla orange or rarely yellow, the tube 

 17.5-20 mm. long, the corolla lobes 3-3.5 (-5.5) mm. long, ovate or lanceolate, 

 marginally papillate-puberulent externally; filaments 4-6.5 mm. long, swollen and 

 geniculate, canaliculate or subappendiculate, pilosulous at their bases, the staminal 

 traces pilosulous; style 16.5-18.5 mm. long; fruit 8-12 mm. long, white; seeds 

 7-9, 3-5 mm. long. 



In Huehuetenango the crushed leaves are used with soap and 

 cold water to remove stains from clothing (fide Steyermark) . 



Oestrum aurantiacum is a common shrub in Guatemala usually 

 collected at middle elevations. 



Cestrum cobanense Francey, Candollea 6: 372. 1935. 

 Wet forests, 1,500 m.; Alta Verapaz (type from Coban, Tuerck- 

 heim II 2372}. Endemic. Known only from the type collection. 



A shrub, the branches slender, puberulent; leaves lance-ovate or narrowly 

 elliptic, 5.5-9 cm. long, 1.5-2 cm. wide, glabrous above, sparsely puberulent only 

 on the midvein below, apex acuminate, rounded or very shortly attenuate at the 

 base; petioles very short, 2-4 mm. long, puberulent; inflorescences terminal or 

 axillary, paniculate, puberulent, the bracts linear, puberulent, caducous; pedicels 

 0.5-1 mm. long; calyx 1.5 mm. long, glabrous except for the ciliolate lobes, the 

 lobes 0.5 mm. or less long, deltoid; corolla tube 16.5-18 mm. long, glabrous exter- 

 nally, ampliate just below the throat of the corolla tube, the lobes 3-4 mm. long, 

 lanceolate, marginally papillate-puberulent; filaments 2.5-3.5 mm. long, append- 

 aged at their bases, the appendages about 0.2 mm. long, entire and glabrous; style 

 15.5-18.5 mm. long; fruit unknown. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 23 



Cestrum cobanense is distinguished from other species in Gua- 

 temala by its very short petioles, puberulent branches, and ap- 

 pendiculate filaments. 



Cestrum dasyanthum Donn.-Sm. Bot. Gaz. 23: 11. 1897. 

 Chipshi; chipshin (Huehuetenango) . 



Moist forests, 2,300-3,000 m.; Huehuetenango (type from be- 

 tween San Martin and Todos Santos, Nelson 3622}. Endemic. 



A shrub 1.5-6 m. tall, the branches pilosulous, the hairs brownish; leaves ovate 

 to elliptic, 5-13.5 cm. long, 2-4.5 cm. wide, pilosulous above and below, more 

 densely so on the veins below, apex acute, obtuse at the base, the petioles 1-2.5 cm. 

 long; axillary leaves early deciduous, nearly orbicular, 4.5-6 mm. long, 4-5 mm. 

 wide, subsessile, apex obtuse; inflorescences axillary and terminal, paniculate or 

 racemose, few- to many-flowered, the rachis pilosulous, the hairs brownish, the 

 bracts linear, caducous; pedicels 0.5-1 mm. long, pilosulous, sometimes sparsely 

 so; calyx 5-6 mm. long, pilosulous, the lobes about 0.5 mm. long, rounded and 

 mucronate or mucronate, the calyx tube cleft down both sides one-third to nearly 

 one-half to the base; corolla yellow, the tube 11.5-15 mm. long, pilosulous exter- 

 nally, gradually ampliate above, the lobes 4-4.5 mm. long, ovate, pilosulous or 

 villosulous externally; filaments 3.5-4 mm. long, slightly swollen, geniculate and 

 pilosulous at their bases, the staminal traces pilosulous, appendages absent; anthers 

 about 1 mm. long; style 10.5-13 mm. long; fruit about 12 mm. long; seeds 13, 

 3.5-4 mm. long. 



The pubescent corolla tube readily separates this species from 

 all others in Guatemala except Cestrum mortonianum which has 

 branched and stellate hairs. 



Cestrum dumetorum Schlecht. Linnaea 7: 61. 1832. Huele 

 de noche (Jutiapa); huele de noche negro (Guatemala). 



Moist or wet thickets, 600-1,205 m.; Guatemala; Jutiapa; Santa 

 Rosa. Southern Mexico to Costa Rica. 



A shrub or small tree, 2-3 m. tall, the branches pilosulous or glabrous; leaves 

 ovate-lanceolate or elliptic-lanceolate, 5-14.5 cm. long, 2-6.5 cm. wide, glabrous 

 or puberulent above, puberulent on the veins below and with a small tuft of hairs 

 in the axis of the secondary veins, the hairs sometimes inconspicuous; petioles 

 1.5-2.5 cm. long; axillary buds lanate; inflorescences terminal and axillary, 

 paniculate, the sessile flowers several to many; calyx 4-6 mm. long, villosulous- 

 pilosulous, the tube with one or two clefts about one-third the length of the tube, 

 shallowly 5-lobate, the lobes unequal, 0.5 mm. long or less; corolla pale green, the 

 tube 9.5-11 mm. long, about twice as long as the calyx, glabrous externally, 

 gradually ampliate above, the lobes 5-6 mm. long, lanceolate, marginally puberu- 

 lent externally; filaments 1-2 mm. long, appendaged at their bases, the appendages 

 0.2-0.5 mm. long, entire and glabrous, the staminal traces pilosulous; style 10- 

 11.5 mm. long; fruit 5-8 mm. long; seeds 4, 3-4 mm. long. 




24 FIELDIANA: BOTANY, VOLUME 24 



Cestrum dumetorum is a very distinctive species in which the calyx 

 tube is about one-half the length of the corolla tube, filaments lack- 

 ing appendages, and leaves with a small tuft of hairs in the axils of 

 the mid-vein below. The lanate axillary buds are also a character- 

 istic of this species. 



Cestrum elegant issimum Morton, J. Arnold Arbor. 17: 347. 

 1936. Cerezo bianco (Quezaltenango) . 



Moist dense forests or mixed forests, 1,200-2,000 m.; Solola; Que- 

 zaltenango; Suchitepe"quez (type from Finca Moca, Skutch 2071). 

 Endemic. 



A slender shrub 1-5 m. tall, the branches glabrous; leaves glabrous above and 

 below, oblong lanceolate to oblong-elliptic, sometimes lanceolate, 6-17.5 cm. long, 

 1.5-4.5 cm. wide, apex acuminate, rounded or obtuse at the base, the petioles 

 short, 5-10 mm. long; axillary leaves orbicular or lanceolate, 0.3-1.5 cm. long, ses- 

 sile and somewhat clasping; inflorescences terminal, paniculate, lax, few- to many- 

 flowered, the rachis dark purple, pilosulous to inconspicuously puberulent, the 

 bracts dark purple, linear, caducous; pedicels glabrous or inconspicuously puberu- 

 lent, 1-1.5 mm. long; calyx 2-3 mm. long, dark purple, glabrous or inconspicu- 

 ously and sparsely puberulent, the lobes deltoid and acuminate, about 0.5 mm. 

 long, ciliolate; corolla greenish or greenish-yellow, the tube 18.5-21 mm. long, 

 glabrous externally and internally, ampliate just below the mouth of the corolla 

 tube, the lobes 5-8 mm. long, narrowly lanceolate or oblong-lanceolate, marginally 

 papillate-puberulent externally; filaments 2.5-3.5 mm. long, appendaged at their 

 bases, appendages glabrous and entire; style 17-19 mm. long; fruit 1-1.5 cm. long; 

 seeds 4, 6-7 mm. long. 



The axillary leaves and appendiculate filaments distinguish C. 

 elegantissimum from all other Oestrums in Guatemala except C. 

 luteovirescens, which has very narrow leaves and a fasciculate in- 

 florescence. 



Cestrum evanidum Morton, J. Arnold Arbor. 17: 346. 1936. 



Moist forests or dense mixed forests, 1,300-1,600 m.; Sacatepe"- 

 quez; Chimaltenango; Quezaltenango; Suchitepe"quez (type from 

 Finca Moca, Skutch 2071 A) ; San Marcos. Endemic. 



A slender shrub or a small tree, 2.5-6.5 m. tall, the young branches incon- 

 spicuously puberulent, soon glabrate; leaves ovate-lanceolate to narrowly elliptic 

 or elliptic-oblong, 8-19 cm. long, 2-5.5 cm. wide, glabrous above and below, apex 

 acuminate, rounded to obtuse at the base; petioles 0.5-1.5 cm. long, glabrous; 

 inflorescences lax, terminal or axillary, paniculate, dichotomously branched, 

 glabrous, the rachis slender, the bracts minute, caducous; pedicels 1-1.5 mm. 

 long, glabrous; calyx green, 2-2.5 mm. long, the lobes about 0.5 mm. long, deltoid, 

 apex pilosulous; corolla green or greenish-white, the tube slender, 20-23 mm. 

 long, gradually ampliate upwards, the lobes 4.5-5 mm. long, lance-oblong, exter- 




GENTRY & STANDLEY: FLORA OF GUATEMALA 25 



nally marginally papillate-puberulent; filaments 2.5-3.5 mm. long, appendaged at 

 their bases, the appendages deeply parted to entire, glabrous; anthers about 1 

 mm. long; style 19-22 mm. long; fruit about 6 mm. long; seeds 5, 3-5 mm. long. 



Cestrum evanidum is probably closely related to C. nocturnum. It 

 is separated from that species by the lax, dichotomously branched 

 inflorescence and pedicellate flowers. 



Cestrum formosum Morton, J. Arnold Arbor. 17: 348. 1936. 



In thickets or pine-fir forests, 2,665-3,200 m. ; Quiche" (type from 

 south slope of Volcan Atitlan, Skutch 2143); Solola; Quezaltenango. 

 Endemic. 



A shrub 4.5-5.5 m. tall, sometimes a tree 8 m. tall, the branches glabrous; 

 leaves elliptic or lanceolate, 6-12 cm. long, 1.5-3.5 cm. wide, glabrous above and 

 below, apex acuminate, acute or obtuse at the base; petioles long, 1.5-2.5 cm. 

 long, glabrous; inflorescences terminal and axillary, paniculate or racemose, 

 glabrous, few-flowered, the bracts linear; pedicels 3-5.5 mm. long; calyx 4-5 mm. 

 long, green, glabrous except for the ciliolate lobes, the lobes 1 mm. long or less, 

 unequal, the tube with two clefts about one-fourth to one-third the length of the 

 tube; corolla greenish-yellow, the tube 10-11 mm. long, glabrous externally, 

 gradually ampliate upwards, the lobes 4-5 mm. long, broadly ovate and mucro- 

 nate, marginally puberulent externally; filaments 1.5 mm. long, glabrous, the 

 staminal traces pilosulous, appendages absent; style 11-12 mm. long; fruit 7 mm. 

 long; seeds 5, 4 mm. long. 



Cestrum franceyi Morton, J. Arnold Arbor. 17: 345. 1936. C. 

 pedunculatum Francey, Candollea 7: 66. 1936, non Sesse* & Mocino, 

 1894. 



Known only from the type collection, Pansamala, Alta Verapaz, 

 Tuerckheim 833). Endemic. 



A shrub ('.')> the branches minutely puberulent; leaves elliptic-lanceolate 

 or ovate-lanceolate, 9-10.5 cm. long, 2.5-4.5 cm. wide, glabrous above and below, 

 apex acuminate or cuspidate, rounded at the base; petioles 6-11 mm. long; inflor- 

 escences axillary and terminal, 5.5-10.5 cm. long, long pedunculate, the rachis 

 minutely puberulent, the bracts linear, minutely puberulent, the flowers short 

 pedicellate, the pedicels 0.3 mm. long; calyx 3.5 mm. long, minutely puberulent 

 externally, the lobes ovate-triangular, 0.5-1.5 mm. long, the margin glabrous; 

 corolla tube 17.5 mm. long, the corolla lobes 5 mm. long, ovate or triangular- 

 ovate, marginally pilosulous externally, glandular internally; filaments 3 mm. 

 long, gibbose or appendaged and densely pilosulous at their bases, the staminal 

 traces pilosulous; style 17.5 mm. long; fruit unknown. 



Cestrum fraternum Morton, J. Arnold Arbor. 17: 345. 1936. 

 Moist thickets, 2,340 m.; Quiche" (type from Nebaj, Skutch 1682). 

 Endemic. Known only from the type collection. 




26 



FIELDIANA: BOTANY, VOLUME 24 



FIG. 5. Cestrumformosum. A, habit, X 3 ->; B, flower, X 2; C, calyx dissected, 

 X2. 



A shrub 2.5 m. tall, the young branches puberulent, soon glabrate; leaves 

 narrowly elliptic, 7.5-17.5 cm. long, 3-5.5 cm. wide, glabrous above, sparsely 

 puberulent on the veins below, apex acuminate, the base acute or obtuse; petioles 

 1.5-2 cm. long, glabrous; inflorescences terminal and axillary, paniculate, puberu- 

 lent, the bracts minute, caducous; pedicels 3-5 mm. long; calyx 3-3.5 mm. long, 

 green, the lobes about 0.5 mm. long, broadly deltoid, slightly mucronate, ciliolate; 

 corolla pale yellow, the tube sometimes tinged with maroon, 16-17 mm. long, 

 glabrous externally, gradually ampliate above, the lobes 3.5-4 mm. long, broadly 

 ovate and mucronate, inconspicuously papillate externally; filaments 4.5-5 mm. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 27 



long, swollen and pilosulous at their bases, the staminal traces pilosulous; append- 

 ages absent; style 17 mm. long; fruit unknown. 



Cestrum glanduliferum Kerber ex Francey, Candollea 6: 386. 

 1935. 



Moist forests, 1,050 m.; Chimaltenango, near Sibaja, Standley 

 62283. Known in Guatemala only from this collection. Southern 

 Mexico. 



A shrub or tree, 6.5 m. tall, the branches glabrous (in ours) or sparsely and 

 minutely glandular; leaves elliptic-oblong or elliptic-lanceolate, 11-26.5 cm. long, 

 3.5-10.5 cm. wide, subcoriaceous, glabrous above, minutely glandular-puberulent 

 below, (glabrous below in ours), apex acute or short-acuminate, narrowly rounded 

 to acute at the base; petioles 1-2 cm. long; inflorescences axillary, racemose, few- 

 flowered, the rachis pilosulous, and shorter than the petioles, the bracts linear, 

 pilosulous, persistent or caducous, the flowers subsessile and short pedicellate; 

 calyx 3.5-4 mm. long, glabrous or inconspicuously glandular, the lobes triangular, 

 about 0.5 mm. long, the apex pilosulous; corolla green or grayish-green, the tube 

 slender, 14 mm. long, gradually ampliate upwards, the lobes 3-3.5 mm. long, ob- 

 long, glabrous externally and internally; filaments 2 mm. long, appendaged at 

 their bases, the appendages slightly notched to parted to their bases or entire, very 

 inconspicuously papillate-puberulent, the corolla tube otherwise glabrous; style 

 11.5 mm. long; fruit about 6 mm. long; seed one, 4.5 mm. long. 



Cestrum guatemalense Francey, Candollea 6: 98. 1935. Huele 

 de noche; huele zorra (Chimaltenango). 



Wet thickets or moist forests, sometimes in Cupressus forests, 

 2,000-3,800 m.; Jalapa; El Progreso; Baja Verapaz; Guatemala; 

 Sacatepe"quez ; Chimaltenango; Totonicapan; Quezaltenango; Hue- 

 huetenango (type from mountains near Hacienda Chaucol, Nelson 

 3666 bis) ; San Marcos. Southern Mexico (Chiapas) . 



A shrub or small tree 1.5-8.5 m. tall, the branches pilosulous or glabrous, some- 

 times densely pubescent; leaves elliptic to elliptic-oblong or lance-elliptic, some- 

 times ovate, 6-12 (-17) cm. long, 2-4 (-6) cm. wide, glabrous above except on the 

 mid-vein, pilosulous below especially on the veins or in age almost glabrous, apex 

 acuminate or narrowly acute, acute or obtuse, sometimes rounded at the base; 

 petioles 1-2 (-3) cm. long; axillary leaves sometimes present; inflorescences axil- 

 lary and terminal, racemose or paniculate, sparsely to densely pubescent, the 

 bracts linear, caducous; pedicels 0.5-3 mm. long, glabrous; calyx 3.5-4 mm. long, 

 glabrous except for the ciliolate lobes, the lobes broadly deltoid or ovate and 

 short-mucronate, 0.2-0.5 mm. long; corolla pale yellow to greenish-yellow or 

 purplish-yellow, the tube 14-18 mm. long, glabrous externally, gradually ampliate 

 upwards, or sometimes abruptly so just below the mouth of the corolla tube, the 

 lobes 3-4 mm. long, ovate and short-mucronate, marginally pubescent externally; 

 filaments 1.5-3 (-3. 5) mm. long, glabrous or pilosulous at the base, appendages 

 absent, sometimes the base of the filaments swollen, the staminal traces pilo- 

 sulous or very rarely glabrous; style 13-17 mm. long; fruit about 12 mm. long, 

 white; seeds 7, 2.5-5 mm. long. 




28 FIELDIANA: BOTANY, VOLUME 24 



Cestrum guatemalense var. gracile Morton, J. Arnold Arbor. 

 17: 349. 1936. Hediondilla (San Marcos). The type is from Calel, 

 Totonicapan, Shannon (Donnell- Smith 282). Distinguished from 

 variety guatemalense by the pilosulous calyx. 



A common shrub in the western and central mountains of Gua- 

 temala. Cestrum guatemalense and C. pacayense are closely related 

 and may prove to be a single species. 



Cestrum lanatum Mart. & Gal. Bull. Acad. Brux. 12, pt. 2: 

 146. 1845. 



Moist or dry thickets, sometimes along rivers or in pastures, 

 800-2,200 m.; Chiquimula; El Progreso; Guatemala; Solola; Qui- 

 che"; Santa Rosa; Escuintla; Sacatepe"quez. Southern Mexico to 

 Costa Rica. Panama. 



A shrub or sometimes a small tree, 2-5 (-8. 5) m. tall, the young branches 

 densely covered with branched (with rays along their stalks) hairs; leaves lanceo- 

 late to elliptic or ovate, 4.5-20 cm. long, 2-6.5 cm. wide, with branched and stel- 

 late hairs above, soon glabrate, or sometimes the hairs persistent, often rugulose 

 above, densely pubescent below, the hairs branched and stellate, soft to the 

 touch or sometimes slightly asperous, apex acute or acuminate, rounded or ob- 

 tuse at the base; petioles 5-25 mm. long, densely covered with branched hairs; 

 axillary leaves sometimes present, lance-ovate to ovate, 10-20 mm. long, 5-10 mm. 

 wide, apex obtuse, short petiolate; inflorescences axillary and terminal, panicu- 

 late, usually dense and many-flowered, the rachis densely covered with branched 

 hairs, the bracts linear, densely covered with branched hairs, caducous, the flowers 

 sessile or subsessile; calyx 3.5-4 mm. long, densely covered with branched hairs, 

 the lobes about 0.5 mm. long; corolla pale yellowish-green or greenish, the tube 

 10-11 mm. long, glabrous externally (in ours), gradually ampliate above, the 

 lobes 2.5-3 mm. long, ovate to ovate-lanceolate, marginally pilosulous externally; 

 filaments 3-3.5 mm. long, slightly swollen and geniculate, gibbose or subappendicu- 

 late at their bases, pilosulous on about the lower one-third of the filament, the 

 staminal traces often pilosulous; style 10-11.5 mm. long; fruit about 7 mm. long; 

 seeds 2-5, 3-3.5 mm. long. 



Cestrum lanatum can be easily recognized in Central America by 

 the short corolla tube, geniculate filaments, and most of its parts 

 being densely covered with branched or branched and stellate hairs. 



It is closely related to C. mortonianum, but that species has a 

 longer corolla tube, appendiculate, and straight filaments. In Gua- 

 temala C. lanatum is readily distinguished from C. mortonianum by 

 its glabrous corolla tube. 



The crushed leaves give off a strong and disagreeable odor but the 

 flowers are fragrant. The leaves are sometimes placed in hen's nests 

 to drive off some of the parasites that infest the nests. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 29 



Called huele de noche and polo hediondo in El Salvador. 



Cestrum luteovirescens Francey, Candollea 6: 349. 1935. 



Moist thickets or mixed forests, 1,200-2,200 m.; Guatemala; 

 Sacatepe'quez (type from Capetillo, Donn.-Sm. 2679); Solola. El 

 Salvador. 



A slender shrub 1.5-3.5 m. tall, the branches glabrous; leaves narrowly 

 lanceolate to lanceolate-linear or rarely narrowly elliptic, 4-17.5 cm. long, 1-2.5 

 cm. wide, rarely to 3 cm. wide, glabrous above and below, apex narrowly acute 

 or acuminate, sometimes acute, obtuse or sometimes acute at the base; small 

 axillary leaves often present, lanceolate to ovate-lanceolate, 3-30 mm. long, short 

 petiolate or subsessile; inflorescences terminal, fasciculate, few-flowered, the bracts 

 linear, the flowers subsessile or short-pedicellate, subtended by leaves, the leaves 

 sparsely glandular, the pedicel 0.5 mm. long; calyx 3.5-4.5 mm. long, sparsely and 

 inconspicuously glandular, the lobes lanceolate, 0.8-1.5 mm. long, ciliolate; corolla 

 greenish or greenish-yellow, the tube 17-18.5 mm. long, glabrous externally, grad- 

 ually ampliate upwards, the corolla lobes 4-5 mm. long, ovate or lanceolate, mar- 

 ginally glandular-puberulent externally; filaments 2.5-3 mm. long, appendaged 

 at their bases, the appendages 0.8 mm. long, entire or slightly notched, pilosulous; 

 style 16.5-18 mm. long; fruit about 8 mm. long, white; seeds 4, 3.5-4.5 mm. long. 



Cestrum luteovirescens and C. skutchii are the only two species 

 in Guatemala with the larger leaves very long and narrow, almost 

 never exceeding 2.5 cm. in width. It is separated from C. skutchii 

 by flower color, longer calyx lobes, and glandular calyx. 



Cestrum megalophyllum Dunal in DC. Prodr. 13, pt. 1: 638. 

 1852. C. baenitzii Lingelsh. Fedde Repert. Sp. Nov. 7: 248. 1909. 



Moist or wet forests or mixed forests, 1,500 m. or less; Alta Vera- 

 paz; Izabal; Suchitepe"quez ; Quezaltenango; San Marcos. British 

 Honduras. Southern Mexico to Costa Rica. Panama. West In- 

 dies. Northern South America. 



A shrub 2-3 m. tall, sometimes a tree to 10 m. tall, the branches puberulent 

 or glabrous; leaves oblanceolate or elliptic, sometimes elliptic-oblong, 16.5-26 

 (-32) cm. long, (3-) 5-8 cm. wide, glabrous above and below, the apex acuminate 

 or acute, the base cuneate; petioles 1.5-2 cm. long; inflorescences axiliary, race- 

 mose, few-flowered, the axis equalling or shorter than the petioles, pubescent, 

 usually densely so, the bracts usually persistent, minute and densely pubescent, 

 the hairs brownish; pedicels 1 mm. long or less; calyx 2.5-3 mm. long, greenish, 

 glabrous or pilosulous, the lobes small, 0.5 mm. long or less, triangular, apex 

 acute, ciliolate and pilosulous apically; corolla greenish-white, the tube 11-12 

 mm. long, gradually ampliate above, glabrous externally, the lobes 3-3.5 mm. long, 

 lanceolate, acute, marginally puberulent externally; filaments 3.5-4 mm. long, 

 pilosulous at their bases, appendages absent; style 9-10.5 mm. long; fruit 6.5-9 

 mm. long, black or violaceous; seeds 3, 4.5-5.5 mm. long. 




FIG. 6. Cestrum mortonianum. A, branch, X Hi B, calyx with immature 

 fruit, X 3*4; C, dissected corolla, X 2%, with detail of ovary, X 5, and stamen, 

 X 6; D, corolla with detail of hairs, X 4. 



30 




GENTRY & STANDLEY: FLORA OF GUATEMALA 31 



In Guatemala, this species may be separated from all others on 

 the basis of the inflorescence and the filaments pilosulous at their 

 bases and the absence of appendages. 



Cestrum mortonianum J. L. Gentry, Phytologia 26:267. 1973. 



Moist forests or thickets, 1,050-2,500 m.; Chimaltenango; Que- 

 zaltenango; San Marcos (type from northwestern slopes of Volcan 

 Tajumulco, Steyermark 36509}. Endemic. 



A shrub or small tree, 2.5-5 m. tall, the branches with branched (with rays 

 along their stalks) and simple, puberulent hairs; leaves ovate to elliptic, 7-26 

 cm. long, 3-10 cm. wide, sparsely pubescent with branched and a few simple hairs 

 above, more densely pubescent below, the hairs branched and short stipitate- 

 stellate, apex acuminate or acute, acute or obtuse at the base; petioles 1-3.5 

 cm. long; inflorescences axillary, few- to many-flowered, the rachis with branched 

 hairs, a few simple hairs intermixed, the bracts linear, caducous, the flowers 

 sessile or subsessile; calyx 5-5.5 mm. long, sometimes cleft on one side, covered 

 with branched hairs, a few simple ones intermixed, the lobes 0.2-0.5 mm. long; 

 corolla pale green, the tube 16-16.5 mm. long, covered with branched hairs 

 externally, gradually ampliate above, the lobes 4.5-5 mm. long, ovate, pubescent 

 externally or sometimes only marginalljr pubescent; filaments 4 mm. long, appendi- 

 cmlate at their bases, the appendages entire, 0.3 mm. long, densely pilosulous, the 

 staminal traces densely pilosulous; style 15-15.5 mm. long; fruit immature. 



This species is closely related to Cestrum lanatum. It can be 

 readily distinguished from that species by the appendiculate and 

 relatively straight filaments. 



Cestrum nocturnum L. Sp. PI. 191. 1753. C. nocturnum var. 

 mexicanum 0. E. Schulz in Urban, Symb. Antill. 6: 256. 1909. Dama 

 de noche; huele de noche. 



Moist or wet thickets or forests, sometimes in open places, 1,800 

 m. or less; Pete"n; Alta Verapaz; Izabal; Jalapa; Guatemala; Saca- 

 tepe"quez; Chimaltenango; Quiche"; Huehuetenango; Escuintla; 

 Suchitepe"quez. British Honduras. Southern Mexico to Costa Rica. 

 Panama. West Indies. 



A shrub 1-4 m. tall, the young branches sparsely pilosulous or glabrous; 

 leaves ovate or lanceolate to elliptic or elliptic-oblong, mostly 8-16 cm. long, 

 2.5-6 cm. wide, glabrous above and below, apex acuminate, rounded to acute or 

 short attenuate at the base; petioles 1-1.5 cm. long, glabrous; inflorescences axil- 

 lary and terminal, racemose and paniculate, usually many-flowered, the rachis 

 pilosulous or sometimes glabrous, the bracts linear, caducous, pilosulous or some- 

 times glabrous, the flowers subsessile and short pedicellate; calyx 2-3 mm. long, 

 glabrous, shallowly cleft on two sides, the lobes triangular or mucronate, 0.2- 

 0.5 mm. long, ciliolate; corolla greenish-yellow or pale green, the tube slender, 17- 

 20 mm. long, gradually ampliate upwards, the corolla lobes 3-3.5 mm. long, 




32 FIELDIANA: BOTANY, VOLUME 24 



lance-oblong, marginally pilosulous externally; filaments 3-3.5 mm. long, append- 

 aged at their bases, the appendages mostly entire to slightly notched or sometimes 

 cleft to the base, glabrous or inconspicuously pilosulous, the staminal traces 

 glabrous or inconspicuously pilosulous; style 16.5-19.5 mm. long; fruit 7-9 mm. 

 long, white; seeds 5, 3.5-4 mm. long. 



This species has sweet-scented flowers. In Central America the 

 berries are said to be poisonous. 



Oestrum nocturnum is a common lowland species. It is fairly uni- 

 form throughout its wide range. 



Known in British Honduras by the following names: chacayum, 

 dama de noche, sapillo, and "night bloom." 



Oestrum pacayense Francey, Candollea 6: 90. 1935. C. keller- 

 manii Francey, I.e. 89 (type from Volcan de Pacaya, Kellerman 6453). 

 Hediondillo (San Marcos) ; huele de noche (Guatemala) . 



Moist or wet, usually dense forests, 1,800-3,300 m.; Guatemala 

 (type from Volcan de Pacaya, Kellerman 6404-)', Chimaltenango; 

 Solola; Quezaltenango; San Marcos. El Salvador. Honduras. 



A shrub 2-5 m. tall or sometimes a tree 12 m. tall, with a dense crown, the 

 branches glabrous; leaves narrowly elliptic to elliptic or lanceolate, 5-17.5 cm. 

 long, 2-4.5 cm. wide, glabrous above and below, apex acuminate or narrowly acute, 

 attenuate to obtuse at the base; petioles 1-2.5 cm. long, glabrous; axillary leaves 

 rarely present; inflorescences axillary and terminal, paniculate or racemose, 

 glabrous or sometimes sparsely pilosulous, few-to many-flowered, the bracts 

 linear; pedicels 1.5-3.5 mm. long; calyx 2.5-3 mm. long, glabrous, except for the 

 ciliolate lobes, the lobes 0.5 mm. long or less, broadly ovate and shortly mucronate 

 or only mucronate; corolla pale yellow or pale greenish-yellow, the tube slender, 

 15.5-18 mm. long, glabrous externally, abruptly ampliate just below the mouth 

 of the corolla tube, the lobes 2-3 mm. long, ovate and short-mucronate, papillate- 

 puberulent on the margins externally; filaments 2-4.5 mm. long, glabrous or 

 pilosulous at the base, appendages lacking, the staminal traces pilosulous; style 

 15.5-18 mm. long; fruit 8-10 mm. long, white; seeds 7, 4 mm. long. 



Probably not distinct from C. guatemalense. 



Cestrum racemosum R. & P. Fl. Peruv. 2: 29, t. 154. 1799. 

 Oestrum panamense Standl. J. Wash. Acad. Sci. 15: 460. 1925. C. 

 racemosum var. panamense Francey, Candollea 6 : 274. 1935. syn. nov. 

 Dama de noche; Ixcajaguay (Pete"n). 



Moist or wet forests, sometimes in cafetales as shade trees or in 

 dry quebradas, 35-1,000 m., rarely higher; Pete"n; Izabal; Quezal- 

 tenango; San Marcos. British Honduras. Southern Mexico. Hon- 

 duras; Nicaragua; Costa Rica. Panama. Colombia. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 33 



A shrub 1-10 m. tall or a tree sometimes 26 m. tall with a trunk 6.5 dm. in 

 diameter, the branches glabrous or very sparsely puberulent; leaves lanceolate 

 or lance-oblong, 7-22.5 cm. long, 2.5-7 cm. wide, glabrous above and below, the 

 lateral veins prominent and broadly curved to the leaf margin, the apex acuminate 

 to long acuminate, the base mostly rounded ; petioles 1-1.5 cm. long; inflorescences 

 axillary and terminal, racemose or paniculate, few- to many-flowered, the rachis 

 pilosulous, the bracts linear, pilosulous, persistent or caducous; pedicels 0.5 

 mm. long, pilosulous, or the flowers sessile; calyx 1.5-2 mm. long, pilosulous, the 

 lobes triangular, 0.2-0.5 mm. long, ciliolate; corolla greenish-yellow or pale green, 

 the tube slender, 11-12 mm. long, glabrous externally, very gradually ampliate 

 until just below the mouth of the corolla tube, the corolla lobes 2.5-3 mm. long, 

 oblong-lanceolate, marginally puberulent externally; filaments 1-2 mm. long, ap- 

 pendages absent, the staminal traces inconspicuously pilosulous; style 11-12 mm. 

 long; fruit 5-7.5 mm. long; seeds 4-8, 2.5-4 mm. long. 



Called "night bloom" in British Honduras. 



The treatment of Oestrum panamense as a variety of C. racemo- 

 sum may be correct, but its elevation to specific rank does not appear 

 to be justified. 



Cestrum regelii Planchon in Fl. Serres, 9: 229, t. 91>6. 1854. C. 

 chiapense Brandegee, Univ. Calif. Publ. Bot. 6: 192. 1915. C. jacal- 

 tenanginum Loes. Verh. Bot. Ver. Brandenb. 65: 98. 1923 (type from 

 Jacaltenango, Huehuetenango, C. & E. Seler 261+1}. 



Moist forests, 1,465-1,800 m., rarely higher; Guatemala; Saca- 

 tepe"quez; Huehuetenango; San Marcos. Southern Mexico (Chia- 

 pas). 



A shrub to 3.5 m. tall, the young branches mostly densely covered with simple 

 multicellular hairs; leaves ovate to ovate-elliptic or lanceolate, 4-12 cm. long, 

 1.5-4.5 cm. wide, pilosulous above and below, more densely so on the veins be- 

 low, apex acute or acuminate, acute or rounded at the base; petioles 1-2 cm. 

 long; inflorescences axillary and terminal, fasciculate, and few-flowered or only 

 with a few solitary flowers, subtended by leaves, the bracts foliaceous, lanceolate 

 or linear; calyx 5-10 mm. long, densely covered with multicellular hairs, the 

 lobes lanceolate to oblong-lanceolate, or linear, 2-5 mm. long, narrowly acute, 

 pilosulous internally; corolla bright orange, the tube 15.5-20 mm. long, externally 

 glabrous, ampliate upwards, ventricose just below the mouth of the corolla tube, 

 the corolla lobes 1-1.5 mm. long, ovate and mucronate, marginally puberulent 

 externally; filaments 9.5-11 mm. long, swollen, canaliculate and inconspicuously 

 pilosulous at their bases, appendages absent; style 16-18 mm. long; fruit black or 

 violaceous. 



Oestrum scandens Vahl, Ecolg. Amer. 1: 24. 1796. C. perilam- 

 banon Loes. Verh. Bot. Ver. Brandenb. 65: 98. 1923 (type from Chi- 

 quimula, C. & E. Seler 3381}. Chunumo coyote (Chiquimula). 



Wet or dry thickets or lowland forests, just above sea level to 

 900 m.; Chiquimula; Jutiapa; Santa Rosa; Escuintla; Retalhuleu. 




34 FIELDIANA: BOTANY, VOLUME 24 



Southern Mexico. El Salvador to Costa Rica. Colombia and Vene- 

 zuela. 



Scandent shrubs, 2.5 m. long or less, the branches puberulent or glabrous; 

 leaves ovate to ovate-lanceolate, 7-16.5 cm. long, 3.5-6 cm. wide, glabrous above 

 and below, apex acuminate or acute, rounded or subcordate at the base; petioles 

 1-1.5 cm. long; inflorescences lax, axillary and terminal, paniculate or sometimes 

 racemose, the axis 2.5-15 mm. long, sparsely pilosulous, the bracts small, ciliolate, 

 caducous; pedicels 0.5-1.5 mm. long; calyx 3.5-4 mm. long, green, the lobes about 

 1 mm. long, triangular, acute, ciliolate; corolla pale green or green, the tube 18- 

 20 mm. long, slender, ampliate at the throat of the corolla tube, glabrous, the lobes 

 7-9.5 mm. long, narrowly oblong-lanceolate, acute, ciliolate; filaments 0.5 mm. long 

 or less, glabrous, appendages absent; style 17-19 mm. long; fruit 7-10 mm. long, 

 blue or white; seeds 5-8, 5 mm. long. 



Cestrum scandens is easily recognized by its scandent habit, very 

 long and narrow corolla lobes, and the stamens inserted just below 

 the mouth of the corolla tube. 



Cestrum skutchii Morton, J. Arnold Arbor. 17: 347. 1936. 

 Aguacatillo de montana (Quezaltenango) . 



Second growth thickets, 1,500-1,710 m.; Quezaltenango (type 

 from Volcan de Zunil, Skutch 988). Endemic. 



A shrub 1.5-3.5 m. tall, glabrous throughout; leaves lanceolate-linear, 14.5- 

 24 cm. long, 1.5-2 (-2. 5) cm. wide, glabrous above and below, apex very narrowly 

 acute, acute or very shortly attenuate at the base; petioles 8-10 mm. long; inflo- 

 rescences axillary and pseudo-terminal, fasciculate, and 3-9 flowered, or a few 

 flowers solitary, axis dark purple, the bracts linear, dark purple, glabrous; pedicels 

 4-5 mm. long, dark purple, cleft about one-third the length of the calyx on two 

 sides, the lobes short-mucronate, 0.5 mm. or less long, unequal, ciliolate; corolla 

 white, more or less suffused with purple, the tube 19-20 mm. long, gradually am- 

 pliate above, the lobes 5-6 mm. long, broadly ovate, acute, inconspicuously papil- 

 late marginally; filaments 3.5 mm. long, appendaged at their bases, the appendages 

 entire, about 0.3 mm. long, pilosulous; style 18-19 mm. long; fruit pale green, 8 

 mm. long. 



Cestrum skutchii is similar to C. luteovirescens and it can be sep- 

 arated from that species by the characters stated in the key. 



Cestrum sp. 



Wet forests, 1,500 m. Known only from the following collection, 

 Huehuetenango, Sierra de los Cuchumatanes, 2 miles southeast of 

 Maxbal, about 17 miles north of Barillas, July 15-16, Steyermark 

 48708. 



A vine, the branches glabrous; leaves lance-elliptic, 5-11 cm. long, 1.5-3 cm. 

 wide, glabrous above and below, apex long acuminate, obtuse at the base; petioles 

 3-7 mm. long, glabrous; inflorescences axillary, consisting of a single, subsessile 

 flower on a long peduncle, the peduncles 1.5-3.5 cm. long, glabrous, one of the 




GENTRY & STANDLEY: FLORA OF GUATEMALA 35 



flowers sessile in the leaf axil; fruiting calyx 3-3.5 mm. long, glabrous, the lobes 

 0.5 mm. long or less, broadly deltoid; fruit 6.5 mm. long; seeds 7, 2-3.5 mm. long. 



This specimen seems to represent an undescribed species, but we 

 hesitate to describe it on inadequate material. The inflorescence is 

 unique among the species known in Guatemala. 



Morton (1936) suggested that C. psittacinum Stapf should prob- 

 ably be included in the Oestrums of Guatemala. The species was 

 described from cultivated material of unknown origin. We have 

 not included it in our treatment because of its unknown origin and 

 the lack of specimens in herbaria. 



CYPHOMANDRA Sendtner 



Shrubs or small trees, unarmed; leaves alternate, entire, 3-lobed, or pinnatifid, 

 often large, sometimes dimorphic, long petiolate; inflorescences cymose, mostly 

 beside the leaf axil, the rachis nodose, the flowers pedicellate; calyx campanulate, 

 5-lobed; corolla subrotate, pink to purplish, or greenish-purple, the limb deeply 

 5-parted, the lobes valvate in bud; stamens 5, exserted; filaments short and broad; 

 anthers oblong or attenuate, dehiscent by two apical pores or slits, these often 

 continued as longitudinal introrse slits part way down the thecae, the connective 

 prominent, adaxially thickened; ovary bilocular, the ovules numerous, the disk 

 annular, entire or dentate; style thick or elongate and filiform; stigma scarcely or 

 strongly dilated; fruit a berry, often large, ovoid, oblong, or elliptic; seeds numer- 

 ous, compressed; embryo strongly curved or subspiral. 



About 60 species all in tropical America. Two or three other 

 species are found in southern Central America. 



Ovary covered with short, glandular and eglandular hairs, the pubescence per- 

 sisting in fruit; stigma strongly dilated; anther connective 1.5-2 mm. wide 

 at the base C. rojasiana. 



Ovary and fruit glabrous; stigma scarcely dilated; anther connective 0.8-1.5 mm. 

 wide C. betacea. 



Cyphomandra betacea (Cav.) Sendtn. Flora 28: 172. 1845. 

 Solatium betaceum Cav. Icon. 6: 15, t. 52^- 1801. Granadilla; pix, 

 caxlan pix (Quecchi); tomate; tomate de arbol; tomate extranjero; 

 tomate granadilla. 



A native of South America that is cultivated for its edible fruit 

 in many parts of tropical America. Planted occasionally in the 

 mountains of Guatemala at middle elevations. The fruit is often 

 seen in the markets of Huehuetenango, Quezaltenango, Guatemala, 

 and Coban. It is somewhat more acid than the common tomato, 

 Lycopersicon, and is used in much the same manner, principally for 

 salads and dukes. 




D 



FIG. 7. Cyphomandra rojasiana. A, habit, X %; B, corolla, X 1}^; C, style, 

 X 2%; D, stamen, abaxial view, X 2J^; E, stamen, lateral view, X 2>; F, calyx, 

 X 3; G, mature fruit, X K. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 37 



Called tomate de polo in Honduras. 



The correct name for this species may be C. crassifolia (Ortega) 

 Kuntze. 



Cyphomandra rojasiana Standl. & Steyerm. Field Mus. Bot. 

 22: 377. 1940. 



In thickets or second growth, 1,100-1,500 m.; Quezaltenango; 

 Suchitepe"quez (type from Finca Moca, Skutch 1472). Mexico 

 (Chiapas) . 



A shrub 2-3.5 m. tall, sometimes straggling, the branches covered with short, 

 glandular and eglandular hairs, glabrate in age; leaves membranceous, entire, 

 broadly ovate, 12-30(-45) cm. long, 9-18(-27) cm. wide, the hairs short, glandular 

 and eglandular above and below, apex acuminate, rounded, and cordate at the 

 base; inflorescences mostly axillary or sometimes extra-axillary, cymose, few- 

 flowered, the rachis nodose; peduncles 2-3.5 cm. long, accrescent in age, the hairs 

 short, glandular and eglandular; pedicels 8-15 mm. long, in fruit to 4.5 cm. long, 

 the hairs short, glandular, and eglandular; calyx campanulate, 2-3 mm. long, the 

 hairs short, glandular and eglandular, the lobes ovate, 1.5-2 mm. long, apex mu- 

 cronate, densely pubescent; corolla limb parted nearly to the base, the lobes ob- 

 long or lanceolate, 8-12 mm. long, acute, externally green, internally purple, 

 shining and bordered with green, ciliate, otherwise glabrous; filaments 1.5 mm. 

 long; anthers 4.5-6 mm. long, the connective gibbose, projecting below the base 

 of the anther sacs, 1.5-2 mm. wide at the base; ovary covered with short, glandu- 

 lar and eglandular hairs; style thick, 3.5-5 mm. long; stigma strongly dilated; 

 fruit elliptic or ovate-elliptic, 5 cm. long, 3-3.5 cm. wide, pubescence of glandular 

 and eglandular hairs, light yellow with rich green stripes and splotches at both 

 ends; seeds 3.5-5 mm. long. 



This species is closely related to the common tree tomato, Cy- 

 phomandra betacea, and may be only a variety of that species. 



The stems and leaves give off a disagreeable odor when crushed, 

 fide Skutch. 



Called huevo de gato in Chiapas. 



DATURA L. 



References: W. E. Safford, Synopsis of the genus Datura. J. 

 Wash. Acad. Sci. 11: 173-189. 1921. A. G. Avery et al., Blakeslee: 

 The Genus Datura. The Ronald Press. New York. 289 pp. 1959. 



Herbs, shrubs, or small trees, glabrous or pubescent; leaves alternate, entire 

 to sinuate, or shallowly lobed, membranaceous, petiolate; inflorescences consist- 

 ing of a solitary flower, the flowers pedicellate and in the forks of the branching 

 stem, small or very large and showy, erect or pendulous; calyx long tubular, 5- 

 lobed or spathe-like and cleft down one side, sometimes circumscissle near the 




38 FIELDIANA: BOTANY, VOLUME 24 



base and leaving a persistent flaring collar under the fruit; corolla funnelform or 

 narrow funnelform, white, yellow, pink, or red, the limb 5-lobed or 10-toothed, the 

 lobes acuminate to long caudate, plicate in bud, the tube long and slender; sta- 

 mens 5, inserted near the middle of the corolla tube, included (in ours) or slightly 

 exserted; filaments slender, pubescent or glabrous below; anthers linear, free or 

 rarely coherent, longitudinally dehiscent; ovary bilocular (sometimes falsely tetra- 

 locular), the ovules numerous; style filiform, included; stigma bilobed; fruit a 

 capsule, 4-valvate or breaking open irregularly, armed with spines or unarmed; 

 seeds numerous, laterally compressed or angulate; embryo curved. 



About 25 species, mainly in warm-temperate or tropical regions 

 of the world. Only one other species, Datura metel L., a native of 

 India, occurs spontaneously in Centra America, others, e. g., D. 

 arborea, probably are cultivated occasionally. Apparently all spe- 

 cies of Datura have alkaloids with narcotic properties. Several 

 species have been used in religious and ceremonial rituals by the 

 native people of the New World. Mr. T. E. Lockwood is currently 

 working on a revision of section Brugmansia (tree Daturas). He is 

 recognizing this section at the generic rank. 



Shrubs or small trees; flowers pendulous, 22-30 cm. long; calyx completely decidu- 

 ous or persistent, not circumscissle and forming a flaring collar under the 

 fruit; fruit unarmed. 

 Calyx spathe-like, terminating in an acuminate point; anthers free.Z). Candida. 



Calyx tubular, lobed at the apex; anthers coherent D. suaveolens. 



Herbs; flowers erect, 4.5-18 cm. long; calyx circumscissle near the base and leaving 



a persistent flaring collar under the fruit; fruit armed with spines. 

 Fruiting pedicels erect; fruits dehiscing regularly, 4-valvate; leaves sparsely 



pubescent or glabrate below D. stramonium. 



Fruiting pedicels nodding; fruits breaking open irregularly; leaves commonly 

 densely pubescent below. 



Corolla 4.5-6 cm. long D. pruinosa. 



Corolla 14-18 cm. long D. inoxia. 



Datura Candida (Pers.) Safford, J. Wash. Acad. Sci. 11: 182. 

 1921. D. arborea R. &. P. Fl. Peruv. 2: 15, t. 127. 1799, non D. ar- 

 borea L. 1753. Brugmansia Candida Pers. Syn. PL 1: 216. 1805. 

 Campana; florifundio; floripondio; kampani (Quecchi); krevapunta; 

 trompetero. 



Moist or wet thickets, 360-2,500 m.; Alta Verapaz; Quiche"; 

 Quezaltenango; Santa Rosa; Suchitepe"quez. Perhaps a native of 

 Peru. It is cultivated in many parts of Guatemala and naturalized 

 in some areas. Widely planted in tropical regions as an ornamental 

 for its large and showy white flowers. 



A shrub or small tree, 2-5 m. tall, the branches pilosulous or puberulent; 

 leaves entire or sometimes repand, broadly ovate to ovate-oblong or ovate- 

 elliptic, mostly 15-30 cm. long, 6-12 cm. wide, pilosulous above and below, apex 




FIG. 8. Datura Candida. A, habit, X Y^', B, portion of corolla showing sta- 

 mens, X 1 A; G, pistil, X 1 A. 



39 




40 FIELDIANA: BOTANY, VOLUME 24 



narrowly acute or acuminate, base rounded or sometimes acute, often unequal; 

 petioles mostly 4.5-12 cm. long, pilosulous; flowers pendulous, very fragrant; 

 pedicels 3.5-5 cm. long, pilosulous; calyx 10-15 cm. long, spathe-like, cleft down 

 one side but otherwise entire, persisting around the mature fruit, pilosulous or 

 glabrate, acuminate; corolla white, the tube 22-30 cm. long, margin of the limb 

 between the lobes entire, the lobes spreading, long caudate, 5.5-7 cm. long; sta- 

 mens included, inserted just below the middle of the tube; filaments 4-4.5 cm. long, 

 villous below, the staminal traces villosulous; anthers free, 2.5-3 cm. long; style 

 included 17-19 cm. long, the stigma oblong, 5-7 mm. long; fruit unarmed, fusiform, 

 20 cm. long, 2 cm. diameter at the middle, seldom maturing; seeds 6-10 mm. long, 

 angulate, rugose. 



Datura Candida is often confused with D. arborea, a name fre- 

 quently applied to any white flowered tree-datura. 



Flowers sometimes are placed beside the pillows of persons trou- 

 bled with insomnia because of the popular belief that the fragrance 

 of the flowers induces sleep. 



Called floripondio in El Salvador and Honduras. 



Datura inoxia Miller, Gard. Diet. ed. 8, Datura no. 5. 1768. D. 

 meteloides DC. in Dunal, Prodr. 13(1): 544. 1852. Tapa (Jutiapa); 

 tapate (Zacapa). 



Damp thickets or waste places, 200-850 m.; Zacapa; El Progreso; 

 Chiquimula. United States, mostly the west and southwest. Mex- 

 ico. Honduras. West Indies. South America. 



Herb 1 m. tall or less, the stems densely pilosulous; leaves subentire to re- 

 pand or sinuate, sometimes shallowly lobed, lanceolate to broadly ovate, mostly 

 10-22 cm. long, 4.5-10.5 cm. wide, pilosulous above, more densely so below, 

 especially on the veins, apex acuminate or narrowly acute, base unequal, short 

 attenuate, acute or sometimes rounded; petioles 3.5-8.5 cm. long, densely pilo- 

 sulous; flowers erect, fragrant; pedicels 1-2 cm. long, densely pilosulous; calyx 10- 

 12 cm. long, pilosulous, circumscissle just above the base of the tube and falling 

 off with the corolla, the base persistent and accrescent, forming a flaring collar at 

 the base of the fruit, the lobes 1.5-2.5 cm. long, narrowly long acuminate; corolla 

 white, 10-lobed, the tube 14-18 cm. long, the lobes spreading, caudate, 8-10 mm. 

 long; stamens included, inserted above the middle of the corolla tube; filaments 

 4.5-5.5 cm. long, glabrous, the staminal traces glabrous; anthers free, 1-1.4 cm. 

 long; style included, 13.5-16.5 cm. long, fruit nodding, globose, 2.5-3 cm. in diame- 

 ter, breaking open irregularly, armed with numerous, long and slender, pubescent 

 spines; seeds compressed, 4-4.5 mm. long. 



Datura inoxia was described from a cultivated plant grown in 

 England. The seeds were sent from Mexico. It is frequently cul- 

 tivated as an ornamental in gardens, now widely distributed in such 

 places as the Canary Islands, the Near East, India, and China. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 41 



Datura inoxia has been divided into at least three subspecific 

 taxa by A. S. Barclay, in herb. We have only the typical form of the 

 species in Guatemala. It has been frequently confused with D. 

 metel L., a native of Asia, now introduced into the Americas, from 

 which it can be distinguished by its usually dense pubescence of the 

 herbage, 10-lobed corolla limb and the fruits armed with long and 

 slender spines. Datura metel has a 5-lobed corolla limb and the fruits 

 armed with tubercles or short, broad spines. It often has a double 

 corolla. 



Datura pruinosa Greenm. Proc. Am. Acad. Sci. 33: 486. 1898. 

 Known only from the following collection in Guatemala; Zacapa, 

 180 m., Kellerman 7017. Mexico. 



Herb, the stems pilosulous; leaves repand to shallowly lobed, ovate to ovate- 

 oblong, 6-14.5 cm. long, 3.5-8.5 cm. wide, pilosulous above and below, apex acute 

 or acuminate, base unequal or sometimes equal, acute to rounded; petioles 1.5- 

 4.5 cm. long, pilosulous; flowers erect; pedicels 0.4-0.8 mm. long, densely pilo- 

 sulous; calyx 3-4.5 cm. long, pilosulous, circumscissle just above the base of the 

 tube and falling off with the corolla, the base persistent and accrescent, forming a 

 flaring collar at the base of the fruit, the calyx lobes 1 cm. long, narrowly acute; 

 corolla white, 5-lobed, the tube 4.5-6 cm. long, the lobes spreading, caudate, 6 mm. 

 long; stamens included, inserted at about the middle of the corolla tube; filaments 

 1.2-1.6 cm. long, glabrous; anthers free, 2. 5-3. 5 mm. long; style included, 3-3.5 cm. 

 long; fruit nodding, globose, 2-4 cm. in diameter, breaking open irregularly, armed 

 with numerous, long and slender, pubescent spines; seeds compressed, 3-4 mm. 

 long. 



The Kellerman collection cited above was annotated by A. S. 

 Barclay as Datura leichhardtii F. v. Muell. in Benth. using the name 

 pruinosa at the subspecific rank. Datura leichhardtii is based upon 

 plants collected in Australia. We are maintaining the use of the 

 name D. pruinosa for these Mexican and Guatemalan plants until 

 the type can be studied. 



Datura stramonium L. Sp. PI. 1: 179. 1753. Machul (Huehue- 

 tenango); tapa (Chimaltenango, Jutiapa); vuelvete loca (Chimal- 

 tenango, Jutiapa, Solola). 



In old fields, waste ground, cultivated fields or thickets, 850- 

 2,500 m.; Jalapa; Chimaltenango; Solola; Quezaltenango; Huehue- 

 tenango; Jutiapa; Sacatepe"quez. British Honduras. United States. 

 Mexico. El Salvador to Costa Rica. West Indies. South America. 

 Old World. Widely distributed in temperate and warm temperate 

 regions of the world. 




42 FIELDIANA: BOTANY, VOLUME 24 



A weedy herb 1 m. or less tall, the young stems pilosulous, glabrate in age; 

 leaves sinuate to shallowly lobed, the lobes acute, ovate to ovate-oblong or el- 

 liptic, 8-22 cm. long, 3.5-14 cm. wide, pilosulous or glabrous above and below, 

 apex narrowly acute, base unequal or equal, short attenuate, acute or sometimes 

 rounded; petioles 1.5-5.5 cm. long, glabrous or pilosulous; flowers erect; pedicels 

 0.5-1 cm. long, pilosulous; calyx 2.5-3.5 cm. long, pilosulous or glabrate, circum- 

 scissle just above the base of the tube and falling off with the corolla, the base 

 persistent and accrescent, forming a flaring collar at the base of the fruit, the calyx 

 lobes 3-5 mm. long, acuminate or long acuminate; corolla white or violet, 5-lobed, 

 the tube 6-9 cm. long, the lobes spreading, caudate, 8-10 mm. long; stamens in- 

 cluded, inserted below the middle of the corolla tube; filaments 2.2-2.5 cm. long, 

 sparsely villosulous at the base; anthers 3.5-5 mm. long; style included, 4-6 cm. 

 long; fruit erect, ovoid, 2.5-3.5 cm. in diameter, dehiscing regularly, 4-valvate, 

 armed (in ours), the spines numerous, long and slender or sometimes unarmed; 

 seeds compressed, 3-3.5 mm. long. 



Datura stramonium has been separated into at least three sub- 

 specific taxa by A. S. Barclay, in herb. 



It is reported that cigarettes made from the dried leaves are 

 sometimes smoked in Guatemala to relieve asthma. 



Called thorn-apple or Jimson (a corruption of Jamestown) weed 

 in the United States. Called tapa or tapate in Central America. 



Datura suaveolens Humb. & Bonpl. ex Willd. Enum. Hort. 

 Berol. 227. 1809. 



A native of Brazil now commonly cultivated in tropical regions 

 and becoming naturalized in some areas. Only one collection is 

 known from Guatemala. Quezaltenango, along Quebrada San Geron- 

 imo, 1,300-2,000 m., Steyermark 33456. 



A shrub or small tree, 1-6.5 m. tall, the young branches pilosulous or puberu- 

 lent; leaves entire or sometimes repand, ovate to elliptic, mostly 15-36 cm. long, 

 4.5-14 cm. wide, pilosulous above and below, glabrate in age above, apex acumi- 

 nate or acute, the base obtuse or acute, sometimes short attenuate, often unequal; 

 petioles mostly 2-10 cm. long, pilosulous; flowers very fragrant and pendulous; 

 pedicels 2.5-5.5 cm. long, puberulent or glabrate; calyx 5-lobed, 9-12 cm. long, 

 tubular, glabrous, deciduous, the lobes 2-3.5 cm. long, short acuminate; corolla 

 white, the tube 22-28 cm. long, the lobes spreading, long caudate, 3.5-4.5 cm. long; 

 stamens included, inserted just below the middle of the tube; filaments 3.5-4.5 cm. 

 long, villous below, the staminal traces villosulous; anthers coherent, 3.5-4 cm. long; 

 style included, 18-22 cm. long, the stigma oblong, 5-5.5 mm. long; fruit unarmed, 

 fusiform, 20 cm. long, 2.5 cm. in diameter at the middle; seeds 8-12 mm. long, an- 

 gulate and rugose. 



JALTOMATA Schlechtendal 



Herbs; leaves alternate, often in pairs, entire or sinuate-dentate, petiolate; 

 inflorescences axillary and umbellate, with a long peduncle, the flowers pedicellate; 




GENTRY & STANDLEY: FLORA OF GUATEMALA 43 



calyx broadly campanulate, 5-lobed, accrescent in fruit and spreading beneath the 

 berry or spreading campanulate; corolla rotate, greenish-white or pale green, 

 5-lobed, the lobes valvate in bud; stamens 5; filaments filiform, dilated and some- 

 what swollen at their bases, pubescent or sometimes glabrous; anthers longi- 

 tudinally dehiscent; ovary bilocular, the ovules numerous, the disc annular; style 

 slightly dilated above; stigma capitate, emarginate; fruit a berry; seeds numerous, 

 laterally compressed; embryo strongly curved. 



About 15 species ranging from the southwestern United States 

 to South America and the West Indies. Only the following two 

 species are known from Central America. 



Jaltomata Schlecht. is the correct name that applies to the plants 

 traditionally referred to as Saracha R. & P. See comments by J. L. 

 Gentry, Phytologia 27: 286-288. 1973, for an explanation of the 

 use of the name Jaltomata Schlecht. 



Plants small, the stems prostrate or decumbent, 2 dm. long or less; pedicels 2, 

 rarely 3; leaves small, 1.5-3.5 cm. long J. confinis. 



Plants large, the stems erect or decumbent, 6-9 dm. long; pedicels mostly 3-5; 

 leaves large, 5.5-18 cm. long J. procumlcns. 



Jaltomata confinis (Morton) J. L. Gentry, Phytologia 27: 286. 

 1973. Saracha confinis Morton, Proc. Biol. Soc. Wash. 51: 77. 1938. 



Open places on roadside banks or grassy hillsides, 2,400-3,800 m.; 

 Chimaltenango (type from Santa Elena, Skutch 447} ; Huehuetenango. 

 Endemic. 



Prostrate or decumbent herbs, usually pilosulous throughout, the stems 

 several, usually branching, 1.2-2.2 dm. long; leaves ovate to ovate-rhomboid, 

 1.5-3.5 cm. long, 2 cm. wide or less, acute, the base attenuate; petiole 1-10 mm. 

 long; inflorescences umbellate with 2 or rarely 3 flowers; peduncles 2-4 cm. long; 

 pedicels 8-20 mm. long; calyx 4-5 mm. long, accrescent in fruit and spreading 

 campanulate, about 1 cm. wide, the lobes ovate, acute; corolla greenish-white, the 

 limb about 1 cm. wide; filaments 2.5-3 mm. long, densely hairy below; anthers 

 about 1.5 mm. long; style about 2 mm. long; berry (immature) about 9 mm. 

 in diameter. 



Jaltomata procumbens (Cav.) J. L. Gentry, Phytologia 27: 

 287. 1973. Saracha procumbens (Cav.) R. & P. Fl. Peruv. 2: 43. 1799. 

 Atropa procumbens Cav. Icon. PI. 1 : 53, t. 72. 1791. Saracha Jaltomata 

 Schlecht. Hort. Hal. 10. 1839. Cinco negros', hierab ixbolul; madre 

 jilete; belladona cimarrona. 



Moist or wet thickets or forests, often a weed in cultivated fields, 

 800-2,500 m.; Alta Verapaz; Izabal; Guatemala; Chimaltenango; 

 Solola; Quezaltnenago; Huehuetenango; Escuintla; Suchitepe"quez ; 

 San Marcos. Southwestern United States. Mexico to Costa Rica. 

 Panama. South America. 




FIG. 9. Jaltomata procumbens. A, habit, X K; B, fruit and calyx, X 2 and 

 X 3; C, corolla dissected, X 3. 



44 




GENTRY & STANDLEY: FLORA OF GUATEMALA 45 



Erect or decumbent herbs, usually branched above, essentially glabrous 

 throughout or pilose, the stems thick and somewhat succulent, 6-9 dm. tall or 

 sometimes taller; leaves ovate to elliptic, 5. 5-18 (-35) cm. long, 3-8 (-19) cm. wide, 

 acuminate, attenuate at the base; petiole 0.5-5 cm. long, sometimes slightly 

 winged; inflorescences umbellate, with 3-5(-15) or rarely 2 flowers; peduncles 2-4 

 cm. long; pedicels 1-2 cm. long; calyx 4-5 mm. long, accrescent in fruit, and widely 

 spreading beneath the berry, 2-2.5 cm. wide, the lobes ovate, acute; corolla 

 greenish-white or pale green, the limb 1-2.5 cm. wide; filaments 3.5-4.5 mm. long, 

 densely pubescent or sometimes glabrous; anthers 2-2.5 mm. long; style about 

 5 mm. long; berry purple or black, 10-12 mm. in diameter, subglobose, very juicy; 

 seeds 1.5-2 mm. long, laterally compressed, reticulate-foveate. 



The young leaves are used as a pot herb by the Indians. 



JUANULLOA Ruiz & Pavon 



Shrubs, mostly epiphytic; leaves alternate or in pairs, entire, petiolate; in- 

 florescences cymose, lateral or pseudoterminal, the flowers large and pedicellate; 

 calyx campanulate or tubular, 5-lobed; corolla tubular, reddish to orange (in 

 ours), or whitish, yellow to brown, the limb 5-lobed, the lobes small and broad, 

 imbricate in bud, the tube slightly ampliate above the middle and contracted at 

 the apex; stamens 5, mostly included; filaments inserted near the base of the 

 corolla tube, densely hairy at the point of insertion; anthers longitudinally dehis- 

 cent, oblong or linear, apiculate; ovary bilocular, the ovules numerous, the disk 

 thick and annular; style filiform; stigma bilobed, slightly dilated; fruit a berry, 

 sometimes almost dry, indehiscent; seeds large, numerous and compressed; 

 embryo slightly or strongly curved. 



Species about 10, in the tropical rain forests from southern Mex- 

 ico and Central America and in South America along the Andean 

 countries to Bolivia. Only the following species is known in Mexico 

 and Central America. 



Juanulloa mexicana (Schlecht.) Miers, Ann. Mag. Nat. Hist. 

 4: 188. 1849. Laureria mexicana Schlecht. Linnaea 8: 513. 1833. J. 

 sargii Donn.-Sm. Bot. Gaz. 18: 5,t.l. 1893 (type from near Escuintla, 

 330 m., Donnell-Smith 1467). 



Nearly always an epiphyte on trees, Pacific slope, 300-1,000 m.; 

 Escuintla; Quezaltenango. Southern Mexico. Honduras; Costa Rica. 

 Panama. 



An epiphytic shrub, rarely terrestrial, the branches thick and spreading, with 

 corky bark, 4 m. long or less, the young branches usually densely pubescent with 

 stellate and branched (rays along their stalks) hairs; leaves mostly crowded near 

 the ends of the branches, sometimes in pairs, membranaceous or subcoriaceous, 

 obovate or elliptic, 10-20 cm. long, 3. 5-10 (-13) cm. wide, obtuse or acute, cuneate 

 at the base, sparsely pubescent with branched hairs or glabrate above, usually 

 more densely pubescent with branched hairs or somewhat glabrate below, the 




46 



FIELDIANA: BOTANY, VOLUME 24 



FIG. 10. Juanulloa mexicana. Plate I from the Botanical Gazette for the year 

 1893, used by permission of the University of Chicago. Magnifications may be 

 calculated from the description. 



petiole 0.5-15 mm. long; inflorescence cymose, pseudoterminal or lateral; pedicels 

 thick, 1-2 cm. long, densely covered with ochraceous, very short simple and 

 branched hairs; calyx narrowly campanulate, 2.5-3 cm. long, costate, orange, 

 densely covered with ochraceous, very short simple and minute branched hairs, 

 red within, cleft for half its length or more, the lobes oblong-lanceolate or lanceo- 

 late, equal or unequal, acute; corolla red or orange, 4-5 cm. long, densely covered 

 with yellowish very short simple and minute branched hairs, the lobes 2-2.5 mm. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 47 



long, obtuse; stamens included or rarely slightly exserted; filaments 2-2.4 cm. long, 

 inserted 4.5-5 mm. from the base of the tube, densely hairy at the point of inser- 

 tion; anthers 11.5-12.5 mm. long, apiculate; style included, about 4 cm. long; 

 berry globose-ovoid, about 2 cm. long, surpassed by the calyx; seeds 4-4.5 mm. 

 long, oblong, reticulate-foveate. 



' i-a 



LYCIANTHES (Dunal) Hassler 



Reference: G. Bitter, Die Gattung Lycianthes, Abh. Naturwiss. 

 Vereine Bremen 24: 292-520. 1920. 



Shrubs, vines, or herbs, unarmed; glabrous or pubescent, the hairs simple or 

 often variously stellate or branched; leaves simple, alternate, entire, the upper 

 leaves often in pairs; inflorescences axillary, the flowers mostly pedicellate; calyx 

 campanulate, truncate, often bearing 5 or 10 linear-subulate to knob-like append- 

 ages at or just below the margin of the calyx; corolla subrotate, white to purple, 

 or pale yellow, the lobes plicate or sometimes valvate in bud; stamens 5, sometimes 

 unequal in length; filaments short and equal or sometimes 1 or 3 filaments longer 

 than the others; anthers ellipsoid to oblong, dehiscent by small apical introrse 

 pores; ovary bilocular, the disk inconspicuous; stigma capitate, entire or obscurely 

 bilobed; ovules numerous (in ours); fruit a berry, usually globose and fleshy, 

 sometimes ellipsoid; seeds numerous (in ours), compressed; embryo strongly curved 

 or spiral. 



A genus of about 140 species, in the tropics of both hemispheres. 

 Other species are known from southern Central America. Lycianthes 

 was established principally by the work of Bitter, who published a 

 monograph in 1920. While the segregate genus has been accepted 

 by some recent authors, it has been criticized by others, notably by 

 the late C. V. Morton. For a discussion of the subject (unfavorable 

 to maintenance of Lycianthes) see C. V. Morton, Contr. U. S. Natl. 

 Herb. 29: 54-55. 1944. 



Stems and leaves with stellate and/or variously branched hairs, sometimes incon- 

 spicuously pubescent or nearly glabrous throughout; simple hairs rarely 

 present. 



Appendages absent on the truncate calyx; leaves whitish below. . . .L. hypoleuca. 

 Appendages present on the truncate calyx; leaves green, pale yellow or reddish 



below. 

 Appendages of the calyx relatively long, 8-12 mm. long, covered with long 



simple, furcate and dichotomously branched hairs L. purpusii. 



Appendages of the calyx short, linear or sometimes knob-like, 4 mm. long or 

 less, covered with relatively short stipitate-stellate or sessile-stellate 

 hairs. 



Appendages of the calyx about as broad as long, knob-like and glandular. 



L. ocellata. 

 Appendages of the calyx linear or subulate-linear, longer than broad, not 



knob-like nor glandular. 

 Stamens subequal in length; stems and lower surface of leaves densely 



covered with reddish to reddish-brown hairs. 



Leaves broadly ovate to ovate or ovate-elliptic; fruits 10-15 mm. in 

 diameter. . L. limitanea. 




48 FIELDIANA: BOTANY, VOLUME 24 



Leaves lanceolate to narrowly elliptic; fruits 7 mm. in diameter. 



L. cuchumatanensis. 



Stamens unequal in length, one anther evidently projecting above the 

 others; stems and lower surface of leaves pubescent with yellowish to 

 yellowish-brown hairs or sometimes glabrous. 



Corolla purplish; long stamen exceeding other four by (2-) 3-4 mm.; 

 pubescence of the calyx minute, nature of the hairs discernible 



only with the aid of a microscope L. lenta. 



Corolla white; long stamen exceeding other four by 1-1.5 mm.; pubes- 

 cence of the calyx long, nature of the hairs nearly always dis- 

 cernible with the naked eye in flowering specimens. 

 Larger leaves narrow, narrowly elliptic to lanceolate or lance-elliptic, 

 moderately to sparsely pubescent or glabrous below; flowering 

 pedicels 15-25 mm. long; ranging from 1,300-2,400 m. 



L. chiapensis. 



Larger leaves broad, elliptic to ovate-elliptic or ovate-lanceolate, 



densely to moderately pubescent below; flowering pedicels 10-16 



mm. long; Caribbean lowlands, below 1,000 m. . . .L. armentalis. 



Stem and leaves with simple hairs, sometimes inconspicuously pubescent or 



glabrous throughout; stellate hairs absent. 

 Appendages absent on the truncate calyx or inconspicuous (knob-like). 



Leaves lustrous, in pairs, very different in size and shape; plants glabrous 



throughout, usually epiphytic L. nitida. 



Leaves dull, solitary or in pairs, different in size, but similar in shape; plants 



pubescent, rarely glabrous throughout, terrestrial. 



Appendages of the calyx absent; the calyx sometimes inconspicuously 

 nerved; just above sea level to 900 m., rarely higher; widespread. 



L. synanthera. 



Appendages of the calyx present (knob-like); 1,500-3,000 m.; endemic. 

 Leaves evidently puberulent below; filaments 1 mm. long, the anthers 



acute; young branches sparsely puberulent L. gongylodes. 



Leaves glabrous or inconspicuously puberulent below; filaments 3-3.5 



mm. long, the anthers obtuse; young branches scurfy . L. ceratocalycia. 



Appendages conspicuously present (linear or subulate-linear) on the truncate 



calyx. 



Stamens unequal in length, one anther evidently projecting above the others. 

 Appendages of the calyx connate at their bases, strongly reflexed in fruit. 



L. connata. 

 Appendages of the calyx distinct at their bases, erect to spreading in fruit. 



Plants glabrous throughout L. chiapensis. 



Plants evidently pubescent, at least in part. 



Plants herbaceous; inflorescences consisting of a solitary flower, the 

 pedicels 6-8 cm. long; anthers 5.5 mm. long or more; fruit ellip- 

 soid, acute at the apex L. ciliolata. 



Plants woody vines or shrubs; inflorescences consisting of 2-15 flowers, 

 the pedicels 0.5-3 cm. long; anthers 4 mm. long or less; fruit glo- 

 bose. 



Long calyx appendages 9.5-11 mm. long Lycianthes sp. 



Long calyx appendages 1-4 mm. long. 



Leaves densely pubescent below, the hairs long; long calyx append- 

 ages 1-2 mm. long; seeds entire, 2-2.5 mm. long. 



L. arrazolensis. 



Leaves sparsely pubescent below, the hairs short; long calyx 

 appendages mostly 2.5-4 mm. long; seeds slightly notched, 



3-4 mm. long L. tricolor. 



Stamens subequal in length. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 49 



Leaves glabrous above and below or with hairs in tufts in the axils of the 

 mid-vein below; inflorescences consisting of 3-5 flowers. 



Corolla 7.5-8 mm. wide; leaves glabrous below L. orogenes. 



Corolla 12 mm. wide; leaves with hairs in tufts in the axils of the mid-vein 

 below L. barbatula. 



Leaves pubescent above and below, the hairs sometimes inconspicuous, 

 without hairs in tufts in the axils of the mid-vein below; inflorescences 

 usually consisting of 1-2 flowers. 



Appendages of the calyx 9 mm. long or more, densely pubescent, the hairs 

 long, at least some glanduliferous and 2 mm. long or more. 



L. gorgonea. 



Appendages of the calyx 5 mm. long or less, if densely pubescent the hairs 

 0.5 mm. long or less, eglandular. 



Corolla limb 1-1.2 cm. wide, nearly parted to the base; leaves generally 

 narrowly elliptic, inequilateral. 



Stem with spreading hairs; pedicels 1 cm. long or less; appendages 

 of the calyx 2.5 mm. long or more L. amatitlanensis. 



Stem with appressed or subappressed hairs; pedicels 2-2.5 cm. long; 

 appendages of the calyx 1 mm. long or less L. inconspicua. 



Corolla limb 2-3.5 cm. wide, shallowly lobed to lobed one-half way to 

 the base; leaves generally ovate to lanceolate, equilateral. 



Corolla limb 2 cm. wide; anthers coherent; leaves solitary; below 

 1,000 m., Caribbean lowlands L. stephanocalyx. 



Corolla limb 2.5-3.5 cm. wide; anthers free; leaves mostly in pairs; 

 ranging from 2,200 to 3,900 m., mountains of Guatemala and 

 Chiapas L. quichensis. 



Lycianthes amatitlanensis (Coult. & Donn.-Sm.) Bitter, Abh. 

 Naturwiss. Vereine Bremen 24: 441. 1920. Solanum amatitlanense 

 Coult. & Donn.-Sm. Bot. Gaz. 37: 420. 1904 (syntypes, Barranca de 

 Eminencia, Guatemala, Donn.-Sm. 1457; Cubilqiiitz, AltaVerapaz, 

 Tuerckheim 7753 & 8^88). 



Wet forests, 200-900 m., rarely higher; Pete"n; Alta Verapaz; 

 Izabal; Chimaltenango; Huehuetenango; Escuintla. Mexico (Chi- 

 apas) ; Atlantic coast of Honduras. Panama. 



Erect suffrutescent herbs or shrubs, 1-1.5 m. tall, simple or little branched, 

 the stems hirsute; leaves in pairs, very unequal in size, sparsely hirsute above, 

 densely so below, especially on the veins, the hairs pale yellow, the larger leaves 

 in equilateral, elliptic or lanceolate to lance-oblong, 10-24 cm. long, 3-8 cm. wide, 

 long acuminate, cuneate and unequal at the base, the petioles 6-10 mm. long, 

 densely hirsute; smaller leaves lanceolate to suborbicular, 1-3 cm. long, 8-15 mm. 

 wide, acute, subsessile; inflorescences consisting of 1-5 flowers; pedicels 6-10 mm. 

 long, densely hirsute; calyx campanulate, 1.5-2 mm. long, densely hirsute, with 

 10 linear appendages, these 2.5-3 mm. long, subequal; corolla pale yellow or white, 

 the limb 13-16 mm. wide, parted to near the base, the lobes lanceolate, 6-7 mm. 

 long, pilose with multicellular hairs externally; stamens equal, the filaments 1.5 

 mm. long, the anthers 3-4 mm. long, acute; style 5-6 mm. long, exceeding the 

 stamens by about 1 mm.; berry red, subglobose, glabrous, 5-6 mm. in diameter. 



See comments under Lycianthes inconspicua. 




50 FIELDTANA: BOTANY, VOLUME 24 



Lycianthes armentalis J. L. Gentry, Phytologia 26: 269. 1973. 



In advanced deciduous forest, below 1,000 m.; Pete*n. British 

 Honduras. Southern Mexico, Yucatan Peninsula (type from Quin- 

 tana Roo, C. L. & A. A. Lundell 7800). 



A clambering shrub, the branches slender, 1-7 m. long, with short, stipitate- 

 stellate hairs, a few shorter, simple hairs intermixed, the hairs reddish-brown 

 to yellowish-brown; leaves mostly in pairs, unequal in size, similar in shape, 

 elliptic to ovate-elliptic or ovate-lanceolate, sometimes suborbicular, sparsely 

 and inconspicuously short stipitate-stellate pubescent above, densely short stipi- 

 tate-stellate pubescent below, the hairs yellowish-brown, the larger leaves 4-6.5 

 (-14.5) cm. long, 2.5-4.5(-6) cm. wide, acute or acuminate, the base obtuse, the 

 petioles 3-12 mm. long; inflorescences consisting of 3-5 flowers, the pedicels 10- 

 16 mm. long, short stipitate-stellate pubescent, with a few shorter, simple hairs 

 intermixed; calyx 3 mm. long, short stipitate-stellate pubescent, with 10 alternately 

 subequal appendages, the long appendages 2.5-3.5 mm. long, the short ones 2-2.5 

 mm. long; corolla probably white, the limb 15 mm. wide, lobed to the middle, the 

 lobes 3.5 mm. long, sparsely stellate-pubescent externally; stamens unequal, the 

 filaments of the short stamens 1.5 mm. long, the fifth one 3 mm. long, the anthers 

 3.5-4 mm. long, sparsely stellate-pubescent abaxially or glabrous; style 7.5-8 mm. 

 long, exceeding the long stamen; berry subglobose 7-9 mm. in diameter; seeds 

 2.2-3 mm. long. 



This species has been confused with Lycianthes sideroxyloides 

 Schlecht. in herbaria. It differs from that species by its longer calyx 

 appendages, unequal stamens, and fewer flowers in the inflorescences. 



Lycianthes arrazolensis (Coult. & Donn.-Sm.) Bitter, Abh. 

 Naturwiss. Vereine Bremen 24: 388. 1920. Solanum arrazolense 

 Coult. & Donn.-Sm. Bot. Gaz. 37: 421. 1904. 



Moist thickets or forests, sometimes in mixed forests, 1,000- 

 3,000 m.; Alta Verapaz; Chiquimula; Baja Verapaz; Guatemala 

 (type from Arrazola, Heyde & Lux 4-736); Sacatepe"quez; Chimal- 

 tenango; Solola; Quezaltenango ; Huehuetenango; Suchitepe"quez ; 

 San Marcos. Mexico. Honduras. 



Erect or subscandent shrubs, rarely herbaceous, 1-3.5 m. tall, often much 

 branched, the young branches densely pilose, occasionally sparsely pilose or 

 merely puberulent or pilose and puberulent; leaves in pairs or solitary, similar 

 in shape, unequal in size, densely pilose above and below, more densely so below, 

 sometimes velutinuous, rarely sparsely pubescent, the larger leaves 6.5-16.5 cm. 

 long, 3-7.5 cm. wide, acuminate, attenuate at the base, equal or unequal, the pet- 

 ioles 5-15(-20) mm. long; smaller leaves 3-8.5 cm. long, 1-3.5 cm. wide; inflores- 

 cences consisting of 3-15 flowers, the pedicels 5-20 mm. long; calyx campanulate, 

 2.5-3 mm. long, with 10 appendages, alternately subequal, the long appendages 

 1-2 mm. long, the short ones 0.5-1.5 mm. long; corolla white, the limb 1.2-2.5 cm. 

 wide, scarcely lobed, the lobes 1-2 mm. long, densely pubescent externally; sta- 

 mens unequal, the filaments of the short stamens 1.5-2 mm. long, the fifth one 3-4 




FIG. 11. Lycianthes barbatula. A, habit, X 3^; B, flower, X 2; C, corolla dis- 

 sected, X 3; D, calyx, style, and stigma, X 3; E, anther, abaxial view, X 6. 



51 




52 FIELDIANA: BOTANY, VOLUME 24 



mm. long, the anthers of the short stamens 2.5-3 mm. long, the fifth one 2.5-3.5 

 mm. long; style 7-8.5 mm. long, exceeding the longest stamen about 1 mm.; berry 

 red, globose, 5-6 mm. in diameter; seeds 2-2.5 mm. long, entire. 



Lycianthes arrazolensis can in most cases be separated from L. 

 tricolor on the basis of the more densely pubescent leaves. However, 

 a few specimens are difficult to place. The size and margin of the 

 seeds have proven to be useful characters in separating these species. 

 In L. arrazolensis the seeds are 2-2.5 mm. long and entire, whereas 

 in L. tricolor the seeds are 3-4 mm. long and slightly notched. 



Lycianthes barbatula Standl. & Steyerm. Field Mus. Bot. 23: 

 228. 1947. 



Moist or wet thickets or forests, 1,200-2,650 m.; Chimaltenango 

 (Volcan de Fuego) ; Suchitepe'quez (type from Volcan de Santa Clara, 

 Steyermark If6653). Endemic. 



Shrubs 3-4.5 m. tall, the branches slender, glabrous; leaves thick-membranace- 

 ous, mostly in pairs, unequal in size and different in shape, glabrous above, the 

 hairs in tufts in the axis of the mid-vein, sometimes along the mid-vein with long, 

 pale brownish hairs, otherwise glabrous below, the larger leaves elliptic to elliptic- 

 oblong, 5-18 cm. long, 2-6.5 cm. wide, acuminate or acute, short attenuate at the 

 base, sometimes unequal, the petioles 5-20 mm. long; smaller leaves broadly ovate 

 or suborbicular, 1-3 cm. long, 8-20 mm. wide, obtuse or rounded at the apex; 

 inflorescences consisting of 3-6 flowers, the pedicels slender, 2-2.5 cm. long, glab- 

 rous; calyx broadly campanulate, 2-2.5 mm. long, glabrous, with 5 (-7) linear, sub- 

 equal appendages, 2-3 mm. long; corolla white, the limb 12 mm. wide, lobed not 

 quite to the middle, glabrous, the lobes lanceolate, about 4 mm. long; stamens 

 equal, the filaments 3 mm. long; anthers 3 mm. long; style 7.5 mm. long, exceeding 

 stamens; berry white, globose, glabrous, nearly 1 cm. in diameter. 



Lycianthes barbatula has its nearest relative in L. orogenes from 

 which it is distinguished by the leaves with hairs in tufts in the axils 

 of the mid-vein below, longer calyx appendages, and longer fila- 

 ments. Both of these species are rather poorly known. Further 

 collections may warrant the treatment of L. orogenes as a synonym 

 of L. barbatula. 



Lycianthes ceratocalycia (Donn.-Sm.) Bitter, Abh. Naturwiss. 

 Vereine Bremen 24: 498. 1920. Brachistus ceratocalycius Donn.-Sm. 

 Bot. Gaz. 48: 297. 1909. 



Open areas in forests, 1,500-1,766 m., possibly higher; Alta Vera- 

 paz )type from near Coban, Tuerckheim II 2060); Quiche"; Huehue- 

 tenango. Endemic. 



A straggling shrub 2 m. tall, the young branches inconspicuously pubescent 

 with minute incurved or subappressed hairs, somewhat scurfy; leaves solitary or 




GENTRY & STANDLEY: FLORA OF GUATEMALA 53 



in pairs, then unequal in size, glabrous above and below, the larger leaves nar- 

 rowly elliptic or lanceolate, 7.5-12 cm. long, 2-3 cm. wide, acuminate, the base 

 attenuate, sometimes inequilateral, the petioles 8-12 mm. long, puberulent; smaller 

 leaves similar or different in shape, 3.5-5 cm. long; inflorescences consisting of 

 4-12 flowers, rarely solitary, the pedicels 1-2 cm. long, puberulent; calyx cam- 

 panulate, 3-3.5 mm. long, glabrous, with 5-10 knob-like appendages, these 

 0.3-0.5 mm. long; corolla pale lavender, the limb 2-2.4 cm. wide, parted to near 

 the base, the lobes lanceolate, 8-12 mm. long, puberulent or nearly glabrous ex- 

 ternally; stamens equal, the filaments 3-3.5 mm. long, the anthers 4-5 mm. long, 

 about 2 mm. wide, obtuse at the apex; style about 9 mm. long, exceeding the 

 stamens by 2-3.5 mm.; berry unknown. 



Lycianthes ceratocalycia is distinguished from other species of the 

 genus in Guatemala by its somewhat scurfy young branches, nodular 

 calyx appendages, broad anthers, and long filaments. 



Lycianthes chiapensis (Brandegee) Standl. Field Mus. Bot. 11: 

 173. 1936. Solanum chiapense Brandegee, Univ. Calif. Publ. Bot. 

 6: 192. 1915. L. nyssifolia Bitter, Fedde Repert. Sp. Nov. 20: 366. 

 1920 (type from Las Nubes, Suchitepe"quez, Bernoulli & Cario 2397) . 

 L. chiapensis var. sparsistellata Standl. & Steyerm. Field Mus. Bot. 

 22: 274. 1940 (type from Cerro Tixixi, north of Jocotan, Chiquimula, 

 Steyermark 31555], syn. nov. 



Usually in dense wet forests, sometimes in mixed forests, 1,300- 

 2,400 m.; Zacapa; Chiquimula; Quezaltenango; Huehuetenango ; 

 Suchitepe'quez; San Marcos. Mexico (type from Finca Irlanda, 

 Chiapas, Purpus 7328). 



Vines or scandent shrubs, with short stipitate-stellate, sometimes furcate or 

 branched hairs or essentially glabrous throughout, the slender branches to 6 m. 

 long; leaves solitary or sometimes in pairs, unequal in size, similar or different in 

 shape, usually more densely pubescent below, or glabrous, sometimes the hairs 

 very inconspicuous, the larger leaves narrowly elliptic or lanceolate to lance- 

 elliptic, 3.5-11 cm. long, 1.5-3(-4.5) cm. wide, acuminate or narrowly acute, acute 

 or short attenuate at the base, equal or unequal, the petioles 5-20 mm. long; smal- 

 ler leaves similar in shape or sometimes suborbicular, 2-5(-8) cm. long, l-2(-3) cm. 

 wide, acute or obtuse; inflorescences consisting of 2-3 (-5) flowers, the pedicels 

 1.5-2.5 cm. long; calyx campanulate, 3-6 mm. long, with short stipitate-stellate 

 and a few branched hairs or glabrous, with 10 alternately subequal appendages, 

 the long appendages 1.5-2(-4) mm. long, the short ones somewhat lower on the 

 calyx and 0.5-1.5(-3) mm. long; corolla white, the limb 2 cm. wide, lobed to about 

 the middle, pubescent with minute stellate, branched and simple hairs externally; 

 stamens unequal, the filaments of the short stamens 1.5 mm. long, the fifth 2.5 

 mm. long, the anthers 4.5-5 mm. long; style 7-10 mm. long, shorter than to ex- 

 ceeding the long stamen; berry orange-red, obovoid or subglobose, glabrous, 7-12 

 mm. in diameter, the seeds 3.5-4.5 mm. long, reticulate-foveate. 




54 FIELDIANA: BOTANY, VOLUME 24 



More material is required for a better understanding of this highly 

 variable species. Lycianthes nyssifolia may prove to be a distinct 

 species. 



Lycianthes ciliolata (Mart. & Gal.) Bitter, Abh. Naturwiss. 

 Vereine Bremen 24: 410. 1920. Solanum ciliolatum Mart. & Gal. Bull. 

 Acad. Brux. 12, pt. 1: 140. 1845. L. ciliolata var. pratorum Bitter, 

 1. c. 411 (syntypes from Fatal, Baja Verapaz, Tuerckheim 1434 & II 

 2317). L. guatemalensis Bitter, 1. c. 419 (syntypes from Retalhuleu, 

 Bernoulli & Carlo 2384; San Sebastian, Bernoulli & Cario 24-04, 

 Retalhuleu). Chile cerrano. 



Wet thickets or fields, sometimes in oak-pine woods, 2,400-2,800 

 m., sometimes lower; Baja Verapaz; Retalhuleu; Huehuetenango. 

 Southern Mexico. 



Plants herbaceous, 1 m. tall or less, the stem branched, sparsely pilose, the 

 hairs multicellular and incurved; leaves solitary below, in pairs above, then un- 

 equal in size, similar in shape, pilosulous above, sparsely so below, the hairs multi- 

 cellular, the larger leaves ovate to ovate-lanceolate or ovate-rhomboid, 4.5-12.5 

 cm. long, 2-6 cm. wide, acuminate or acute, the base attenuate, the petioles 5-15 

 mm. long, sparsely pilosulous, winged nearly to the base; smaller leaves 3-7 cm. 

 long, 1.5-3.5 cm. wide; inflorescences consisting of a solitary flower, the pedicels 

 sparsely pilosulous or glabrous, reflexed in age, 6-8 cm. long; calyx campanulate, 

 3.5 mm. long, pilosulous, 10-nerved, the 10 appendages linear-subulate, 5-8 mm. 

 long; corolla white with lavender, the limb 2.5-4(-5) cm. wide, the lobes short, 

 about 2 mm. long, puberulent externally; stamens unequal, the filaments of the 

 short stamens 3-3.5 mm. long, the fifth one 6-7 mm. long, the short anthers 5- 

 5.5 mm. long, the long one 5.5-6 mm. long; style 11 mm. long, slightly shorter 

 than to barely exceeding the longest stamen; berry ellipsoid, 2-3.5 cm. long, apex 

 acute. 



Lycianthes ciliolata is most likely to be confused with L. quichensis 

 in general aspect, but it is readily distinguished by the unequal 

 stamens, ellipsoid fruits, and habit. 



Lycianthes guatemalensis is known to us only from the original 

 description and a photograph of a syntype. It appears to fall within 

 the morphological range of L. ciliolata, however, further collecting 

 may prove it to be a distinct species. 



Lycianthes connata J. L. Gentry, Phytologia 26: 271. 1973. 



Cloud forest, 2,000-3,000 m.; Huehuetenango (type from the 

 Sierra de los Cuchumatanes, Cruz de Limon, Steyermark 49828). 

 Mexico (Chiapas). 



Shrub 2-7 m. tall, the slender branches sparsely and inconspicuously pubes- 

 cent, the hairs incurved or glabrous; leaves in pairs or solitary, mostly similar in 




GENTRY & STANDLEY: FLORA OF GUATEMALA 



55 



FIG. 12. Lycianthes connata. A, branch, X %; B, fruit, X 2, and seeds, X 3; 

 C, calyx, X 2 1 A; D, dissected corolla, X 2>; E, flower and detail of anther, X 2>. 



shape, unequal in size, sparsely and inconspicuously pubescent above and below, 

 the hairs incurved on the veins below or glabrous, the larger leaves elliptic, 7-15 

 (-26.5) cm. long, 3-4(-8) cm. wide, acuminate, attenuate at the base, equal or 

 unequal, the petioles 3-10(-40) mm. long; smaller leaves 4-9 cm. long, 2-3.5 cm. 

 wide; inflorescences consisting of 1-4 flowers, the pedicels slender, 2.5-5 cm. long; 

 calyx campanulate, 2.5-3 mm. long, glabrous or sparsely pubescent, with 10 sub- 

 ulate-linear appendages, alternately subequal, connate at their bases, spreading or 

 slightly reflexed, the long appendages 2.5-4 mm. long, the short ones 1-2 mm. long; 




56 FIELDIANA: BOTANY, VOLUME 24 



corolla white, the limb 2-3 cm. wide, glabrous externally, scarcely lobed, the lobes 

 2 mm. long; stamens unequal, the filaments of the short stamens 1-2 mm. long, the 

 fifth one 2.5-3 mm. long, the anthers 4.5-5.5 mm. long; style 9.5-10.5 mm. long, 

 exceeding the stamens by about 1.5 mm.; berry orange, globose, 8.5-10 mm. in 

 diameter, the connate appendages strongly reflexed; seeds 2-2.5 mm. long. 



A collection from Alta Verapaz, Tuerckheim 3935, has been ten- 

 tatively placed here. It is morphologically similar to Lycianthes con- 

 nata but differs in characters of the calyx appendages. This may 

 prove to be a distinct taxon. 



Lycianthes cuchumatanensis J. L. Gentry, Phytologia 26: 

 273. 1973. 



1,500-2,600 m. Endemic. Huehuetenango (type from Cerro Huitz, 

 Sierra de los Cuchumatanes, Steyermark 48925) . 



A vine, the young branches, petioles, pedicels, and calyces densely pubescent 

 with ferruginous, stellate, short stipitate-stellate and branched hairs; leaves sub- 

 coriaceous, solitary, lanceolate to lance-elliptic, 5-10 cm. long, 2-3.5 cm. wide, 

 acuminate, cuneate at the base, equal or unequal, moderately pubescent above, 

 densely so below, especially on the veins, the petioles 5-10 mm. long; inflorescences 

 consisting of 2-4 flowers, the pedicels 1 cm. long; calyx campanulate, 3.5 mm. 

 long with 10 spreading, linear appendages, 1.5 mm. long; corolla limb 1.5 cm. 

 wide, lobed to the middle, the lobes lanceolate, 4 mm. long, densely pubescent 

 with minute stellate hairs externally; stamens equal, the filaments 1 mm. long, the 

 anthers 3.5 mm. long, free, sparsely pubescent below; style 7 mm. long, exceeding 

 the stamens by 1.5 mm.; berry subglobose, glabrous, about 7 mm. in diameter; 

 seeds 2.5-3 mm. long. 



Closely related to Lycianthes limitanea. 



Lycianthes gongylodes J. L. Gentry, Phytologia 26: 274. 1973. 



Cloud forest, 2,600-3,000 m.; Huehuetenango (type from the 

 Sierra de los Cuchumatanes, 4 miles east of San Mateo Ixatan on 

 road to Barillas, Breedlove 8771). Endemic. 



Shrubs 1.5-3.5 m. tall, the young branches sparsely puberulent; leaves soli- 

 tary or in pairs, then similar in shape, unequal in size, puberulent above and below, 

 more densely so below on the veins, the larger leaves elliptic to elliptic-ovate or 

 ovate, 10.5-17.5 cm. long, 3.5-6.5 cm. wide, acuminate, the base shortly attenuate, 

 equal or unequal, the petioles 1-2.5 cm. long; smaller leaves 7-10 cm. long, 3.5-4 

 cm. wide; inflorescences consisting of 4-8 flowers, the pedicels 1-1.5 cm. long; calyx 

 campanulate, 3 mm. long, glabrous, with 10 knob-like appendages, 0.5 mm. long 

 or less; corolla white, the limb 2 cm. wide, lobed to below the middle, the lobes 6 

 mm. long, ciliate, sparsely pubescent externally; stamens equal, the filaments 1 

 mm. long, the anthers 5 mm. long; style 7 mm. long, exceeding the stamens by 1 

 mm.; berry orange, subglobose, glabrous, about 8 mm. in diameter; seeds 2-2.5 

 mm. long, reticulate-foveate. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 57 



Lycianthes gorgonea Bitter, Fedde Repert. Sp. Nov. 20:364. 

 1920. L. cuspidata (Morton) Standl. & Steyerm. Field Mus. Bot. 

 23: 18. 1943. Solanum cuspidatum Morton in Lundell, Contr. Univ. 

 Mich. Herb. 4: 25. 1940 (type from Arenal-Valentin road, El Cayo 

 District, British Honduras, Lundell 6172), syn. nov. 



Moist or wet thickets, 200-335 m.; Pete"n (type from Sacluc, 

 Bernoulli & Cario 2357} ; Alta Verapaz. British Honduras. 



Erect or scandent shrubs, densely pubescent throughout with the hairs of 

 varying lengths, at least some hairs glandular, the stems slender, 1-3 m. long or 

 tall; leaves mostly in pairs, very unequal in size, different in shape, densely 

 pubescent above and below, the larger leaves ovate-lanceolate, or lanceolate, 6- 

 10 cm. long, 2-4 cm. wide, long acuminate, obtuse or acute at the base, sometimes 

 unequal, the petioles 6 mm. long or less; smaller leaves broadly ovate or suborbi- 

 cular, 1-3 cm. long, 1-1.8 cm. wide, obtuse or acute, the petioles 2-3 mm. long; 

 inflorescences consisting of 1 or 2 flowers, the pedicels 2.5-3.5 cm. long, slender, 

 densely pubescent; calyx campanulate, 2 mm. long, densely pubescent, with 10 

 filiform appendages, alternately subequal, densely pubescent, the longest append- 

 ages 11-15 mm. long, the shortest ones 9-12 mm. long; corolla white, the limb 

 about 2.5 cm. wide, the lobes broad triangular, about 4 mm. long, sparsely pilose 

 externally; stamens subequal, the filaments 1-1.5 mm. long, anthers coherent, 4- 

 4.5 mm. long; style about 5.5 mm. long; berry red globose, 8-10 mm. in diame- 

 ter, greatly surpassed by the accrescent calyx appendages. 



Lycianthes gorgonea is distinguished from other members of the 

 genus in Central America by the following combination of charac- 

 ters: densely pubescent throughout with pale, translucent, simple 

 hairs of varying lengths, at least some hairs glandular (the glands 

 often only discernible with difficulty); leaves mostly in pairs and 

 unequal in shape and size; calyx appendages very long, greatly ex- 

 ceeding the fruits. 



A species known only from the Caribbean lowlands. 



Lycianthes hypoleuca Standl. Trop. Woods 9: 12. 1927. 



In advanced forest, usually on limestone hillsides, 150-700 m.; 

 Peten; Alta Verapaz. British Honduras (type from Orange Walk 

 District, Winzerling V-llf). 



A scandent shrub or vine, sometimes an erect shrub, 5 m. tall, the branchlets 

 slender, flexuous, dark reddish brown, when young sparsely tomentose with min- 

 ute, whitish, stellate hairs; leaves solitary or in pairs, then similar in shape, some- 

 what smaller in size, green above and appearing glabrous but bearing scattered 

 minute stellate hairs, the lower surface whitish or silvery, densely covered with a 

 minute tomentum of appressed stellate hairs, elliptic or elliptic-ovate, mostly 5- 

 7.5 cm. long, 2-3.5 cm. wide, acute to acuminate, attenuate at the base, the 

 petioles slender, 1 cm. long or less; inflorescences consisting of mostly 2-4 




58 FIELDIANA: BOTANY, VOLUME 24 



flowers, sometimes only 1, the pedicels slender, 2-3 cm. long, sparsely and min- 

 utely stellate pubescent; calyx campanulate, 2.5-3 mm. long, sometimes slightly 

 lobed, sparsely to densely pubescent with minute, stellate hairs, appendages 

 absent; corolla white, the limb about 16 mm. wide, slightly lobed, the lobes lance- 

 olate, the apex pubescent with minute brownish branched hairs externally; sta- 

 mens equal, the filaments 1.5 mm. long, the anthers 3.5-4 mm. long; style 8.5 mm. 

 long, exceeding stamens by 3-4 mm.; berry red, subglobose, glabrous, 7-8 mm. in 

 diameter. 



This species of the Caribbean lowlands can be readily distin- 

 guished from the other species of the genus in Central America by 

 the whitish or silvery tomentum of stellate hairs on the lower surface 

 of the leaves. 



Lycianthes inconspicua Bitter, Fedde Repert. Sp. Nov. 20: 368. 

 1924. Chiltepe de montana. 



In forests, 1,300-1,870 m.; Quezaltenango (type from Santa 

 Maria, Bernoulli & Cario 2373} ; San Marcos. Endemic. 



Shrub 1.5-3.5 m. tall, the branches slender, with appressed and sub-appressed 

 pale yellowish hairs; upper leaves in pairs, very unequal in size, sparsely strigillose 

 above, hirsutulous below, especially on the veins, the hairs subappressed and pale 

 yellow, the larger leaves lanceolate to broadly lanceolate or elliptic to elliptic- 

 oblong, 3-20 cm. long, 1-4.5 cm. wide, acuminate, cuneate at the base, unequal, 

 the petioles 2-10 mm. long; smaller leaves ovate to lanceolate or oblong, 1-7.5 cm. 

 long, 5-25 mm. wide, acute or obtuse, subsessile; inflorescences consisting of 1-3 

 flowers, the pedicels slender, strigose, solitary, 2-2.5 cm. long; calyx campanulate, 

 1.5-3 mm. long, strigillose, the 10 appendages subulate-linear, unequal, the al- 

 ternate ones subequal, 0.4-1 mm. long or appendages 5; corolla white, the limb 

 9-12 mm. wide, deeply lobed, the lobes lanceolate, 4.5-5 mm. long, sparsely pubes- 

 cent with appressed and subappressed hairs externally; stamens equal, the fila- 

 ments 1 mm. long, the anthers 2.5 mm. long; style 3.8-4.5 mm. long, slightly ex- 

 ceeding the stamens; berry red, subglobose, glabrous, about 8 mm. in diameter. 



Lycianthes inconspicua is apparently related to L. amatitlanensis, 

 and is distinguished from that species by the characters listed in the 

 key. It is also found at somewhat higher elevations. 



Lycianthes lenta (Cav.) Bitter, Abh. Naturwiss. Vereine Bre- 

 men 24: 364. 1920. Solanum lentum Cav. Icon. 4: 4, t. 308. 1797. L. 

 variifolia Standl. Field Mus. Bot. 4: 259. 1929 (type from Tower 

 Hill, British Honduras, Karling 13), syn. nov. 



Wet to dry thickets, often on rocky slopes, sometimes on sand 

 dunes, near sea level to 1,000 m., rarely higher; Pete"n; Alta Verapaz; 

 Izabal; Zacapa; El Progreso; Chiquimula; Jalapa; Guatemala; Que- 

 zaltenango; Huehuetenango; Jutiapa; Santa Rosa; Escuintla; Re- 




GENTRY & STANDLEY: FLORA OF GUATEMALA 59 



talhuleu; San Marcos. British Honduras. Southern Mexico to Costa 

 Rica. Cuba. 



A woody vine or slender shrub, the young branches 2-3 m. long, covered with 

 minute, sessile and short stipitate-stellate hairs; leaves solitary or in pairs, then 

 unequal in size, similar in shape, the hairs short stipitate-stellate above and below, 

 more dense and usually velvety and soft to the touch below, sometimes with a few 

 simple and furcate hairs present with the stellate ones, the larger leaves usually 

 ovate to ovate-deltoid or elliptic, sometimes suborbicular or obovate, 2. 5-10 (-14) 

 cm. long, 1.5-4 (-6) cm. wide, acute or sometimes obtuse, usually subcordate to 

 nearly truncate or short attenuate at the base, equal or unequal, the petioles 5- 

 25 mm. long; smaller leaves 2.5-6 cm. long, 1-2.5 cm. wide; inflorescences con- 

 sisting of 2-6(-9) flowers, the pedicels 1-2 cm. long; calyx campanulate, 2-3.5 mm. 

 long, the hairs sessile-stellate or short stipitate-stellate, the 10 appendages linear- 

 subulate, 2-4 mm. long, alternately subequal; corolla purplish, the limb 2-3.5 cm. 

 wide, shallowly lobed, densely stellate-pubescent externally; stamens unequal, 

 the filaments of the short stamens 1-1.5 mm. long, the fifth one (2.5-)3.5-5.5 mm. 

 long, the anthers of the short stamens 4-5.5 mm. long, the long one (3.5-)4.5-6 

 mm. long; style about 1 cm. long, exceeding the long stamen by 1.5 mm. to slightly 

 shorter; berry red or orange, 8-12 mm. in diameter; seeds 2-2.5 mm. long, reticu- 

 late-foveate. 



Lycianthes lenta is a wide ranging species; southern Mexico to 

 Costa Rica and Cuba, being collected mostly below 1,000 m. It is 

 quite variable in both leaf shape and pubescence. Bitter (1920) 

 recognized six infraspecific taxa in his treatment of Lycianthes. It 

 is confused sometimes with L. sideroxyloides Schlecht., a species not 

 known to occur in Central America. 



We have included here two collections from British Honduras, 

 Gentle 3506 and Schipp 311. They differ from Lycianthes lenta by 

 their white flowers, somewhat larger leaves and more densely pubes- 

 cent pedicels and calyx. These collections may represent an un- 

 described taxon. 



Known in El Salvador by the names manzana, montes, quaco, and 

 quesillo. 



Lycianthes limitanea (Standl.) J. L. Gentry, Phytologia 26: 

 275. 1973. Solanum limitaneum Standl. Carnegie Inst. Wash. Publ. 

 461:85. 1935. 



Moist or wet thickets or forests, 500-1,800 m.; Izabal; Huehue- 

 tenango. British Honduras (type from Camp 33, British Honduras- 

 Guatemala boundary, Schipp S-681). 



A large, woody vine, unarmed, 10 m. long or less, the thick branches, petioles, 

 pedicels and calyces densely covered with reddish-brown, short stipitate-stellate 

 and stipitate-branched hairs; leaves solitary or sometimes in pairs, similar in 




60 FIELDIANA: BOTANY, VOLUME 24 



shape, nearly equal in size, the larger leaves broadly ovate to ovate or ovate-el- 

 liptic, (6.5-)10-17.5 cm. long, (3-)6-9.5 cm. wide, acuminate, rounded at the base 

 and equal, short stipitate-stellate pubescent, lustrous and asperous above, densely 

 covered with reddish-brown, short stipitate-stellate hairs below, especially on the 

 veins; petioles thick, 1-2.5 cm. long; inflorescences few-flowered; corollas unknown; 

 fruiting pedicels thick, 1-2 cm. long; calyx in fruit 6-7 mm. long, 10-12 mm. wide, 

 broadly campanulate, with 5 (?) appendages, 1-1.5 mm. long; fruit yellow, globose, 

 1-1.5 cm. in diameter, glabrous; seeds 3-3.5 mm. long. 



Lycianthes limitanea is known from only fruiting specimens. It 

 is closely related to L. cuchumatanensis and is distinguished by the 

 leaf shape and the much larger fruits. 



Lycianthes nitida Bitter, Abh. Naturwiss. Vereine Bremen 

 24: 501. 1920. Solanum calochromum Blake, Contr. U. S. Natl. Herb. 

 24: 21. 1922 (type from along trail from Hacienda El Limon to El 

 Paraiso, Copan, Honduras, Blake 7370). 



Mostly wet lowland forests, 25-350 m., rarely higher; Izabal; 

 Alta Verapaz (syntypes Cubilquitz, Tuerckheim II 59 and Donn.-Sm. 

 7637). Huehuetenango. British Honduras. Atlantic coast of Hon- 

 duras; Costa Rica. 



An erect or scandent shrub, usually epiphytic, glabrous throughout, the 

 branches 1-5 m. long; leaves sub coriaceous, lustrous above, in pairs, very unequal 

 in size, different in shape, the larger leaves elliptic to oblong 12-26 cm. long, 4- 

 8.5 cm. wide, acuminate or abruptly acuminate, cuneate and somewhat unequal 

 at the base, sometimes inequilateral, the petioles 1.5-2.5 cm. long; smaller leaves 

 obovate or suborbicular, mostly 4-7 cm. long, 3-5 cm. wide, rounded and often 

 with a short abrupt apex; inflorescences consisting of several flowers, the pedicels 

 1.2-1.5 cm. long; calyx campanulate, 2-3 mm. long, appendages absent; corolla 

 purple or lavender, sometimes pinkish, the limb 1.6-2.2 cm. wide, parted to near 

 the base, the lobes lanceolate, 6-9.5 mm. long; stamens equal, the filaments about 

 1 mm. long, the anthers coherent, 6-7 mm. long; style about 1 cm. long; berry 

 orange or red, globose, 6-8 mm. in diameter, glabrous. 



Lycianthes nitida is characterized by its habit, large subcoriaceous 

 and lustrous leaves that are very unequal in size and different in 

 shape, and by the absence of calyx appendages. 



Lycianthes ocellata (Donn.-Sm.) Morton & Standl. Field Mus. 

 Bot. 22: 274. 1940. Solanum sideroxyloides var. ocellatum Donn.-Sm. 

 Bot. Gaz. 14: 28. 1889. L. sideroxyloides subsp. ocellata Bitter, Abh. 

 Naturwiss. Vereine Bremen 24: 405. 1920. L. sideroxyloides var. 

 transitoria Bitter, 1. c. (type from Pansamala, Alta Verapaz, Tuerck- 

 heim 923), syn. nov. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 61 



Wet forests or thickets, sometimes in wet pine forests, 1,260- 

 1,500 m., Alta Verapaz (type from Pansamala, Tuerckheim 1155}; 

 Quiche"; Huehuetenango. Endemic. 



Shrub or vine, 3 m. tall, the young branches, petioles, pedicels and calyces 

 densely pubescent with ferruginous, sessile-stellate, short stipitate-stellate and 

 branched hairs; leaves subcoriaceous and lustrous, solitary, lanceolate to ovate, 

 sometimes elliptic-ovate, 5-10.5 cm. long, 2.5-4.5 cm. wide, acuminate, rounded or 

 cuneate at the base, equal, sparsely, short stipitate-stellate pubescent above and 

 below, more densely so on the veins, the petioles 5-15 mm. long; inflorescences 

 consisting of 6-12 flowers, the pedicels about 5 mm. long; calyx campanulate, 3 

 mm. long, with 10 thick, knob-like, black appendages alternately subequal in size, 

 0.5-1 mm. long; corolla limb 1.5 cm. wide, lobed to below the middle, the lobes 

 lanceolate, 6 mm. long, densely pubescent with minute stellate hairs externally; 

 stamens equal, the filaments 1 mm. long, the anthers 4 mm. long; style 7 mm. long, 

 exceeding the stamens by 1.5 mm. long; berry subglobose, glabrous, about 7 mm. 

 in diameter; seeds about 2.5 mm. long. 



Lycianthes ocellata is distinguished from other species in Guate- 

 mala by the reddish, stellate pubescence and the black, knob-like 

 calyx appendages. 



Variety transitoria differs by its more densely pubescent leaves 

 below. The stamens are somewhat unequal but this could be due 

 to immaturity. Further collections may warrant the recognition of 

 variety transitoria as a distinct species. 



Lycianthes orogenes Standl. & Steyerm. Field Mus. Bot. 23: 

 229. 1947. 



Moist or wet forests or thickets, sometimes in mixed forest, 1,000- 

 1,800 m., rarely higher; Sacatepe"quez ; Chimaltenango (type from 

 Volcan Fuego, Steyermark 52101+); Quezaltenango. Endemic. 



Shrubs 1.5-3 m. tall or sometimes a small tree, glabrous throughout, the 

 branches slender; leaves in pairs, unequal in size, mostly different in shape, mem- 

 branaceous or thick-membranaceous, the larger leaves elliptic or oblong-elliptic, 

 sometimes lance-elliptic, 7-19.5 cm. long, 3-5.5 cm. wide, acuminate, attenuate 

 at the base, the petioles 1-2.5 cm. long; smaller leaves ovate to lance-ovate or el- 

 liptic, 3-5.5 cm. long, 2-3 cm. wide, acute, the petioles 5 mm. long or less; inflores- 

 cences consisting of 3-5 flowers, the pedicels slender, 2.5-3.5 cm. long; calyx cam- 

 panulate, 2.5-3 mm. long, with 5 linear, subequal appendages, 1-1.5 mm. long; 

 corolla white, the limb 7.5-8 mm. wide, lobed not quite to the middle, the lobes 

 lanceolate, 2 mm. long; stamens equal, the filaments 1.5-2 mm. long, the anthers 

 about 3.5 mm. long; style 6.5 mm. long, exceeding the stamens; berry green, 

 globose, about 6 mm. in diameter. 



Apparently closely related to Lycianthes barbatula. See com- 

 ments under that species. 




62 FIELDIANA: BOTANY, VOLUME 24 



Lycianthes purpusii (Brandegee) Bitter, Abh. Naturwiss. Ver- 

 eine Bremen 24: 382. 1920. Solatium purpusii Brandegee, Univ. 

 Calif. Publ. Bot. 6: 62. 1914. L. purpusii var. extensidentata Bitter, 

 Fedde Repert. Sp. Nov. 20: 365. 1924 (type from San Francisco 

 Miramare, Quezaltenango, Bernoulli & Cario 2334-) L. vulpina 

 Standl. Field Mus. Bot. 4: 321. 1929 (type from Lancetilla Valley 

 near Tela, Honduras, Standley 5^356), syn. nov. 



Moist or wet forests, 50-1,000 m.; Alta Verapaz; Izabal; Quezal- 

 tenango. British Honduras. Mexico (Chiapas). Atlantic coast of 

 Honduras. 



A scandent shrub or vine, 3-8 m. long, pubescent with stiff and spreading, red- 

 dish-brown or yellowish, long, stipitate furcate and variously branched hairs, sim- 

 ple hairs sometimes present among the branched ones; leaves in pairs or solitary, 

 unequal in size, pubescence of furcate and simple hairs above, the hairs mostly 

 short stipitate stellate and furcate below, the larger leaves elliptic or ovate, some- 

 times obovate, 9-18 cm. long, 4-8 cm. wide, acuminate, cuneate or rarely sub- 

 cordate at the base, unequal; smaller leaves similar in shape, 1.5-3.5 cm. long, 1.5- 

 2.5 cm. wide, subsessile or petiole to 5 mm. long; inflorescences consisting of 1-3 

 flowers, the pedicels slender, 1-2 cm. long; calyx campanulate, 2-2.5 mm. long, 

 densely pubescent with simple, furcate and dichotomously branched hairs, the 10 

 linear appendages, 8-12 mm. long, subequal or unequal, accrescent in fruit; corolla 

 white, the limb 2.5-3 cm. wide, scarcely lobed, the lobes 1-2 mm. long, externally 

 pilosulous; stamens unequal, the filaments of the short stamens 1-1.5 mm. long, 

 the fifth one 4-4.5 mm. long, the anthers of the short stamens coherent on adjacent 

 sides, 4.5-5 mm. long, the long anther 5-6 mm. long; style about 11 mm. long, ex- 

 ceeding the long stamen by about 2 mm.; berry red or orange-red, globose, 1-1.5 

 cm. in diameter, exceeded by the accrescent calyx appendages. 



Lycianthes purpusii is separated from other species in Guatemala 

 by its reddish-brown or yellowish, furcate and dichotomously 

 branched hairs, the shallowly lobed corolla limb, and the long calyx 

 appendages. 



Lycianthes quichensis (Coult. & Donn.-Sm.) Bitter, Abh. 

 Naturwiss. Vereine Bremen 24: 428. 1920. Solanum quichense Coult. 

 & Donn.-Sm. Bot. Gaz. 37: 422. 1904. L. obliquifolia Standl. Field 

 Mus. Bot. 23: 101. 1940 (type from Volcan de Tacana, Chiapas, 

 Matuda 2938). Chilete; coshel; coxel; flor de rosa; quilete; tomatillo 

 bianco (fide Aguilar) . 



Moist or wet forests or thickets, often in Cupressus or Abies for- 

 ests, 2,200-3,900 m.; Baja Verapaz; Quiche; Huehuetenango; Saca- 

 tepe"quez; Chimaltenango; Solola; Totonicapan; Quezaltenango; San 

 Marcos. Mexico (Chiapas). 




GENTRY & STANDLEY: FLORA OF GUATEMALA 63 



An erect shrub 1-5 m. tall, much branched, the young branches pilose, the 

 hairs multicellular, spreading or incurved; leaves membranaceous, in pairs, un- 

 equal in size, similar in shape or the leaves solitary, pilose, often more densely pubes- 

 cent below, especially on the veins, ciliolate, the larger leaves lanceolate to ovate 

 or elliptic, sometimes broadly ovate, 5-20(-25) cm. long, 2.5-8(-14) cm. wide, 

 acuminate or acute, cuneate at the base, equal or unequal, the petioles 1-6 cm. 

 long; smaller leaves 2.5-9 cm. long, 1.5-4 cm. wide; inflorescences consisting of 

 1-2 flowers, the pedicels pilose, 2-6 cm. long; calyx campanulate, 3.5-4 mm. long, 

 pilose, 10-nerved, the 10 appendages linear-subulate, the alternate ones subequal, 

 the long appendages 3-5 mm. long, the short ones 2-4 mm. long; corolla lavender 

 to purple, sometimes blue, the limb 2.5-3.5 cm. wide, puberulent externally, 

 scarcely lobed, the lobes about 1 mm. long; stamens equal, the filaments 2 mm. 

 long, the anthers 5-6 mm. long; style 1-1.2 cm. long, exceeding stamens by 1-2 

 mm.; berry red or orange-red, subglobose, 8-10 mm. in diameter. 



A shrubby, montane species of Chiapas and western Guatemala 

 with large and brightly colored corollas. 



Lycianthes stephanocalyx (Brandegee) Bitter, Fedde Repert. 

 Sp. Nov. 18: 315. 1922. Solanum stephanocalyx Brandegee, Univ. 

 Calif. Publ. Bot. 6: 374. 1917. S. solitarum Blake, Contr. U. S. 

 Natl. Herb. 24: 21. 1922 (type from Rio Mojanales, Izabal, Blake 

 7489). L. solitaria Standl. J. Wash. Acad. Sci. 17: 15. 1927, syn. nov. 

 S. hondurense Morton in Lundell, Contr. Univ. Mich. Herb. 4: 26. 

 1940 (type from Chalillo Crossing, El Cayo District, British Hon- 

 duras, Lundell 6512). L. hondurensis Standl. & Steyerm. Field Mus. 

 Bot. 23: 18. 1943, syn. nov. 



Wet thickets, 150 m. or below; Izabal. British Honduras. South- 

 ern Mexico. Honduras. 



An erect herb, sometimes suffrutescent, 1-3 m. tall, the young branches 

 sparsely pubescent with small, incurved hairs, glabrate in age; leaves membranace- 

 ous, in pairs, unequal in size, similar or sometimes different in shape or the leaves 

 solitary, pilosulous to nearly glabrous above, pubescent with incurved or spreading 

 hairs below, ciliolate, the larger leaves elliptic to lance-elliptic or ovate to ovate- 

 lanceolate, 5-15 cm. long, 2-5 cm. wide, acuminate, short attenuate or cuneate at 

 the base, the petioles 0.5-2(-5) cm. long; smaller leaves 1.5-5 cm. long, 0.8-2.5 

 cm. wide; inflorescences consisting of a single flower, the pedicels slender, 2.5- 

 5.5 cm. long, glabrous to sparsely pubescent; calyx campanulate, 2.5-3 mm. long, 

 inconspicuously 10-ribbed, sparsely pubescent on the ribs, with 10 alternately 

 subequal appendages, the longest appendages 5 mm. long, the shortest ones 3.5- 

 4.5 mm. long, pubescent with incurved or spreading hairs; corolla white, the limb 

 2-2.5 cm. wide, lobed about one-half way to the base, glabrous; stamens equal, the 

 filaments 1 mm. long, the anthers coherent, 6-6.5 mm. long; style 6-6.5 mm. long; 

 berry red, subglobose, 5-6 mm. in diameter; seeds 1.5-2 mm. long. 



Lycianthes stephanocalyx is a species of the Caribbean lowlands. 




64 FIELDIANA: BOTANY, VOLUME 24 



Lycianthes synanthera (Sendt.) Bitter, Abh. Naturwiss. Ver- 

 eine Bremen 24: 499. 1920. Solanum heteroclitum Sendt. Flora 29: 

 193 [177]. 1846. S. synantherum Sendt. 1. c. 194[1781. Brachistus 

 escuintlensus Coult. Bot. Gaz. 16: 144. 1891 (type from Escuintla, 

 Donn.-Sm. 2267}. S. mitratum Greenm. in Donn.-Sm. Bot. Gaz. 37: 

 211. 1904. L. heteroclita Bitter, I.e. 494, syn. nov. L. heteroclita subsp. 

 coalescens Bitter, I.e. 496 (syntypes from Alta Verapaz) . L. mitrata 

 Bitter, I.e. 500. Bassovia escuintlensis Standl. Contr. U.S. Natl. 

 Herb. 23: 1304. 1924. S. escuintlense A. T. Hunziker, Kurtziana 5: 

 166. 1969. L. escuintlensis D'Arcy, Phytologia 25: 116. 1973, syn. 

 nov. Chilete dulce; chilete. 



Wet thickets or forests, just above sea level to 900 m., rarely 

 higher; Pete"n; Alta Verapaz; Izabal; Quezaltenango ; Huehueten- 

 ango; Retalhuleu; Escuintla; Santa Rosa. Based on material col- 

 lected by Friedrichsthal supposedly in Guatemala, Friedrichsthal 

 1292. British Honduras. Southern Mexico to Costa Rica. Panama. 



Erect herbs or shrubs, 1-3.5 m. tall, sometimes a small tree, the young branch- 

 lets inconspicuously puberulent; leaves solitary or in pairs, then unequal in size, 

 similar in shape, minutely puberulent on the veins above and below, sometimes 

 sparsely so, the larger leaves ovate-lanceolate to broadly ovate or elliptic, some- 

 times narrowly elliptic, 12-30 cm. long, 4.5-15 cm. wide, acuminate, the base 

 attenuate, sometimes unequal, the petioles 1-7 cm. long, minutely puberulent; 

 smaller leaves 3.5-18 cm. long, 2-10 cm. wide; inflorescences consisting of 4-12 

 flowers, the pedicels 1-2 cm. long, inconspicuously puberulent; calyx campanulate, 

 2-3 mm. long, appendages absent, sometimes inconspicuously 10-nerved; corolla 

 white or lavender, the limb 1.5-3 cm. wide, parted to just below the middle, the 

 lobes oblong-lanceolate, 6-9.5 mm. long, inconspicuously puberulent externally, 

 pilose at the apex; stamens equal, the filaments 1.5-2 mm. long, the anthers 

 coherent below or sometimes for their entire length or free, 5-6.5 mm. long; style 

 10-12.5 mm. long, exceeding stamens by 3-5 mm.; berry red or orange, subglobose, 

 glabrous, 7-10 mm. in diameter. 



A more thorough study is required for a better understanding of 

 this complex. Brachistus escuintlense has been treated as a distinct 

 species by D'Arcy (1973). 



Called quilete in Chiapas. 



Lycianthes tricolor (Sesse" & Moc. ex Dun.) Bitter, Abh. Natur- 

 wiss. Vereine Bremen 24: 385. 1920. Solanum tricolor Moc. & 

 Sesse" ex Dun. in Poiret Encycl. Suppl. III. 756. 1813. L. tricolor var. 

 flavidipila Bitter, I.e. 387 (syntypes from Volcan de Agua, Lehmann 

 I486; Santa Maria, Raimann s. n., Guatemala), syn. nov. L. tricolor 

 var. primoaurata Bitter, I.e. 388 (type from Volcan de Fuego, Salvin 

 s. n., Chimaltenango), syn. nov. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 65 



Wet thickets or forests, 1,800-2,500 m., sometimes higher; Qui- 

 che"; Huehuetenango ; Jalapa; Sacatep^quez; Chimaltenango ; Solola; 

 Escuintla; San Marcos. Southern Mexico. El Salvador. 



A vine or shrub, sometimes scandent, 1-3 m. tall, the young branches densely 

 pubescent with incurved or appressed hairs, sometimes the hairs spreading and 

 appressed; leaves in pairs or solitary, similar in shape, unequal in size, sparsely 

 hirsutulous or pilosulous above and below, the hairs often pale yellowish, the larger 

 leaves 6.5-15(-18.5) cm. long, 3-7(-9) cm. wide, acuminate or acute, the base 

 short attenuate, equal or unequal, the petioles 5-25(-40) mm. long; smaller leaves 

 2.5-8(-ll) cm. long, 1.5-3(-5) cm. wide; inflorescences consisting of 3-6(-10) 

 flowers, the pedicels (1.5-)2-3 cm. long; calyx campanulate, 3 mm. long, with 10 

 alternately subequal appendages, the long appendages (1.5-)2.5-4 mm. long, the 

 short ones (l-)2-3.5 mm. long; corolla white or purplish, the limb 1.5-2.5 cm. wide, 

 densely pubescent externally, scarcely lobed, the lobes about 1 mm. long; stamens 

 unequal, the filaments of the short stamens 1.5-2 mm. long, the fifth one 3.5-4 

 mm. long, the anthers 3.5-4 mm. long; style about 8 mm. long, exceeding the 

 longest stamen by about 1 mm. ; berry red, globose, about 1 cm. in diameter, the 

 seeds 3-4 mm. long, slightly notched. 



See comments under Lycianihes arrazolensis. 



Lycianthes sp. 



Known only from the following collection (Huehuetenango, Paso 

 del Boqueron, along Rio Trapichillo, below La Libertad, 1,200-1,300 



m., Steyermark 51177}. 



Shrub, 1-1.5 m. tall, the branches sparsely pubescent with appressed or sub- 

 appressed hairs, glabrescent; leaves mostly in pairs, unequal in size, similar or dif- 

 ferent in shape, sparsely and inconspicuously pubescent above and below, the 

 larger leaves 8-14.5 cm. long, 3-5 cm. wide, acuminate, long-attenuate at the base, 

 the petioles 1-1.5 cm. long; smaller leaves 4.5-7 cm. long, 2-4.5 cm. wide; inflores- 

 cences consisting of 2 flowers, the pedicels 1.5-2 cm. long, sparsely pubescent with 

 appressed hairs; immature fruiting calyx 3 mm. long, pubescent with appressed 

 hairs, 10-nerved, the 10 appendages alternately subequal, the long ones 9.5-11 mm. 

 long, the short ones 5-6.5 mm. long; berry immature, apparently globose. 



This specimen seems to represent an undescribed species but we 

 hesitate to describe it on inadequate material. 



LYCOPERSICON Miller 



Reference: C. H. Muller, A revision of the genus Lycopersicon, 

 U. S. Dept. Agr. Misc. Publ. No. 382. 29 pp. 1940. 



Herbs; leaves alternate, odd-pinnate or sometimes bipinnate, petiolate, inter- 

 stitial leaflets present, the leaflets shallowly lobed, serrate, crenate, or rarely entire, 

 petiolulate; inflorescences cymose or racemose, lateral and opposite the leaves or 

 internodal; pedicels articulate near or above the middle; calyx campanulate, 




66 FIELDIANA: BOTANY, VOLUME 24 



5-parted nearly to the base, the lobes lanceolate, often accrescent in fruit; corolla 

 subrotate, yellow, the limb shallowly lobed or parted (in ours); stamens 5, rarely 

 more, subsessile or with short filaments; anthers coherent, forming a tube around 

 the style, the thecae elongate, apically appendaged with sterile tips, abaxially 

 dehiscent by longitudinal slits; ovary bilocular, the ovules numerous, the disk 

 inconspicuous; style elongate, included in the anther tube or exserted; stigma 

 capitate; fruit a berry, globose, fleshy and juicy, usually red or yellow; seeds 

 numerous (in ours), compressed, each enclosed in a mucilaginous sheath, nearly 

 glabrous or pubescent; embryo curved. 



A genus of about eight species; native of western South America. 

 It is of great economic importance because it contains the common 

 tomato, Lycopersicon esculentum. Only the following species is 

 known in North America. 



Lycopersicon esculentum Miller, Gard. Diet. ed. 8. Lycopersi- 

 con No. 2, 1868. Solanum lycopersicum L. Sp. PI. 185. 1753. Tomate; 

 Pix (Quiche") ; Pixb, Pixp (Quiche*, Poconchi) ; Ixpix (Chuje) ; Xcoya 

 (Mame) ; Pai'c (Ixil) ; Tomate de rinon; Icpic. 



A native of South America but now widely cultivated in nearly 

 all parts of the world. It is grown in almost all inhabited parts of 

 Guatemala except at higher elevations. 



The tomato is one of the most important vegetables in Guate- 

 mala. They are added as flavoring to almost every kind of dish, 

 and also cooked alone, or eaten in salads. At a pension in Quezal- 

 tenango a favorite dessert is made by boiling whole well-shaped 

 tomatoes in thin syrup, in such a manner that each fruit preserves 

 its shape. 



Lycopersicon esculentum var. cerasiforme (Dunal) A. Gray, 

 Syn. Fl. 2, pt.l: 226. 1886. L. cerasiforme Dunal, Hist. Solan. 113. 

 1813. Tomate. 



Dry or damp thickets, or in fields or waste ground, especially 

 about dwellings, just above sea level to 2,500 m.; Zacapa; Chiqui- 

 mula; Jalapa; El Progreso; Guatemala; Sacatepe*quez ; Quezalten- 

 ango; Jutiapa; Santa Rosa; Escuintla; Retalhuleu, also probably in 

 most of the other departments. Mexico to Costa Rica. West In- 

 dies. South America. 



An herb, the stems somewhat succulent, puberulent to pilose with jointed 

 hairs and glandular-puberulent; leaves large, mostly 10-25 cm. long, petiolate, 

 inconspicuously to densely puberulent; interstitial leaflets present; leaflets mostly 

 9, ovate to lanceolate, mostly 2-8 cm. long, 0.5-3 cm. wide, repand to parted, apex 

 acute, petiolulate; inflorescence racemose; peduncle 1-3.5 cm. long, pilose with 




FIG. 13. Lycopersicon esculentum var. cerasiforme. A, habit, X ^; B, fruit, 

 X 1 and X 1M; G, flower, X 3}/; D, corolla showing connivent anthers, X 3>; 

 E, stamen tube dissected showing sterile tips of the anthers, X 3>; F, calyx and 

 pistil, X 3>6. 




68 FIELDIANA: BOTANY, VOLUME 24 



jointed hairs and glandular-puberulent; pedicels 5-15 mm. long, pilose with jointed 

 hairs and glandular-puberulent; calyx deeply parted, the lobes 3.5-5 mm., lance- 

 olate, accrescent in fruit, acute; corolla parted to below the middle, the lobes 5-6 

 mm. long, narrowly lanceolate, reflexed; anthers 5-6 mm. long; style included or 

 exserted beyond the anther tube; fruit a berry, globose, red or yellow, mostly 1.5-2 

 cm. in diameter. 



Variety cerasiforme differs from variety esculentum by its small 

 cherrylike fruits, mostly 1.5-2 cm. in diameter. The fruits of this 

 small tomato are abundant in all markets. They are utilized for 

 flavoring food. 



MARGARANTHUS Schlechtendal 



Herbs; leaves alternate, solitary or in pairs, entire, petiolate; inflorescences 

 axillary, consisting of a single flower, the flowers pedicellate; calyx campanulate, 

 5-lobed, accrescent and inflated in fruit; corolla small, pentangular-urceolate, 

 purplish or greenish-yellow, minutely 5-dentate; stamens 5; anthers longitudinally 

 dehiscent; ovary bilocular, on a short gynophore, the ovules numerous, the disk 

 inconspicuous; style filiform, included; stigma discoid; fruit a berry, entirely en- 

 closed in the inflated calyx; seeds numerous, compressed; embryo strongly inwardly 

 curved. 



A genus consisting of only one species, ranging from the south- 

 western United States to Mexico, Honduras, and West Indies. 



Margaranthus solanaceus Schlecht. Ind. Sem. Hort. Halensis 

 8. 1838. 



Moist thickets and weedy fields, 800-1,000 m.; Southwestern 

 United States; Mexico; (Dept. Morazan) Honduras; West Indies. 



Annual herbs, erect or decumbent, the stems 1.5-4 dm. long, sometimes longer; 

 leaves ovate to lanceolate, 2-6 cm. long, 0.5-1.5 cm. wide, acute or acuminate, the 

 base unequal or equal, cuneate, generally glabrous, ciliate, the petioles 5-20 mm. 

 long; flowers with recurved pedicels, 2-3 mm. long; calyx 1.5-2 mm. long, ap- 

 pressed-pubescent, the lobes about 0.5 mm. long, acute; corolla 3.5-4 mm. long, 

 internally pubescent with multicellular hairs, the lobes minute; stamens included; 

 filaments about 1 mm. long; anthers blue, 1-1.2 mm. long; berry ovoid, 6-10 mm. 

 in diameter, included in the accrescent calyx, the calyx 1-1.5 cm. long; seeds about 

 2 mm. long, suborbicular, reticulate-foveate. 



Margaranthus is closely related to Physalis. Fruiting specimens 

 of Margaranthus cannot be distinguished from some of the small fruit- 

 ing species of Physalis, however, the pentangular-urceolate corolla 

 of Margaranthus is quite different from those of Physalis. 



MARKEA L. C. Richard 

 Merinthopodium Donn.-Sm. Bot. Gaz. 33: 11. 1897. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 69 



Reference: J. Cuatrecasas, Notes on American Solanaceae. Fedde 

 Repert. Spec. Nov. 61: 74-86. 1958. 



Shrubs, mostly epiphytic, and often scandent, glabrous or pubescent, the hairs 

 simple; leaves alternate, sometimes in pairs and entire, often coriaceous, petiolate; 

 inflorescences cymose or sometimes the flowers solitary, often pendent on long cord- 

 like peduncles, the flowers large; calyx campanulate, 5-parted; corolla funnel- 

 form, white, yellow, green, or orange, sometimes red, the limb 5-lobed, the lobes 

 valvate or imbricate in bud, the tube ampliate above; stamens 5, inserted below the 

 middle of the corolla tube, exserted or included; filaments filiform, pubescent be- 

 low; anthers oblong or linear, longitudinally dehiscent; ovary bilocular, the 

 ovules numerous, the disk conspicuous or inconspicuous; style elongate, dilated 

 above, included or exserted; stigma bilobed; fruit a berry, leathery, often almost 

 dry; seeds numerous; embryo somewhat curved or almost straight. 



About 20 species in the American tropics. A few more species 

 are known from southern Central America. 



Corolla 10 cm. long M. campanulata. 



Corolla 7.5 cm. long or less. 



Inflorescence consisting of a cluster of flowers produced at the end of long 



pendent, cord-like peduncles. 

 Calyx 3-3.5 cm. long, membranaceous, the lobes lanceolate to narrowly 



lanceolate, the apex narrowly long attenuate M. internexa. 



Calyx 1.2-1.6 cm. long, subcoriaceous, the lobes oblong lanceolate, the apex 



acute or short-acuminate M. neurantha. 



Inflorescence cymose or rarely the flowers solitary, not produced at the end of 



long-pendent, cord-like peduncles. 

 Calyx 3.5-4 cm. long, 5-angled; pedicels 4-7.5 cm. long; branches hirsute, 



tuberculate in age M. uniflora. 



Calyx 1-1.5 cm. long, not angled; pedicels 0.5-1 cm. long; branches glabrous. 



M. megalandra. 



Markea campanulata (Donn.-Sm.) Lundell, Phytologia 1: 339. 

 1939. Merinthopodium campanulatum Donn.-Sm. Bot. Gaz. 47: 257. 

 1909. 



Known only from the type collected near Coban, Alta Verapaz, 

 1,600 m., Tuerckheim II 2391. Endemic. 



An epiphytic shrub (?), the leaves unknown, but said to be oval, coriaceous 

 and acute; pedicels 8 cm. long, sparsely and inconspicuously pubescent; calyx 5.5 

 cm. long, coriaceous, 5-angled, parted to near the base, the lobes lanceolate, apex 

 short acuminate; corolla greenish, 10 cm. long, glabrous externally, the tube 2.5 

 cm. long, the campanulate portion 5.2 cm. long, the lobes lanceolate, 2.2 cm. long; 

 stamens short-exserted, inserted about 2.5 cm. above the base of the corolla; fila- 

 ments 4.5 cm. long, densely villous at their bases; anthers 1.7 cm. long; style 8 

 cm. long; fruit unknown. 



This is a poorly known species, described from incomplete ma- 

 terial. The type collection consists of a single flower. 




70 FIELDIANA: BOTANY, VOLUME 24 



Markea internexa (Blake) Lundell, Phytologia 1: 340. 1939. 

 Merinthopodium internexum Blake, Contr. Gray Herb. 52: 85. 1917. 



Wet forests, 300-400 m.; Alta Verapaz (type from Cubilquitz, 

 Tuerckheim 8435). Endemic. Known only from the type area. 



An epiphytic shrub, the young branches densely or sparsely hispidulous, the 

 bases of the hairs much swollen, becoming tuberculate in age; leaves subcoriace- 

 ous. elliptic to obovate, 9.5-18 cm. long, 4.5-6.5 cm. wide, glabrous above and be- 

 low, apex abruptly acuminate, obtuse to acute at the base, the petioles 5-15 mm. 

 long, glabrous or sparsely pubescent; flowers few, produced at the end of a long 

 cord-like, pendent peduncle; peduncle 22.5-32 cm. long, densely hispidulous, the 

 bases of the hairs much swollen, becoming tuberculate in age; pedicels slender 

 below, thickened above, 3-4 cm. long, sparsely and inconspicuously puberulent; 

 calyx 3-3.5 cm. long, membranaceous, parted to near the base, the lobes lanceolate 

 to narrowly lanceolate, apex narrowly attenuate and puberulent; corolla probably 

 greenish, 5.5-7 cm. long, 15-nerved, the nerves inconspicuously puberulent ex- 

 ternally, the tube 2-2.5 cm. long, the campanulate portion 2.5-3.5 cm. long, the 

 lobes triangular, 1 cm. long, acute; stamens short-exserted, attached about 1 cm. 

 above the base of the tube; filaments 3.5 cm. long, villosulous at their bases, the 

 staminal traces villosulous; anthers 1 cm. long; style equalling the stamens; fruit 

 unknown. 



Markea megalandra (Dunal) D'Arcy, Phytologia 25: 116. 1973. 

 Metternichia megalandra Dunal in DC. Prodr. 13, pt. 1: 595. 1852. 

 Markea leucantha Donn.-Sm. Bot. Gaz. 31: 116. 1901. Merintho- 

 podium leucanthum Blake, Contr. Gray Herb. 52: 77. 1917, syn. nov. 



In moist or wet forests, 1,360 m.; Alta Verapaz. Costa Rica. 

 Panama. Northern South America. 



An epiphytic shrub, the branches stout, glabrous; leaves coriaceous, often 

 lustrous, elliptic to obovate, mostly 6.5-15 cm. long, 2.5-6.5 cm. wide, incon- 

 spicuously glandular-punctate above, inconspicuously glandular-punctate and 

 punctate below, apex acute to obtuse or rounded, sometimes abruptly short- 

 acuminate, acute or obtuse to nearly rounded at the base, the petioles stout, 5-10 

 (-15) mm. long, glabrous; inflorescences consisting of flowers clustered at the ends 

 of leafy branches, the flowers often very numerous; pedicels stout, 5-10 mm. long, 

 pilosulous, sometimes densely so; calyx coriaceous, 1-1.5 cm. long, parted to just 

 below the middle or to near the base, the lobes oblong-lanceolate or lanceolate, 

 6-10 mm. long, pilosulous, the hairs sometimes sparse, short-acuminate; corolla 

 white or yellow, 5-6.5 cm. long, 5-nerved, glabrous externally, the tube about 7 

 mm. long, the campanulate portion 3-4.5 cm. long, the lobes rounded, 1.2-1.5 

 cm. long; stamens exserted, inserted about 6 mm. above the base of the corolla 

 tube; filaments 3.5-4 cm. long, densely villous at their bases; anthers 5-6 mm. long; 

 style 4-5 cm. long; fruit leathery, 1-1.5 cm. long; seeds numerous, 2.5-3 mm. long. 



Markea megalandra is evidently rare in Guatemala, but abundant 

 in the mountains of Costa Rica. It is a very beautiful plant when in 

 full flower. The flowers are strongly fragrant. 




OTAXICA1. fiAXKTTE. -VA7//. 



FIG. 14. Markea neurantha. Plate I from the Botanical Gazette for the year 

 1897, used by permission of the University of Chicago. Magnifications may be 

 calculated from the description. 



71 




72 FIELDIANA: BOTANY, VOLUME 24 



Markea neurantha Hemsley, Biol. Cent.-Am. Bot. 2: 429. 1882. 

 Merinthopodium neumnthum Donn.-Sm. Bot. Gaz. 23: 12, 1. 1. 1897. 

 M. leptesthemum Blake, Contr. Gray Herb. 52: 86. 1917 (type from 

 Toledo, British Honduras, Peck 508). 



Wet or moist forests, 65 m. or less, Izabal, Steyermark 39458. 

 British Honduras. Honduras. Costa Rica. 



An epiphytic shrub, the branches scarred by the prominent leaf scars, his- 

 pidulous, the bases of the hairs much swollen, becoming tuberuclate in age or 

 sometimes glabrous; leaves subcoriaceous, elliptic to obovate, 10-25 cm. long, 

 3-8 cm. wide, glabrous above and below, apex abruptly long acuminate, acute to 

 short attenuate at the base, the petioles 1-4 cm. long, glabrous; flowers few, pro- 

 duced at the end of a long cord-like, pendent peduncle; peduncle 10-70 cm. long, 

 rarely shorter, glabrous (in ours) or hispidulous, the bases of the hairs much swol- 

 len, becoming tuberculate in age, the rachis scarred by the prominent pedicel 

 scars, sometimes branched, hispidulous; pedicels slender below, thickened above, 

 1-3.5 cm. long, puberulent or nearly glabrous; calyx 1.2-2 cm. long, subcoria- 

 ceous, puberulent to nearly glabrous, parted nearly to the base, the lobes oblong, 

 1-1.4 cm. long, apex acute or short acuminate; corolla green or yellowish-green, 

 3.5-5 cm. long, 15-nerved, inconspicuously puberulent and glandular externally, 

 the tube 2-3.5 cm. long, the lobes oblong-lanceolate, 0.8-1.5 cm. long, acute or ob- 

 tuse, the sinuses of the corolla lobes bilobulate or lobulate; stamens short-exserted, 

 attached about 3 mm. above the base of the tube; filaments 2-3.5 cm. long, densely 

 villous at the base and for about one-third their length; anthers 7.5-8.5(-12) mm. 

 long; style 3-3.5 cm. long; fruit 1.5-1.8 cm. long, ovoid; seeds 2 mm. long. 



Markea neurantha has been collected only a few times in Guate- 

 mala and British Honduras but it is very common in Costa Rica. 

 Collectors have often called this epiphytic shrub a large tree because 

 they failed to note the true source of the long, pendent, cord-like 

 peduncles. 



Markea uniflora Lundell, Phytologia 1: 340. 1939. 



In wet forests, 2,000-3,000 m.; San Marcos, southeast slopes of 

 Volcan de Tacana, Steyermark 36465. Mexico (Chiapas), the type 

 from Ciquihuite, Volcan de Tacana, 2,800 m., Matuda 2816. 



An epipihytic scandent shrub, the young branches hirsute, the hairs multi- 

 cellular and with the basal cell swollen, the branches becoming tuberculate in age; 

 leaves coriaceous, elliptic or obovate-elliptic, 4-12 cm. long, 2-6 cm. wide, incon- 

 spicuously glandular, glabrate in age above, hirsute mostly on the veins and in- 

 conspicuously punctate and glandular-punctate below, apex acute to obtuse or 

 nearly rounded, obtuse or short attenuate at the base, the petioles 5-15 mm. long, 

 sparsely hirsute; inflorescences consisting of a solitary flower or in open few-flow- 

 ered cymes; pedicels 4-7.5 cm. long, sparsely hirsute, slender at the base, thickened 

 above; calyx 3.5-4 cm. long, coriaceous, 5-angled, parted to below the middle, the 

 lobes lanceolate, 2.5-3 cm. long, sparsely and very inconspicuously glandular, apex 




GENTRY & STANDLEY: FLORA OF GUATEMALA 73 



acute or very short acuminate; corolla pale green, 7.5 cm. long, 5-nerved, the 

 nerves pilose externally, the tube 2 cm. long, the campanulate portion 3.5 cm. long, 

 the lobes ovate, 2 cm. long, acute or rounded; stamens short-exserted, inserted 2 

 cm. above the base of the corolla; filaments 4 cm. long, densely villous at their 

 bases; anthers about 1.5 cm. long; style about 6 cm. long; ovary glabrous, said to 

 be trilocular, multiovulate; fruit unknown. 



MELANANTHUS Walpers 



Microschwenckia Hemsley, Biol. Cent.-Am. Bot. 2: 438, t. 57 A. 



1882. 



Herbs; leaves alternate, sometimes fasciculate, entire, sessile; inflorescence 

 racemose or spike-like, the flowers small and inconspicuous; calyx tubular, 5-lobed; 

 corolla tubular, purplish-black, the lobes 3-lobulate with the median lobule linear, 

 the 2 shorter lateral lobules only slightly fused with the adjoining lateral lobes of 

 each adjacent corolla lobe, thus the corolla appearing to be 1 5-lobed, but actually 

 5-lobed; stamens 4, didynamous, the 2 lower ones sometimes sterile, included; 

 anthers free, longitudinally dehiscent; filaments filiform, inserted below the middle 

 of the corolla tube; ovary unilocular, uniovulate, the disk cupular; style smooth 

 and slender, included; stigma capitate; capsule septicidal and tardily dehiscent, 

 ovate, oblique, glabrous and tuberculate; 1-seeded; embryo straight. 



A genus of about five species, ranging from southern Mexico to 

 Honduras, Cuba, and Brazil. 



Melananthus guatemalensis (Benth.) Solereder, Ber. Deutsch. 

 Bot. Ges. 9: 84. 1891. Microschwenckia guatemalensis Benth. ex 

 Hemsl. Biol. Cent.-Am. Bot. 2: 438, t. 57 A. 1882. Schwenckia oxy- 

 carpa Robinson, Proc. Am. Acad. 45: 400. 1910 (type from near the 

 Sibun River, British Honduras, Peck 417 A), syn. nov. 



Grassy plains or fields, sometimes in oak or pine forests, 800- 

 1,500 m.; Jalapa; Jutiapa (type from Llanos de Jutiapa, Bernoulli 

 716); Huehuetenango. British Honduras. Southern Mexico. Hon- 

 duras. Cuba and Brazil. 



Slender annual plants, 1-3.5 dm. tall, stem pubescent with inwardly curved 

 hairs; leaves linear or narrowly elliptic, glabrous, only the midvein conspicuous, 

 5-15 mm. long, 2 mm. wide or less, acute; inflorescence racemose or spike-like, the 

 flowers subsessile; calyx 1-1.5 mm. long, puberulent, the lobes less than 0.5 mm. 

 long; corolla purplish-black, 2.5-3 mm. long, the tube slender, 0.5 mm. wide; 

 anthers 2-celled; filaments 0.7-1.2 mm. long; style 1.5 mm. long; capsule hard, 

 2-2.5 mm. long, acute. 



Melananthus is most likely to be confused with Schwenckia when 

 in flower, however, they can be readily distinguished in fruit. In 

 Melananthus the capsule is ovoid, oblique and acute, tuberculate, 

 and with 1 seed whereas in Schwenckia the capsule is globose, equal 




FIG. 15. Melananthus guatemalensis. A, habit, X J^; B, flower, X 10; G, 

 corolla lobes, detail, X 20; D, mature fruit, X 10. Nicandra physalodes. E, 

 habit, X 1 A; F, fruiting calyx, X H. 



74 




GENTRY & STANDLEY: FLORA OF GUATEMALA 75 



and obtuse, smooth and with numerous seeds. Melananthus ap- 

 pears to be anomolous in the Solanaceae but see Solereder in Ber. 

 Deutsch. Bot. Ges. 9: 65-84, t. IS. 1891. 



NICANDRA Adanson, nom. cons. 



Herbs, generally glabrous throughout; leaves alternate, coarsely sinuate- 

 dentate or shallowly lobed, petiolate; inflorescences axillary, consisting of a single 

 flower, the flowers pedicellate; calyx 5-parted, angulate, accrescent and reticulate 

 in fruit, the lobes cordate-sagittate at their bases; corolla broadly campanulate, 

 pale blue, the limb nearly entire to shallowly lobed, plicate in bud; stamens 5, 

 included; filaments inserted near the base of the corolla tube, densely pubescent 

 on the dilated bases; anthers ovate-oblong, longitudinally dehiscent; ovary on a 

 short gynophore, 3-5-locular, the ovules numerous, the disk entire; style thick and 

 elongate, included; stigma subglobose with 3-5 prominent stigmatic areas; fruit a 

 berry, 3-5-locular, enclosed in the accrescent calyx; seeds numerous, compressed; 

 embryo strongly curved. 



A monotypic genus, probably a native of Peru now widely dis- 

 persed in temperate and tropical areas of both hemispheres. 



Nicandra physalodes (L.) Gaertn. Fruct. Semin. PI. 2: 237. 

 1791. Atropa physalodes L. Sp. PI. 181. 1753. Miltomate de culebra; 

 miltomate de monte; tolcic (Quecchi) ; tomate de culebra; tomatillo. 



Moist fields or thickets or in waste ground, often in cafetales, 900- 

 1,800 m., occasionally higher; Huehuetenango; Chimaltenango; 

 Sacatepe"quez; Quiche", Alta Verapaz; Santa Rosa; Jalapa; Chiqui- 

 mula. Eastern United States; Mexico to Costa Rica; West Indies; 

 northern South America. Probably a native of Peru. 



Plants to 1 m. tall, occasionally more, glabrous throughout or sometimes 

 sparsely pilose; leaves usually ciliate, ovate to broadly ovate, mostly 7-25 cm. long, 

 3-10 cm. wide, acute or acuminate, attenuate at the base, the petiole 1-5 cm. long, 

 sometimes narrowly winged; flowers solitary, cernuous, the pedicels 1-2 cm. long; 

 calyx 1-2.5 cm. long, parted to near the base, the lobes ovate-cordate, acuminate 

 at the apex, sagittate at the base; corolla blue or pale blue, 1.5-3 cm. long, the 

 limb 2.5-3.5 cm. broad, nearly entire to shallowly lobed; filaments 5-6 mm. long; 

 anthers 3.5-4 mm. long, apiculate; style 6-7 mm. long; berry 1-2 cm. in diameter, 

 globose; seeds 1.5-1.8 mm. long, compressed, suborbicular, reticulate-foveate. 



Nicandra is very similar to Physalis in general appearance. How- 

 ever, it can be distinguished by the blue corollas and sagittate bases 

 of the calyx lobes. 



Called miltomate and huevo de tortuga in El Salvado. 




FIG. 16. Nicotiana glauca. A, habit, X M; B, flower, X 

 part, X 1 1 A; D, pistil, X \ 1 A; E, cauline leaf, X 1. 



; C, corolla, lower 



76 




GENTRY & STANDLEY: FLORA OF GUATEMALA 77 



NICOTIANA L. 



Reference: T. H. Goodspeed, The Genus Nicotiana, Chron. Bot. 

 16: 1-535. 1954. 



Herbs, rarely shrubs, or small trees, often viscid-glandular or sometimes 

 glabrous; leaves alternate, entire, sessile or petiolate; inflorescence a panicle or 

 false raceme, the flowers pedicellate; calyx tubular, 5-lobed, usually accrescent in 

 fruit; corolla tubular or salverform, white to red, or yellow, the limb 5-lobed, the 

 lobes convolute-plicate in bud; stamens 5, variously inserted on the corolla tube; 

 anthers longitudinally dehiscent; ovary bilocular, the ovules numerous, the disk 

 prominent; style filiform and smooth; stigma capitate, slightly grooved; fruit a 

 capsule, septicidal, the valves bifid, smooth and glabrous; seeds numerous; embryo 

 straight. 



Species about 65, chiefly in the Americas, a few in Australia and 

 the Pacific Islands. Only the following taxa are known in Central 

 America, but several others are native in Mexico and the south- 

 western United States. 



Shrubs or small trees; glabrous throughout; leaves long petiolate N. glauca. 



Herbs; pubescent with usually viscid-glandular hairs; middle and upper leaves 

 sessile. 



Corolla tube 3-4 mm. wide above; stamens inserted near the mouth of the corolla 

 tube, the filaments short, 1 mm. long or less; leaves clasping to auriculate, 

 not decurrent N. plumbaginifolia. 



Corolla tube 7-12 mm. wide above; stamens inserted below the middle of the 

 corolla tube, the filaments long and slender; lower leaves decurrent. 



N. tabacum. 



Nicotiana glauca Graham, Edinburgh New Philos. Journ. 5: 

 175. 1828. Tabacon. 



A native of South America which has become naturalized in Mex- 

 ico. Known only from cultivated plants in Guatemala but to be 

 expected as an escape. El Salvador; Honduras. 



Shrub or small tree, 3-10 m. tall, glabrous; leaves ovate to elliptic or oblong- 

 elliptic, glaucous, 4-12 (-38) cm. long, 2-8(-22) cm. wide, apex acute, the base 

 cuneate, sometimes unequal; petioles 3-6(-10) cm. long; inflorescence paniculate, 

 flowers lax; pedicels 3-10 mm. long; calyx 1-1.5 cm. long, the lobes 2-3 mm. long, 

 acute and much shorter than the tube; corolla yellowish, tubular, 3-3.5 cm. long, 

 villosulous externally, the limb narrow, nearly circular; stamens subequal, extend- 

 ing almost to the mouth of the tube; filaments glabrous, about 2.5 cm. long, at- 

 tached below the middle of the tube, geniculate just above the point of attach- 

 ment; capsule 7-10 mm. long, included or slightly exceeding the calyx; seed some- 

 what angular, laterally compressed, wavy reticulate, about 0.5 mm. long. 







Nicotiana plumbaginifolia Viviani, Elench. PI. Hort. Dinegro, 

 26, L 5. 1802. 




78 FIELDIANA: BOTANY, VOLUME 24 



Wet sand along rivers or in waste places, sea level to 1,500 m.; 

 Izabal; Jalapa; Guatemala; Sacatepe"quez; Quiche"; Escuintla. Uni- 

 ted States (Florida Keys). Mexico. West Indies. South America. 



Annual, 0.3-1 m. tall, the stem slender, sparsely viscid-pilose, becoming 

 rough in age; leaves sparsely viscid-pilose, the basal leaves obovate or oblance- 

 olate, 1-2 dm. long, the lower ones rotund-ovate, sessile, the margin undulate, apex 

 abruptly acuminate, usually twisted, the base clasping to clasping-auriculate, the 

 upper cauline leaves reduced, lanceolate to linear-lanceolate, margin strongly 

 undulate, auriculate; inflorescence a false raceme, occasionally somewhat pani- 

 culate; pedicels 3-7 mm. long; calyx 8-12 mm. long, 10-ribbed, lobed to the mid- 

 dle or more deeply so, the lobes unequal, 4-8 mm. long, linear-subulate; corolla 

 vespertine, salverform, the limb lavender or pink, deeply lobed, the lobes ovate, 

 acute, the tube pale green or purplish, long and slender, 2.5-3.5 cm. long, 1.5- 

 2 mm. wide, ampliate above to 3-4 mm. wide; stamens unequal, attached just be- 

 low the mouth of the tube, four nearly sessile, the fifth stamen about 3 mm. lower 

 on the tube; filaments glabrous, about 1 mm. long or less, shorter than the an- 

 thers; capsule 8-11 mm. long, narrowly ovoid, about the same length as the 

 calyx; seeds round-elliptic, wavy-reticulate, about 0.5 mm. long. 



Nicotiana tabacum L. Sp. PI. 180. 1753. Tabaco; tabaco bobo; 

 cutz, cuutz (Maya) ; iic (Alta Verapaz) ; jic (Quiche") ; mai, met (Quec- 

 chi, Tzental) ; si'cal (Ixil) ; si'ch (Aguacateca) ; sii'c (Mame, Chuje) . 



Cultivated in almost all regions of Guatemala, occasionally 

 escaping cultivation. Believed to be a native of western South 

 America, but introduced into Central America and Mexico in pre- 

 Columbian times. 



Annual 1-3 m. tall, viscid throughout, the stems thick, branches few, leaves 

 ovate to elliptic or obovate, the larger ones at least 5 dm. long, decurrent, sessile 

 and amplexicaul, apex acuminate or acute; panicle with several branches; pedicels 

 5-15 mm. long; calyx 12-18 mm. long, the lobes 4-8 mm. long, narrowly acute, un- 

 equal, shorter than the calyx tube; corolla salverform, the limb white, pink or red, 

 lobed or pentagonal, the tube pale greenish-cream, pink or red, 3.5-4.5 cm. long, 

 3-5 mm. wide below, ampliate above to 7-12 mm. wide; stamens unequal, attached 

 below the middle of the tube, the anthers of the two longer pairs near the mouth 

 of the corolla or slightly exserted, the fifth stamen shorter than either pair; fila- 

 ments 2.5-3.5 cm. long, greatly exceeding the anthers in length, pubescent below; 

 capsule 1.5-2 cm. long, narrowly elliptic or ovoid, surpassing or included in the 

 calyx; seeds spherical or broadly elliptic, 0.5 mm. long, wavy-reticulate. 



PETUNIA Jussieu 



Reference: R. E. Fries. 1911. Die Arten der Gattung Petunia. 

 Kongl. Svenska Vetenskapsakad. Handl. 46. 



Herbs, viscid-pubescent, the stems branched and often decumbent; leaves al- 

 ternate or in pairs, entire, sessile or subsessile; inflorescences axillary, consisting 




GENTRY & STANDLEY: FLORA OF GUATEMALA 79 



of a single flower, the flowers large and showy, long pedicellate; calyx campanulate, 

 deeply 5-lobed; corolla salverform or funnelform, slightly zygomorphic, variously 

 colored, often variegated and sometimes double, 5-lobed, the lobes plicate in bud, 

 the tube ampliate above; stamens 5, 4 didynamous, the fifth one shorter than either 

 pair; filaments filiform, subequally inserted below the middle of the corolla tube, 

 glabrous; anthers longitudinally dehiscent; ovary bilocular, the ovules numerous, 

 the disk prominent, carnose; style filiform and smooth; stigma capitate and slightly 

 grooved ; fruit a capsule, septicidal, the valves bifid, equalling or surpassed by the 

 calyx; seeds numerous; embryo curved or almost straight. 



About 30 species, mostly in South America. Only one species 

 (P. parviflora) is native to North America and apparently does not 

 reach Central America. 



Petunia hybrida Hort. Petunia. 



Commonly cultivated in almost all parts of Guatemala. Oc- 

 casionally escaping from cultivation and becoming established in 

 some areas. This is the showy cultivated Petunia. It is a putative 

 hybrid derived from P. axillaris (Lam.) B.S.P. x P. violacea Lindl. 



Annual herbs; leaves ovate to elliptic, 3-6 cm. long, 1.5-4 cm. wide, acute, the 

 base cuneate; pedicels 2-6 cm. long; calyx 1.2-1.5 cm. long, the lobes 10-13 mm. 

 long, obtuse and unequal, more than 2 times as long as the calyx tube; corollas 

 variously colored, often variegated, sometimes double, the limb 3.5-5 cm. wide, the 

 lobes obtuse, the tube 2.5-4 cm. long; stamens with two long pairs, the fifth one 

 shorter than either pair; filaments somewhat geniculate at the base; style 2.5-3.5 

 cm. long; capsule 1-1.5 cm. long, glabrous, the seeds 0.6 mm. long or less, subglo- 

 bose or ovoid, reticulate-foveate. 



Dr. Paul C. Standley, who originated this flora, professed to 

 despise Petunias and often tried to interest his listeners in a society 

 for the extermination of Petunias, all Petunias. 



PHYSALIS L. 



References: U. T. Waterfall, A taxonomic study of the genus Phy- 

 salis in North America North of Mexico. Rhodora 60: 107-114; 

 128-142; 152-173. 1958; Physalis in Mexico, Central America, and 

 the West Indies, 1. c. 69: 82-120; 203-239; 319-329. 1967. 



Herbs or rarely shrubs; leaves alternate, sometimes in pairs, entire to sinuately 

 lobed, petiolate; inflorescences axillary, consisting of a single flower or sometimes 

 with few to several flowers in axillary fascicles, the flowers pedicellate; calyx cam- 

 panulate or sometimes tubular-campanulate, 5-lobed, much accrescent in fruit; 

 corolla subrotate to campanulate-rotate, or oampanulate, rarely tubular-campanu- 

 late, mostly yellowish, often with 5 dark, contrasting spots near the mouth of the 

 corolla tube, the mouth densely pubescent or sometimes glabrous or somewhat 

 glandular; the limb mostly shallowly 5-parted, the lobes mostly plicate in bud; sta- 




80 FIELDIANA: BOTANY, VOLUME 24 



mens 5; filaments filiform or broad; anthers oblong to linear-oblong, or lanceolate, 

 longitudinally dehiscent, yellow to blue, violet or greenish-blue, or sometimes so 

 tinged; ovary bilocular, the ovules few to many; style filiform, exserted or included; 

 stigma capitate to nearly truncate; fruit a berry, included in the inflated calyx, 

 rarely tightly invested by the calyx, sometimes on a short gynophore, the fruiting 

 calyx 5- or 10-angled or -ribbed or teretish, much enlarged and inflated around the 

 fruit, often invaginated basally; seeds few to many, laterally compressed; embryo 

 curved. 



About 100 species mostly in the New World tropics, especially 

 in Mexico and Guatemala, also extending into temperate regions, a 

 few in the Old World. A few other species are known from other 

 parts of Central America. 



The treatment presented here differs in some respects from that 

 proposed by Waterfall in 1967. 



Physalis is a difficult genus that requires further study. Physalis 

 melanocystis (Robinson) Bitter var. cernua (Donn.-Sm.) Waterfall 

 and P. schraderiana Bernh. = Athenaea viscosa (Schrader) Fern, 

 have been placed in Athenaea. See comments under Athenaea and 

 Margaranthus. 



Corolla limb 5-lobed, the limb lobed one-half way to the base or more; flowers 



(1-) 2-6 in the leaf axils. 

 Plants shrubs; leaves with a few furcate and branched hairs below; Caribbean 



lowlands, below 500 m P. porphyrophysa. 



Plants herbs, sometimes suffrutescent; leaves with simple hairs; montane forest, 

 1,300-2,850 m. 



Calyx lobes prominent, the lobes 2.5-10 mm. long P. amphitricha. 



Calyx nearly truncate to shallowly and broadly lobed, the lobes 1 mm. long 



or less P. calidaria. 



Corolla limb nearly entire to 5-angled, the limb lobed less than one-half way to the 



base; flowers solitary (in ours). 

 Corolla tubular-campanulate, the flowers 18-19 mm. long; filaments 8-9 mm. 



long P. campanula. 



Corolla subrotate to campanulate, the flowers 2-15 mm. long; filaments 0.5-5 



mm. long. 



Fruiting calyces strongly 5-angled or 5-ribbed. 

 Fruiting calyces glabrous or essentially so. 



Fruiting calyces 1-2 cm. long, 0.6-1.5 cm. wide, the lobes 1-2 mm. long; 



corolla limb narrow, 2-3 mm. wide. 



Leaves succulent, the blades of the larger leaves 1.5-2.5 cm. long; 

 petioles 5-10 mm. long; fruiting calyces 1-1.3 cm. long, 0.6-0.8 



cm. wide P. carnosa. 



Leaves not succulent, the blades of the larger leaves 3-6 cm. long; 

 petioles 10-35 mm. long; fruiting calyces 1.5-2 cm. long, 1.2-1.5 



cm. wide P. minuta. 



Fruiting calyces 2.5-5.5 cm. long, 2-3 cm. wide, the lobes 4-13.5 mm. 



long; corolla limb broad, 10-20 mm. wide. 



Corolla maculations prominent; fruiting calyces usually gradually 

 terminated apically 2.5-4 cm. long, the lobes 4-6 mm. long; 

 1,000 m. and below P. cordata. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 81 



Corolla maculations inconspicuous, their edges often diffuse; fruiting 



calyces usually abruptly beaked, (3-) 3.5-5.5 cm. long, the lobes 



(8-) 10-13.5 mm. long; 1,500-2,500 m., rarely lower. P. porrecta. 



Fruiting calyces pubescent, the hairs sometimes short and inconspicuous. 



Fruiting calyces densely pubescent with grayish, short hairs; gynophore 



and inside of fruiting calyx capitate-glandular P. ignota. 



Fruiting calyces variously pubescent but not densely pubescent with 

 grayish, short hairs; gynophore and inside of fruiting calyx not capi- 

 tate-glandular except sometimes in P. pubescens. 

 Fruiting calyx lobes narrow, subulate and long-acuminate, 10-15 mm. 



long. 



Stems, petioles, and pedicels with at least a few long hairs 1.5-3.5 

 mm. long (in ours) ; corolla limb 10-15 mm. wide; flowering pedi- 

 cels 10-75 mm. long P. maxima. 



Stems, petioles and pedicels with short hairs, the hairs less than 1 

 mm. long; corolla limb 6-8 mm. wide; flowering pedicels 2-4 

 mm. long P. nicandroides. 



Fruiting calyx lobes broader, ovate or deltoid to lanceolate or tri- 

 angular, 2-6.5 mm. long. 

 Corolla immaculate or the spots not strongly contrasting. 



P. hylophila. 



Corolla maculate, the spots strongly contrasting with the surround- 

 ing corolla limb. 

 Fruiting calyces narrow, about twice as long as wide; montane 



forests, 2,500-2,900 m P. angustiphysa. 



Fruiting calyces broad, as long as wide to 1.5 times longer than 



wide; 1,000 m. and below. 



Leaf blades thin, more or less translucent when dry; corolla 

 limb 6-8 mm. wide, the throat glabrous or nearly so; fruit- 

 ing calyces 1.2-2.5 cm. long, 1-1.5 cm. wide, the lobes ovate 



to deltoid, the apex acute, 3-3.5 mm. long P. hirsuta. 



Leaf blades thick, opaque; corolla limb 8-15 mm. wide, the 

 throat densely pubescent; fruiting calyces 2-4 cm. long, 1.2- 

 3 cm. wide, the lobes triangular to narrowly lanceolate, the 

 apex narrowly acute to acuminate, (3.5-) 4.5-6.5 mm. long. 



P. pubescens. 

 Fruiting calyx 10-ribbed, 10- angled or teretish. 



Corolla 2-7 mm. long, the limb narrow, 1.5-5.5 mm. wide; anthers small, 



0.4-1.5 mm. long; fruiting calyces 0.5-2 cm. long, 0.5-1.5 cm. wide. 

 Corolla 2-3 mm. long, the limb 1.5-2 mm. wide; anthers 0.4-0.6 mm. long; 

 fruiting calyces 0.5-1 cm. long, 0.5-0.6 cm. wide. . . .P. microcarpa. 

 Corolla 5-7 mm. long, the limb 4.5-5.5 mm. wide; anthers 1.2-1.5 mm. 



long; fruiting calyces 1-2 cm. long, 1-1.5 cm. wide P. lagascae. 



Corolla (6-) 8-15 mm. long, the limb broad, 7-24 mm. wide; anthers large, 

 2-4 mm. long; fruiting calyces 2-3.5 cm. long, 1.5-2.5 cm. wide. 



Stems and fruiting calyces evenly and densely pubescent P. lassa. 



Stems and fruiting calyces glabrous to sparsely pubescent. 



Corolla immaculate or with spots light colored and not strongly con- 

 trasting; fruiting pedicels 20-40 mm. long P. angulata. 



Corolla strongly maculate, the spots sometimes confluent basally; 



fruiting pedicels 3.5-20 mm. long. 



Fruiting pedicels 3.5-8 (-12) mm. long; anthers blue or yellowish 

 with blue margins, 2.5-3 mm. long, often contorted after dehis- 



cence; filaments 2 mm. long P. philadelphica. 



Fruiting pedicels 10-20 mm. long; anthers yellowish, sometimes with 

 a slight bluish tinge, 3-4 mm. long, relatively straight after 

 dehiscence; filaments 2-5 mm. long P. gracilis. 




82 FIELDIANA: BOTANY, VOLUME 24 



Physalis amphitricha (Bitter) Standl. & Steyerm. Field Mus. 

 Bot. 23: 229. 1947. Saracha amphitricha Bitter, Fedde Repert. Sp. 

 Nov. 20:362. 1924. P. arnica Standl. & Steyerm. Field Mus. Bot. 

 23: 231. 1947 (type from between La Vega ridge along Rio Vega and 

 northeast slopes of Volcan Tacana, to 3 mi. from Guatemala-Mexico 

 boundary, in vicinity of San Rafael, San Marcos, Steyermark 36175], 

 syn. nov. Flor de miltomate (Quezaltenango) . 



Dense, moist or wet forests or mixed forests, sometimes on white 

 sand slopes, 1,300-2,700 m.; Quezaltenango (type collected between 

 San Martin, Sacatepe"quez and Las Mercedes, Bernoulli & Cario 

 2335) ; San Marcos. Southern Mexico (Chiapas) . 



Herb 1-3 m. tall, branched, stem usually herbaceous, sometimes suffrutescent, 

 glabrous or nearly so or sometimes covered with multicellular hairs, glabrescent 

 in age; leaves mostly entire or shallowly sinuate, elliptic to ovate or lanceolate, 

 the larger leaves mostly 6-20 cm. long, 3-8 cm. wide, the apex acuminate or long 

 acuminate, the base short attenuate or acute to obtuse, pubescence of short, 

 multicellular hairs above and below, the petioles 1-4 cm. long; inflorescence con- 

 sisting of (l-)2-5 flowers, sometimes only one maturing a fruit, the pedicels 8-20 

 mm. long, thickened above; flowering calyx 7-13.5 mm. long, mostly glabrous, or 

 sometimes densely pubescent with short, multicellular hairs, the lobes ovate to 

 lanceolate, 2.5-10 mm. long; corolla yellowish or greenish-yellow, the limb 25-35 

 mm. wide, the lobes ovate or lanceolate, 9.5-15 mm. long, marked with greenish- 

 purple spots near their bases, or rarely without prominent maculations, the corolla 

 tube, throat and the base of the corolla lobes densely pubescent; filaments 3.5-5 

 mm. long; anthers bluish to purplish, 3-4 mm. long; fruiting calyx 5-ribbed, 5.5 

 cm. long, 3.5 cm. wide, glabrous or essentially so, reticulate-veined, strongly in- 

 flated, the pedicels 17 mm. long; berry about 10 mm. in diameter. 



Physalis amphitricha (Bitter) Standl. & Steyerm., P. calidaria 

 Standl. & Steyerm., and P. porphyrophysa Donn.-Sm. are members of 

 an apparently natural group of species characterized by the deeply 

 lobed corolla limb, the lobes valvate in bud, and the inflorescence 

 consisting of mostly axillary fascicles. Another taxon, P. melano- 

 cystis (Robinson) Bitter var. cernua (Donn.-Sm.) Waterfall, included 

 in this group by Waterfall is treated under the genus Athenaea. See 

 comments under Athenaea. 



Physalis angulata L. Sp. PI. 1: 183. 1753. 



Known in our area from only two collections from British Hon- 

 duras; 1,000 m. or below. Eastern to Central United States. Mexico. 

 El Salvador; Nicaragua; Costa Rica. Panama. West Indies. South 

 America. Old World tropics. 



Herbs 1 m. tall or less, often much branched, the stems glabrous or nearly 

 so; leaves coarsely sinuate-dentate or sometimes entire, ovate to ovate-lanceolate, 




GENTRY & STANDLEY: FLORA OF GUATEMALA 83 



the larger leaves mostly 6.5-15 cm. long, 2.5-8 cm. wide, the apex acuminate, the 

 base acute or obtuse, sparsely pubescent on the veins, otherwise glabrous above 

 and below, the petioles (l-)2.5-5 cm. long; flowers solitary, the pedicels 10-12 mm. 

 long; flowering calyx 3-4 mm. long, pubescent with appressed hairs, the lobes 

 deltoid or triangular, 1.5 mm. long; corolla yellowish, 6-8 mm. long, the limb 7-11 

 mm. wide, immaculate or with indistinct spots, somewhat pubescent in the mouth 

 of the corolla tube; filaments 2.5-3 mm. long; anthers bluish or violet, 1.5-2.5 mm. 

 long; fruiting calyx 10-angled or 10-ribbed, 2.5-3 cm. long, 2-2.5 cm. wide, sparsely 

 pubescent on the veins with short hairs or essentially glabrous, reticulate, the 

 pedicels 20-40 mm. long; berry 10-12 mm. in diameter. 



Physalis angulata L. has been separated by Waterfall into three 

 varieties. We apparently have only variety angulata in our area. 



Physalis angustiphysa Waterfall, Rhodora 69: 228. 1967. 



Known in Guatemala only from the type, La Sierra (Tujimach), 

 across the river from San Juan Atitlan, Sierra de los Cuchumatanes, 

 2,500-2,900 m., Huehuetenango, Steyermark 51977. Mexico. 



Herb 1 m. tall, the stems densely pubescent with long, multicellular hairs of 

 varying length, some capitate-glandular; leaves sinuately lobed, ovate to broadly 

 ovate, the larger leaves 4-9.5 cm. long, 2-4.5 cm. wide, the apex acute, the base 

 rounded, densely pubescent with long, multicellular hairs, some hairs capitate- 

 glandular above and below, the petioles 1-3 cm. long; flowers solitary, the pedicels 

 5-6.5 mm. long; flowering calyx 3-5 mm. long, densely pubescent, the lobes lance- 

 olate, 2-2.5 mm. long; corolla yellow, 8-9 mm. long, the limb 12-14 mm. wide, 

 maculate; filaments 3-4 mm. long; anthers violaceous, 2-2.5 mm. long; fruiting 

 calyx 5-angled, 2.4-2.8 cm. long, 1-1.8 cm. wide, reticulate, pubescent, the pedicels 

 7.5-10 mm. long; berry about 6 mm. in diameter. 



Physalis calidaria Standl. & Steyerm. Field Mus. Bot. 23: 231. 

 1947. P. chimalteca Standl. & Steyerm. I.e. 232 (type from slopes of 

 Volcan de Acatenango, above Las Calderas, Chimaltenango, Stand- 

 ley 61803], syn. nov. 



Wet forests, 2,300-2,850 m.; Chimaltenango; Quiche"; Quezal- 

 tenango (type from Fuentes Georginas, western slope of Volcan de 

 Zunil, Quezaltenango, Standley 67488). Southern Mexico (Chiapas). 



Herb 0.5-2 m. tall, possibly suffrutescent below, the stems glabrous or pubes- 

 cent with short, brownish, incurved hairs, glabrescent in age; leaves entire or with 

 a few angulate lobes, ovate to lance-elliptic or elliptic, the larger leaves 8-20 cm. 

 long, 4-9 cm. wide, the apex acuminate, the base acute or obtuse, sometimes short 

 attenuate, a few scattered, multicellular hairs above, more densely pubescent, 

 especially on the veins below, the petioles 1-3 cm. long; inflorescence consisting of 

 (l-)2-5 flowers, often only one maturing a fruit, the pedicels 8-16 mm. long, 

 thickened above; flowering calyx 3-10 mm. long, glabrous except for the ciliolate 

 margin or densely pubescent throughout, nearly truncate to shallowly and broadly 

 lobed, the lobes 1 mm. long or less; corolla greenish-yellow or pale yellow, the limb 




84 FIELDIANA: BOTANY, VOLUME 24 



20-35 mm. wide, maculate, the lobes ovate, the corolla tube, throat and the base 

 of the corolla lobes densely pubescent; filaments 3-5 mm. long; anthers 2.5-3 mm. 

 long; fruiting calyx 2.5-5 cm. long, 2-3 cm. wide, 10-angled, glabrous, reticulate- 

 veined, strongly inflated; berry 8-10 mm. in diameter. 



Phy salis chimalteca Standl. & Steyerm. has been separated from 

 a closely related species, P. calidaria Standl. & Steyerm., based upon 

 its essentially glabrous calyx. It is treated here as a synonym of P. 

 calidaria. 



Physalis campanula Standl. & Steyerm. Field Mus. Bot. 23: 

 18. 1943. 



Dry banks along stream, 2,200-2,500 m.; San Marcos (type from 

 along Quebrada Canjula, between Sibinal and Canjula, Volcan 

 Tacana, Steyermark 36067). Endemic. Known only from the type 

 collection. 



A branching herb, the stems covered with spreading, multicellular hairs of 

 varying lengths, some capitate-glandular, glabrescent in age; leaves entire or 

 shallowly sinuately lobed, ovate, the larger leaves mostly 5-12 cm. long, 3-5 cm. 

 wide, the apex acuminate or acute, the base rounded, pubescence of multicellular 

 hairs, some capitate-glandular above and below, the petioles 1-4.5 cm. long; flowers 

 solitary, the pedicels 20 mm. long; flowering calyx 10-11 mm. long, densely pubes- 

 cent with multicellular hairs, some capitate-glandular, the lobes triangular, 3.5-5 

 mm. long, long acuminate; corolla tubular-campanulate, pale yellow, immaculate, 

 the tube 18-19 mm. long, pubescent below the middle, slightly ventricose above, 

 the limb obscurely dentate; filaments 8-9 mm. long; anthers bluish, about 3 mm. 

 long; fruiting calyx (only one approaching maturity) 25mm. long, 15 mm. wide, 

 densely pubescent with multicellular hairs, some capitate-glandular, the pedicels 

 25-30 mm. long; berry about 10 mm. in diameter. 



Physalis campanula Standl. & Steyerm. may be separated from 

 all other members of the genus in Central America by its tubular- 

 campanulate corolla. 



Physalis carnosa Standl. & Steyerm. Field Mus. Bot. 23: 19. 

 1943. 



On sand dunes along the coast, 1-2 m.; San Marcos (type from 

 Oc6s, Steyermark 37766). Endemic. Known only from the type 

 collection. 



A much branched herb, prostrate or decumbent, the stems 15-20 cm. long, 

 pubescence of minute, incurved and appressed hairs, glabrescent in age; leaves 

 succulent, entire or shallowly sinuately lobed, mostly ovate to ovate-oblong or 

 elliptic, the larger leaves 12-28 mm. long, 7-12 mm. wide, the apex acute, the 

 base obtuse, nearly glabrous above and below, the hairs minute, incurved and 

 appressed, the petioles 5-10 mm. long; flowers solitary, the pedicels 2.5-3 mm. long; 

 flowering calyx 2-3 mm. long, pubescent with minute, incurved and appressed 




GENTRY & STANDLEY: FLORA OF GUATEMALA 85 



hairs, the lobes narrowly triangular, 0.5-1 mm. long, acute; corolla 3-4 mm. long, 

 long, probably immaculate; filaments about 1.5 mm. long; anthers bluish-green, 

 0.8-1.2 mm. long; fruiting calyx 5-angled, 1-1.3 cm. long, 0.6-0.8 cm. wide, reticu- 

 late, inconspicuously pubescent, the pedicels 4-5 mm. long; berry about 5 mm. in 

 diameter. 



Physalis carnosa Standl. & Steyerm., P. lagascae Roem. & Schult., 

 P. microcarpa Urban & Ekman, and P. minuta Griggs represent the 

 species with relatively small fruiting calyces, 0.5-2 cm. long, 0.5- 

 1.5 cm. wide, that occur or are likely to occur in Guatemala. 



Physalis cordata Miller, Gard. Diet. ed. 8, Physalis no. 14. 1768. 



Wet thickets or rocky river banks, sometimes on sand dunes, 

 1,000 m. and below; Izabal; Jutiapa; Santa Rosa; Escuintla; Retal- 

 huleu; San Marcos. Mexico to Costa Rica. Panama and West 

 Indies. 



Herb 1.5 m. tall or long, erect or sometimes decumbent, the stems pubescent 

 with very short, incurved and inconspicuous hairs; leaves coarsely sinuate-den- 

 tate or entire, ovate, the larger leaves 4-18 cm. long, 2.5-8 cm. wide, the apex 

 acuminate or acute, the base obtuse to subcordate or nearly truncate, pubescent 

 with very short and inconspicuous hairs, these more prominent on the veins above 

 and below, the petioles 2.5-9 cm. long; flowers solitary, the pedicels 5-10 mm. long; 

 flowering calyx 5-6 mm. long, pubescent with short, incurved hairs, the lobes nar- 

 rowly triangular, about 3 mm. long; corolla yellowish, 5-10 mm. long, the limb 

 maculate, 10-20 mm. wide, pubescent internally; filaments 1.5-2 mm. long; an- 

 thers bluish or greenish-blue, about 2 mm. long; fruiting calyx 5-angled, reticu- 

 late, 2.5-4 cm. long, 2-2.5 cm. wide, glabrous, the pedicels 10-25 mm. long; berry 

 6-15 mm. in diameter. 



Physalis cordata Miller had not been recognized in American floras 

 since its description until the work of Waterfall in 1967. Most of 

 the material in herbaria has been identified as P. turbinata Medic. 

 The length of the fruiting calyx lobes has proven to be a useful charac- 

 ter in separating this species from Physalis porrecta Waterfall. 



Physalis gracilis Miers, Ann. Mag. Nat. Hist. ser. 2, 4: 37. 1849. 

 P. luteoanthera Waterfall, Rhodora 69: 113. 1967 (type from lower 

 south facing slopes of Volcan Santa Maria, Steyermark 33726), syn. 

 nov. Bombita (Izabal); kop and miltomate (Alta Verapaz). 



Moist or wet thickets or forests, sometimes in wet pine forests, 

 just above sea level to 1,500 m., rarely higher; Pete"n; Alta Verapaz; 

 Izabal; Baja Verapaz; Guatemala; Quezaltenango; Huehuetenango ; 

 Escuintla; Suchitepe"quez ; Retalhuleu; San Marcos. British Hon- 

 duras. Mexico to Costa Rica. 




86 FIELDIANA: BOTANY, VOLUME 24 



Herbs 1 m. tall or less, erect to decumbent or procumbent, the stems pubes- 

 cent with long, multicellular hairs, sometimes nearly glabrous; leaves entire to 

 undulate or shallowly angulately lobed, ovate to broadly ovate, the larger leaves 

 4-16 cm. long, 2.5-8 cm. wide, the apex acute or acuminate, the base obtuse or 

 rounded, sometimes subcordate, pubescent with multicellular hairs or sometimes 

 nearly glabrous above and below, the petioles 1-6 cm. long; flowers solitary, the 

 pedicels 7-15(-20) mm. long; flowering calyces 5-10 mm. long, pubescent with 

 usually spreading hairs, sometimes glabrous, the lobes ovate-deltoid, 2-3 mm. 

 long; corolla yellow or greenish-yellow, 10-13 mm. long, the limb 12-24 mm. wide, 

 maculate; filaments 2-5 mm. long; anthers yellowish, sometimes with a slight bluish 

 tinge, 3-4 mm. long; fruiting calyx 10-ribbed to nearly terete, 2-3 cm. long, 1.5- 

 2.5 cm. wide, sparsely pubescent, reticulate, the pedicels 10-20 mm. long; berry 

 8-15 mm. in diameter. 



The characters that distinguish Physalis gracilis Miers and P. 

 philadelphica Lam. are subject to some overlap, but they appear to 

 be distinct species. 



Physalis hirsuta Dunal in DC., Prodr. 13, pt. 1: 445. 1852. P. 

 leptophylla Robinson & Greenm. Proc. Am. Acad. 29: 389. 1894, syn. 

 nov. 



Moist thickets or brushy rocky slopes, 400-660 m.; Zacapa; Chi- 

 quimula. Mexico. 



Herb 0.5-1 m. tall, the stems pubescent with multicellular hairs of varying 

 lengths, at least some capitate-glandular; leaves thin, entire or rarely with a few 

 shallow lobes, ovate, 6-16 cm. long, 2.5-6 cm. wide, the apex acuminate, the base 

 rounded, ciliate, sparsely pubescent with short multicellular hairs, these mostly 

 on the veins or nearly glabrous above and below, the petioles 2-8 cm. long; flowers 

 solitary, the pedicels 3-4. 5 mm. long; flowering calyx 2. 5-4 mm. long, densely pubes- 

 cent with spreading hairs, the lobes ovate to deltoid, 1-2.5 mm. long; corolla yel- 

 lowish or cream-colored, 6-8.5 mm. long, the limb maculate, 6-8 mm. wide, gla- 

 brous or nearly so internally; filaments 2-3 mm. long; anthers bluish, rarely yel- 

 lowish in age, 1-1.5 mm. long; fruiting calyx thin, 5-angled, the angles usually not 

 prominent, 1.2-2.5 cm. long, 1-1.5 cm. wide, pubescent with spreading hairs, the 

 pedicels 4-5 (-9) mm. long; berry 5-9 mm. in diameter, sessile or subsessile. 



We are following D'Arcy (personal communication) in the applica- 

 tion of this name. 



Physalis hylophila Standl. J. Wash. Acad. Sci. 14: 243. 1924. 

 Wooded slopes. Mexico. El Salvador. To be expected in Guate- 

 mala. 



Herb 1 m. tall or less, the stems pubescent with multicellular hairs of vary- 

 ing lengths, at least some capitate-glandular; leaves entire or with a few irregular 

 sinuate teeth, ovate, 3.5-6.5 cm. long, 2-2.5 cm. wide, the apex acuminate, the 

 base rounded, pubescent with multicellular hairs, at least some capitate-glandular 




GENTRY & STANDLEY: FLORA OF GUATEMALA 87 



above and below, the petioles 1-2.5 mm. long; flowers solitary, the pedicels 3-5 

 mm. long; flowering calyx 3-5.5 mm. long, densely pubescent with glandular hairs, 

 the lobes ovate or deltoid, about 1 mm. long; corolla yellowish, 6-7 mm. long, the 

 limb immaculate or with spots not strongly contrasting, 7-8 mm. wide, glabrous 

 internally; filaments 3-4 mm. long; anthers yellowish, greenish-yellow or bluish 

 tinged, about 1 mm. long; fruiting calyx 5-angled, the angles not prominent, 1.4- 

 1.6 cm. long, 1-1.2 cm. wide, pubescent with spreading hairs, the pedicels 3-5 mm. 

 long; berry about 6 mm. in diameter. 



Phy salis hylophila Standl. may prove to be a synonym of P. 

 hirsuta Dunal. 



Physalis ignota Britton, Mem. Torrey Bot. Club 16: 100. 1920. 

 P. pentagona Blake, Contr. U.S. Natl. Herb. 24: 20. 1922 (type from 

 Los Amates, Izabal, Blake 7313). Sopldn (Jutiapa). 



Damp or moist thickets, sometimes on gravelly or rocky slopes, 

 45-900 m.; Izabal; Zacapa; Jalapa; Chiquimula; Jutiapa; San Mar- 

 cos. El Salvador to Costa Rica. Panama and West Indies. 



Herb 1 m. tall or less, the stems mostly densely covered with grayish, short 

 and spreading, multicellular hairs; leaves entire or repand, ovate, the larger leaves 

 5-18 cm. long, 3.5-8 cm. wide, the apex acuminate, the base obtuse or acute, pubes- 

 cent with short, multicellular hairs, more dense on the veins above and below, 

 the petioles 2-6 cm. long; flowers solitary, the pedicels 3-8 mm. long; flowering 

 calyx 4.5-6 mm. long, densely pubescent with short, spreading hairs, the lobes 

 narrowly lanceolate or triangular, 2-3 mm. long; corolla yellowish, 6-10 mm. long, 

 the limb immaculate, 5-10 mm. wide, sparsely pubescent internally; filaments 2- 

 2.5 mm. long; anthers yellow or bluish tinged, 2-2.5 mm. long; fruiting calyx 

 strongly 5-angled, reticulate, 3-5 cm. long, 2.5-3.5 cm. wide, mostly densely cov- 

 ered with grayish, short and spreading, multicellular hairs, the pedicels 8-15 mm. 

 long; berry 10-15 mm. in diameter, on a gynophore 1-2 mm. long, capitate- 

 glandular. 



Physalis ignota Britton can be separated from all other species 

 of the genus in Guatemala by the large and strongly 5-angled fruit- 

 ing calyces and the stems, pedicels, and fruiting calyces densely 

 pubescent with grayish, short hairs. 



Physalis lagascae Roem. & Schult. Syst. Veg. 4: 679. 1819. P. 

 micrantha Link, Enum. Hort. Berol. 1: 181. 1821. P. parviculea 

 Blake, Contr. U. S. Natl. Herb. 24: 20. 1922 (type from Los Amates, 

 Izabal, Blake 7318). 



Grassy slopes, 900-1,500 m.; Izabal; Chiquimula. Mexico. El 

 Salvador; Honduras. Panama and Martinique. 



Herb 1.5 m. tall or less, erect, the stems pubescent with long, multicellular 

 hairs; leaves entire or slightly repand, ovate to lanceolate, the larger leaves 2-10 

 cm. long, 1-3.5 cm. wide, apex acuminate or narrowly acute, the base obtuse, 

 rounded or sometimes acute, pubescent with relatively long hairs above and 




88 FIELDIANA: BOTANY, VOLUME 24 



below, the petioles 0.5-2.5 cm. long; flowers solitary, the pedicels 2-5 mm. long; 

 flowering calyx 3-4 mm. long, pubescent with long, spreading hairs, the lobes 

 deltoid, 0.6-1.5 mm. long; corolla 5-7 mm. long, maculate, the spots usually not 

 conspicuous, sometimes appearing immaculate, pubescent internally; filaments 

 1.5 mm. long; anthers bluish or violet, 1.2-1.5 mm. long; fruiting calyx 10-ribbed, 

 1.2-2 cm. long, 1-1.5 cm. wide, sparsely pubescent with long spreading hairs mostly 

 on the ribs, often the ribs with widely spaced small teeth or enations produced 

 mostly by the base of the hairs, the pedicels 3-5 mm. long; berry 5-7 mm. in 

 diameter. 



Physalis lagascae var. glabrescens Schulz in Urban, Sym. An- 

 till. 6: 147. 1906. 



Moist fields or thickets, sometimes in rocky thickets, 200-850 

 m.; Zacapa; Jutiapa; Santa Rosa. Mexico. El Salvador to Costa 

 Rica. Panama and Cuba. 



Variety glabrescens differs from variety lagascae by being glabrous 

 or nearly glabrous. 



Physalis lagascae Roem. & Schult. is often confused with Mar- 

 garanthus. The fruiting calyces are similar but the corollas are very 

 distinctive. See comments under Margaranthus. 



Physalis lassa Standl. & Steyerm. Field Mus. Bot. 23: 19. 1943. 

 Tomatillo (Huehuetenango). 



In damp oak forests or in grassy thickets, 250-2,000 m.; Baja 

 Verapaz; Zacapa (type from between Jalapa and Montana Mira- 

 mundo, Steyermark 32868); Guatemala; Chimaltenango; Huehue- 

 tenango; Jalapa. Mexico. 



Herbs 1 m. tall or less, the stems densely covered with multicellular hairs, 

 sometimes most of the hairs capitate-glandular; leaves entire or sometimes repand, 

 ovate, the larger leaves 3.5-9 cm. long, 2-4 cm. wide, the apex acute or acuminate, 

 the base rounded or subcordate, mostly densely pubescent with multicellular hairs, 

 sometimes most of the hairs capitate-glandular above and below, the petioles 

 1-2.5 cm. long; flowers solitary, the pedicels 5-8 mm. long; flowering calyx 8-10 

 mm. long, pubescent with spreading, multicellular hairs, sometimes most of the 

 hairs capitate-glandular, the lobes ovate or ovate-lanceolate, 2.5-5 mm. long; 

 corolla yellowish, 10-15 mm. long, the limb 12-15 mm. wide, maculate, pubescent 

 in the mouth of the corolla tube; filaments 2.5-3 mm. long; anthers bluish, 2.5-4 

 mm. long; fruiting calyx 10-ribbed or slightly 10-angled, 2.5-3.5 cm. long, 1.5-2.5 

 cm. wide, evenly pubescent, sometimes densely so, reticulate, the pedicels 10-15 

 mm. long; berry 10-15 mm. in diameter. 



The pubescence is variable in this species. The type and a few 

 additional collections have eglandular hairs, whereas two collections, 

 Steyermark 29545 and 50638 have mostly glandular hairs. These two 

 collections may prove to be worthy of subspecific rank. 




D 



FIG. 17. Physalis lagascae var. glabrescens. A, habit, X ^2; B, flower, X 4; 

 C, fruiting calyx, X 2. Schwenckia americana. D, habit, X H; E flower, X 3; 

 F, corolla lobes, detail, X 5. 



89 




90 FIELDIANA: BOTANY, VOLUME 24 



Physalis maxima Miller, Gard. Diet. ed. 8, Physalis no. 15. 

 1768. 



Dry slopes or moist thickets, 600-1,200 m. ; Jalapa; Baja Verapaz; 

 Guatemala; Jutiapa; Escuintla. Mexico. Honduras; Nicaragua; 

 Costa Rica. 



Herb 1.5 m. tall or less, the stem pubescent with short spreading hairs, at 

 least some capitate-glandular, more robust hairs 1.5-3.5 mm. long present (in ours) 

 in addition to the shorter ones on the stem, petioles, and pedicels; leaves coarsely 

 sinuate-dentate to shallowly angulately-lobed or sometimes entire, ovate, the 

 larger leaves mostly 8-30 cm. long, 5-14 cm. wide, the apex acuminate, the base 

 rounded to cordate, pubescent with multicellular hairs above and below, the 

 petioles 3-12 cm. long; flowers solitary, the pedicels 10-25(-75) mm. long, often 

 with long hairs; flowering calyx 5-8 mm. long, often pubescent with long, spreading 

 hairs, the lobes subulate, long-acuminate, 3-6 mm. long; corolla white or whitish, 

 6-8 mm. long, the limb slightly to moderately maculate, 10-15 mm. wide, pubes- 

 cent internally; filaments 2.5-4 mm. long; anthers yellow or slightly bluish-tinged, 

 2.5-3 mm. long; fruiting calyx strongly 5-angled, reticulate, 4-6 cm. long, 2.5-4 

 cm. wide, pubescent, the pedicels 20-30(-80) mm. long; berry 10-20 mm. in 

 diameter. 



Physalis maxima Miller is closely related to P. nicandroides 

 Schlecht., especially variety attenuata Waterfall. It can be separated 

 in most cases from that variety by the long hairs on the stem, peti- 

 oles, and pedicels and by the wider corolla limb. Some specimens 

 are difficult to place. 



The name Physalis maxima Miller had not been used in floristic 

 work since its original description until Waterfall's treatment in 

 1967. Prior to that time, most of the specimens were identified as P. 

 nicandroides Schlecht. 



Physalis microcarpa Urban & Ekman in Urban, PI. Haitiensis, 

 Arkiv Bot. 21A(5): 59. 1927. Miltomate de culebra (Jutiapa). 



In moist thickets, 300-850 m.; Zacapa; Chiquimula; Jutiapa. 

 El Salvador; Honduras. West Indies. 



Herb 0.4 m. tall or less, slightly to much branched, erect or sometimes nearly 

 prostrate, the stems with short, incurved hairs; leaves entire or slightly repand, 

 lanceolate to lance-ovate, the larger leaves 1.5-6 cm. long, 0.5-2 cm. wide, the 

 apex acuminate, sometimes long acuminate, the base obtuse, rounded or acute, 

 pubescent with short, incurved hairs above and below, the petioles 0.4-2 cm. long; 

 flowers solitary, the pedicels 0.5-2 mm. long; flowering calyx 1.2-2 mm. long, 

 pubescent with appressed hairs, the lobes deltoid or ovate, 0.4-0.6 mm. long; 

 corolla 2-3 mm. long, immaculate, slightly pubescent internally; filaments 0.5-0.7 

 mm. long; anthers bluish or violet, sometimes yellowish (perhaps due to fading), 

 0.4-0.6 mm. long; fruiting calyx nearly terete, slightly 10-ribbed, 0.5-1 cm. long, 

 0.5-0.6 mm. wide, sparsely pubescent with relatively short hairs, the pedicels 3-4 

 mm. long; berry 3-4.5 mm. in diameter. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 91 



In addition to the characters in the key, Physalis microcarpa 

 Urban & Ekman differs from P. lagascae Roem. & Schult. by the 

 shorter hairs on the fruiting calyx and the somewhat narrower fruit- 

 ing calyx lobes. Physalis lagascae often has widely spaced small 

 teeth or enations produced mostly by the base of the hairs on the 

 ribs of the fruiting calyx. 



Physalis minuta Griggs, Torreya 3: 138. 1903. 



Sandy fields or a weed in cultivated fields, 850 m. or below. Mex- 

 ico. Honduras; Nicaragua; Costa Rica. Panama. To be expected 

 in Guatemala. 



Herb 0.4 m. tall or less, erect, the stem pubescent with short, incurved hairs; 

 leaves entire or slightly repand, ovate to lanceolate, the larger leaves 3.5-9 cm. 

 long, 1.5-3.5 cm. wide, the apex acuminate or acute, the base obtuse or acute, 

 pubescent with very short, incurved hairs mostly on the veins above and below, the 

 petioles 1-3.5 cm. long; flowers solitary, the pedicels 3-6 mm. long; flowering calyx 

 2-3 mm. long, pubescent with appressed hairs, the lobes triangular, 1-1.5 mm. 

 long; corolla 4-5.5 mm. long, immaculate or with slightly contrasting spots, pubes- 

 cent internally; filaments about 1.5 mm. long; anthers bluish tinged or yellowish, 

 1-2 mm. long; fruiting calyx 5-angled, 1.5-2 cm. long, 1.2-1.5 cm. wide, minutely 

 and sparsely pubescent, the pedicels 4-10 mm. long; berry about 8 mm. in diameter. 



Physalis minuta Griggs is restricted apparently to the Pacific 

 slope in Central America. 



Physalis nicandroides Schlecht. Linnaea 19: 311. 1846. 



Moist thickets or brushy hillsides, often on rocky slopes, some- 

 times in cultivated ground, 850-1,830 m.; Guatemala; Chimalten- 

 ango; Quiche"; Jutiapa. Mexico. Honduras; Costa Rica. 



Herb 1-2 m. tall, the stems, petioles, and pedicels glandular-pubescent, the 

 hairs less than 1 mm. long; leaves coarsely sinuate-dentate to shallowly angulately- 

 lobed, ovate, the larger leaves 6.5-20 cm. long, 3-10 cm. wide, the apex acute or 

 acuminate, the base acute to subcordate, glandular-pubescent above and below, 

 the petioles 2-6.5 cm. long; flowers solitary, the pedicels 2-4 mm. long; flowering 

 calyx 5-6.5 mm. long, glandular-pubescent with spreading hairs, the lobes subulate, 

 long-acuminate, 3-4.5 mm. long; corolla white, greenish-white or yellowish, 4-8 

 mm. long, the limb maculate, but often faded, 6-8 mm. wide, pubescent internally; 

 filaments 2-2.5 mm. long; anthers blue tinged to yellowish, 1.5-2 mm. long; fruit- 

 ing calyx strongly 5-angled, reticulate, 3-4.5 cm. long, 2-4 cm. wide, pubescent, 

 the pedicels short and thick, 5-10 mm. long, (1-)1. 5-1.7 mm. thick; berry 12-20 

 mm. in diameter. 



Physalis nicandroides var. attenuata Waterfall, Rhodora 69: 

 235. 1967. 




92 FIELDIANA: BOTANY, VOLUME 24 



Damp or wet thickets, sometimes on rocky slopes, 350-1,300 m., 

 rarely higher; Zacapa; Chiquimula; Jalapa; Sacatepe"quez; Huehue- 

 tenango; Jutiapa; Santa Rosa. Mexico. Honduras; Costa Rica. 



Variety attenuate, differs from variety nicandroides by its longer 

 and more slender fruiting pedicels, 10-25 mm. long, 0.5-1 mm. thick. 



Physalis philadelphica Lam., Encycl. Method. Bot. 2: 101. 

 1786. P. philadelphica Lam. forma pilosa Waterfall, Rhodora 69: 

 214. 1967 (type from Finca la Alameda near Chimaltenango, Chi- 

 maltenango, Standley 79899], syn. nov. Miltomate; Miltomate Santo 

 Tomas (Chimaltenango) ; tomatillo (Jutiapa) . 



Moist or damp thickets or fields, often a weed in cultivated 

 ground, sometimes in pine-oak forests; just above sea level to 1,830 

 m., rarely higher; Alta Verapaz; Izabal; Chiquimula; Jalapa; Gua- 

 temala; Baja Verapaz; Sacatepe"quez; Chimaltenango; Solola; Qui- 

 che"; Huehuetenango; Jutiapa; Escuintla. Mexico. El Salvador; 

 Costa Rica. Panama and West Indies. 



Herb 1 m. tall or less, the stems sparsely pubescent with mostly short hairs to 

 glabrous; leaves coarsely sinuate-dentate or dentate, sometimes entire, ovate to 

 ovate-lanceolate, the larger leaves mostly 3.5-12.5 cm. long, 1.5-6 cm. wide, the 

 apex acuminate, the base acute or obtuse, glabrous or sometimes with a few hairs 

 on the veins above and below, the petioles mostly 2-5 cm. long; flowers solitary, 

 the pedicels 3-7.5 mm. long; flowering calyx 3.5-6.5 mm. long, pubescent with 

 spreading, multicellular hairs, the lobes ovate, 1.5-3 mm. long; corolla yellowish, 

 8-12 mm. long, the limb 10-18 mm. wide, maculate; filaments about 2 mm. long; 

 anthers blue or yellowish with bluish margins, usually strongly contorted after 

 dehiscence, 2.5-3 mm. long; fruiting calyx 10-ribbed, 2-3 cm. long, 2-2.5 cm. wide, 

 glabrous or sometimes sparsely pubescent, reticulate, the pedicels 3.5-8 mm. long; 

 berry 15-20 mm. in diameter, often nearly filling the fruiting calyx. 



There has been a great deal of confusion in the application of a 

 name for this species and its synonymy. Physalis ixocarpa Brotero 

 ex Horn, and P. aequata Jacq. f. ex Nees were treated as synonyms 

 of P. philadelphica Lam. by Waterfall in 1967. However, recent 

 work by Fernandes (Bol. Soc. Brot. 44: 343-366, 10 plates, 1970) in- 

 dicates that the two names cannot be applied to P. philadelphica but 

 represents a separate species not known to occur in the New World. 



See comment under Physalis gracilis Miers. 



Miltomates (the name is Nahuatl, signifying cornfield tomato) 

 are cultivated frequently in Guatemala for its edible fruit. Large 

 quantities of the fruits are also gathered from spontaneous plants 

 that occur as weeds in vegetable gardens and cornfields. They are 




GENTRY & STANDLEY: FLORA OF GUATEMALA 93 



used mostly for flavoring food, just as tomatoes. Preserves and sim- 

 ilar sweet dishes are also prepared from them. 



Physalis porphyrophysa Donn.-Sm. Bot. Gaz. 61: 377. 1916. 

 In moist forests, 200-400 m.; Pete"n; Zacapa (type from the vi- 

 cinity of Zacapa, Pittier 1 754) Southern Mexico (Chiapas) . 



A shrub 0.5-6 m. tall, the young stems densely pubescent with short, antrorsely 

 curved and appressed, mostly simple hairs, glabrescent; leaves entire to slightly 

 undulate, lanceolate to elliptic, the larger leaves 5-15 cm. long, 2-7 cm. wide, the 

 apex acuminate, the base short attenuate, pubescence of simple, short and mostly 

 incurved hairs above, the hairs short and mostly incurved, simple and with a few 

 furcate and branched hairs among the simple ones below, the petioles 1-3 cm. long; 

 inflorescences consisting of (l-)2-6 flowers, often only one maturing a fruit, the 

 pedicels 5-10 mm. long, filiform, only slightly thickened above; flowering calyx 4-5 

 mm. long, pubescence of short, incurved hairs, the lobes triangular, about 2 mm. 

 long; corollas yellowish, 6.5-7.5 mm. long, the limb 8.5-9.5 mm. wide, maculate; 

 filaments 2.5-3 mm. long; anthers yellowish, about 3 mm. long; fruiting calyx 

 10-ribbed, 2.5-4 cm. long, 1.8-2.5 cm. wide, and strongly invaginated at the base, 

 sometimes purplish at maturity, pubescence of short incurved hairs, the pedicels 

 6-12 mm. long; berry 12-15 mm. in diameter, subsessile or on a short gynophore 

 1.5 mm. long or less. 



Some collectors have described the habit of Physalis porphyro- 

 physa Donn.-Sm. as subscandent. Part of the material cited by 

 Waterfall (1967) in P. melanocystis var. cernua has been included in 

 this species. 



Physalis porrecta Waterfall, Rhodora 69 : 237. 1967. Miltomate. 



Moist or wet thickets, sometimes moist dense forests, 1,550- 

 2,500 m., rarely lower; Chiquimula; Sacatepe"quez; Chimaltenango; 

 Quezaltenango; San Marcos. Mexico. Costa Rica. 



Herb 1.5 m. tall or less, the stems sparsely pubescent with short, multicellular 

 hairs, these usually confined to one side of the stem, glabrescent in age or some- 

 times more or less evenly pubescent, with some of the hairs capitate-glandular; 

 leaves mostly broadly dentate or entire, ovate to ovate-lanceolate, the larger 

 leaves mostly 5-12 cm. long, 4-7 cm. wide, the apex acuminate, the base obtuse 

 or acute, pubescent with multicellular hairs above and below, more sparsely 

 pubescent below, the petioles (2-)3-7 cm. long; flowers solitary, the pedicels 4-5 

 mm. long; flowering calyx 5-7 mm. long, pubescent with multicellular hairs, the 

 lobes lanceolate, 2.5-4 mm. long; corolla pale yellow or greenish-yellow, 7-10 mm. 

 long, the limb 12-15 mm. wide, maculate, these usually inconspicuous, edges of 

 the maculations often diffuse; filaments 2-4 mm. long; anthers yellow to bluish or 

 greenish-blue, about 2 mm. long; fruiting calyx 5-angled, (3-)3.5-5.5 cm. long, 2-3 

 cm. wide, glabrous, reticulate, usually abruptly beaked, the pedicels 10-16 mm. 

 long; berry 12-15 mm. in diameter. 




94 FIELDIANA: BOTANY, VOLUME 24 



Physalis pubescens L. Sp. PI. 1: 183. 1753. Miltomate. 



Moist or wet thickets or fields, open rocky places, sometimes 

 along rivers, 30-1,000 m.; Pete"n; Alta Verapaz; Izabal; Zacapa; Baja 

 Verapaz; Guatemala; Chiquimula; Huehuetenango; Jutiapa; Santa 

 Rosa. British Honduras. Eastern United States. Mexico to Costa 

 Rica. Panama and West Indies. South America. A widespread 

 pantropical species. 



Herb 2 m. or less tall, the stems pubescent with relatively long, multicellular 

 hairs, these sometimes glandular, sometimes glabrate; leaves coarsely sinuate- 

 dentate to entire, ovate, the larger leaves 6-16 cm. long, 2-8 cm. wide, the apex 

 acuminate, the base rounded to subcordate, truncate or acute, pubescent with 

 spreading, multicellular hairs above and below, the petioles 2-7 (-10) cm. long; 

 flowers solitary, the pedicels 3-6 mm. long; flowering calyx 3.5-6 mm. long, densely 

 pubescent with spreading hairs, the lobes narrowly lanceolate or triangular, rarely 

 ovate-deltoid, 1.5-3.5 mm. long; corolla yellowish, 6-12 mm. long, the limb macu- 

 late, 8-15 mm. wide, pubescent internally; filaments 2-3 mm. long; anthers bluish 

 or violet, 1.5-3 mm. long; fruiting calyx 5-angled, reticulate, 2-4 cm. long, 1.2-3 

 cm. wide, pubescent with spreading hairs, the pedicels 5-15 mm. long; berry 10- 

 18 mm. in diameter, sessile or on a gynophore 1-2 mm. long, often capitate- 

 glandular. 



Some of the specimens placed here have been identified as Phy- 

 salis turbinata Medic, by Waterfall. 



SCHWENCKIA L. 



Herbs; leaves alternate, entire, short petiolate; inflorescence paniculate or 

 racemose, bracteate or bracts absent, the flowers small; calyx tubular, 5-lobed; 

 corolla tubular, reddish-purple or purplish-green, the lobes somewhat 3-lobulate 

 with the median lobule clavate, the 2 shorter lateral lobules fused completely or 

 almost so with the adjoining lobes of each adjacent corolla lobe, thus the corolla 

 appearing to be 10-lobed but actually 5-lobed; fertile stamens 2, included, with 3 

 or 2 staminodes; anthers coherent, longitudinally dehiscent; filaments dilated 

 below and ciliate; ovary bilocular, the ovules numerous, the disk cupular; style 

 smooth and slender, included or exserted; stigma capitate; capsule septicidal, 

 smooth and glabrous; seeds numerous and angulate; embryo straight. 



About 20 species, only one additional species in Central America, 

 the others chiefly Brazilian. 



Schwenckia americana L. Syst. Veg. 60. 1774. 



Dry rocky open hillsides or open thickets, sometimes in brushy 

 pine forests, near sea level to 1,500 m., sometimes higher; Zacapa; 

 Jalapa; Jutiapa; Chiquimula. British Honduras. Southern Mexico. 

 Honduras; El Salvador; Costa Rica. Panama. Cuba and South 

 America. Africa. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 95 



Slender annual plants, 2.5-6 dm. tall, stem often densely pubescent with 

 inwardly curved hairs; leaves oblong to linear-lanceolate or elliptic, mostly 2-3.5 

 cm. long, 0.5-2 cm. wide, reduced upwards, hirsutulous, the lateral veins con- 

 spicuous in the larger leaves, acute, attenuate at the base; petioles 0.7 mm. long 

 or less; flowers on filiform pedicels about as long as the calyx; calyx about 4.5 mm. 

 long, sparsely pubescent or glabrous, the lobes 1-1.5 mm. long, acute; corolla 

 reddish-purple or dark purple and greenish, 10-13 mm. long, the tube slender, 

 about 1 mm. wide; filaments about 7.5 mm. long, pubescent below, the staminodes 

 3 mm. long; style 10.5-11 mm. long; capsule 3.5-4 mm. long, obtuse, slightly 

 exceeding the accrescent calyx; seeds angulate, rugulose and somewhat muricate, 

 less than 1 mm. long. 



There is some tendency for the plants in British Honduras to 

 have narrower leaves than in typical Schwenckia americana. The 

 name S. americana var. angustifolia Schmidt is available for this 

 narrow-leaved form of the species. 



See additional comments under Melananthus. 



SOLANDRA Swartz, nom. cons. 



Scandent shrubs, sometimes epiphytic, glabrous or pubescent, the hairs 

 simple or branched; leaves alternate and entire, petiolate, subcoriaceous or mem- 

 branaceous; inflorescences pseudoterminal, consisting of a single flower, the flowers 

 very large and showy; calyx tubular or campanulate, 2-5 lobed or parted; corolla 

 funnelform, white or yellowish, the limb 5-lobed, the lobes imbricate in bud, the 

 tube widely ampliate above; stamens 5, inserted at the point where the corolla 

 tube widens, exserted; filaments glabrous or pubescent at their bases, the staminal 

 traces villous; anthers oblong, longitudinally dehiscent; ovary partly inferior, 

 tetralocular, the ovules numerous, the disc inconspicuous; style filiform, exserted; 

 stigma capitate and dilated; fruit a large, leathery berry; seeds numerous; embryo 

 strongly curved. 



A genus of seven or eight species, all in tropical America. Two or 

 three more species may occur in southern Central America. 



Solandra grandiflora Swartz, Vet. Akad. Handl. Stockh. 8: 

 300, t. 11. 1787. Ingerto de montana (Guatemala). 



Wet or damp forests or on cliffs, 1,170-3,000 m.; Alta Verapaz; 

 Guatemala; Chimaltenango; San Marcos; Suchite"pequez. Southern 

 Mexico. El Salvador; Honduras; Costa Rica. Panama and West 

 Indies. 



An epiphytic or sometimes terrestrial scandent shrub, the branches thick, 

 glabrous; leaves subcoriaceous, lustrous, broadly elliptic to elliptic-ovate or 

 obovate, 10-22 cm. long, 4.5-10 cm. wide, glabrous above, glabrous below or 

 pubescent along the edge of the mid-vein or with tufts of hairs in the axis of the 

 mid-vein below, apex acuminate or sometimes acute, short attenuate or acute at 

 the base; petioles 3.5-9 cm. long, glabrous; inflorescence pseudoterminal, consisting 




FIG. 18. Solandra grandiflora. Habit and flower, X 



96 




GENTRY & STANDLEY: FLORA OF GUATEMALA 97 



of a solitary flower; pedicels thick, 0.8-2 cm. long, glabrous; calyx 6.5-8 cm. long, 

 glabrous, 5-angled, slightly lobed to parted nearly to the base, the lobes usually 

 unequal, the longer ones oblong, apex short acuminate; corolla yellowish, 19-25 cm. 

 long, the tube 16-19.5 cm. long, the lobes subrotund to ovate-rotund, 3.5-5 cm. 

 long, erose and entire, apex rounded; stamens inserted near the middle of the tube, 

 exserted; filaments 9-11 cm. long, glabrous, curved in the upper portion holding 

 the anthers suberect; anthers 10-12 mm. long; style exserted, 18-21 mm. long; 

 ovary tetralocular; fruit ovate-globose, 4.5-5 cm. long, 4-5 cm. wide at the base, 

 acute; seeds numerous, 5 mm. long. 



We are recognizing only one species in Guatemala. However, 

 further collecting and study may warrant the recognition of one or 

 possibly two additional species. The material placed here has been 

 identified as Solandra nitida Zucc. in Roem. in herbaria. 



Solandra grandiflora is planted often as an ornamental in Guate- 

 mala. The large fruits, often found in great abundance on the 

 ground, are said to be edible. 



Called copa de oro in Honduras. 



SOLANUM L. 



References: M. F. Dunal in A. DC. Prodr. 13, pt. 1, Solanum, 

 pp. 27-387. 1852. D. S. Correll, Section Tuberarium of the Genus 

 Solanum of North America and Central America, Agr. Monogr. 

 U.S.D.A. No. 11, 1-243. 1952. C. B. Heiser, Jr., The Solanum 

 nigrum complex in Costa Rica, Ceiba 4: 293-299. 1955. D. S. 

 Correll, The Potato and its Wild Relatives, Texas Research Founda- 

 tion, 606 pp. 1962. J. G. Hawkes, A revision of the tuber-bearing 

 Solanums, 2 ed., Scottish Plant Breed. Sta. Rec. 76-181. 1963. 

 K. P. Roe, A revision of Solanum sect. Brevantherum (Solanaceae) in 

 North and Central America, Brittonia 19: 353-373. 1967; A revision 

 of Solanum section Brevantherum (Solanaceae) I.e. 24: 239-278. 1972. 

 J. Edmonds (ne'e Gray), A synopsis of the taxonomy of Solanum 

 Sect. Solanum (Maurella) in South America, Kew Bull. 27: 95-114. 

 1972. 



Herbs, shrubs, small trees, or vines, armed with prickles or unarmed ; glabrous 

 or pubescent, the hairs often variously stellate or branched; leaves alternate, 

 sometimes in pairs, simple and entire to parted or odd-pinnate; inflorescences 

 simple or compound cymes, sometimes umbelliform or racemose, these pseudo- 

 terminal, lateral and opposite the leaves, or internodal, or axillary; calyx cam- 

 panulate, 5-lobed; corolla subrotate, white, yellow, violaceous, or purple, the lobes 

 plicate or valvate in bud; stamens 5, inserted near the base of the corolla tube; 

 filaments filiform or broad; anthers oblong or attenuate, dehiscent by two terminal 

 pores or slits, these often continued as longitudinal introrse slits part way down 

 the thecae; ovary bilocular, the ovules numerous, the disc inconspicuous; stigma 




98 FIELDIANA: BOTANY, VOLUME 24 



capitate, entire or obscurely bilobed; fruit a berry, usually globose and fleshy, 

 sometimes ovoid or ellipsoid; seeds numerous, compressed, orbicular or subreni- 

 form; embryo strongly curved or spiral. 



A genus of some 1,500 species comprising over one-half of the 

 species in the family, chiefly in tropical and subtropical regions in 

 both hemispheres. There are other numerous species known from 

 Mexico and southern Central America. An economically important 

 genus because it includes all of the cultivated potatoes. Solatium 

 also contains some of the most poisonous members of the family, 

 however, a few species produce edible fruits. 



Solanum is one of the largest genera of plants, as well as one of 

 the most difficult taxonomically. The large number of species 

 partly explains some of the problem. The genus is in need of a tax- 

 onomic revision and would provide a lifetime work for some indus- 

 trious person. The total number of names to consider in such a re- 

 vision is over 3,000. 



The treatments cited above by Correll and Roe have been most 

 helpful in the preparation of the manuscript. We have followed their 

 work with some modifications. 



Anthers long attenuate, the pores small; plants nearly always armed with prickles. 

 Plants vines or rarely erect shrubs; branches armed with mostly short and 

 strongly recurved prickles. 



Leaves glabrous or inconspicuously puberulent, the hairs simple. 



Stamens equal in length; leaves entire S. cobanense. 



Stamens unequal in length; at least the lower leaves pinnatifid or odd- 

 pinnate. 



Corolla blue to lavender, the limb 3.5-5.5 cm. wide, scarcely lobed to 

 nearly entire; branches, leaves, and inflorescences nearly always glab- 

 rous S. wendlandii. 



Corolla white, the limb less than 2 cm. wide, lobed to below the middle; 

 branches, leaves, and inflorescences pubescent, the hairs very short. 



S. molinarum. 

 Leaves stellate-pubescent, the hairs sometimes sparse and inconspicuous. 



Leaves sessile or subsessile, broadly cuneate at their bases, covered with 



short, stipitate-stellate hairs above; plants shrubs S. jamaicense. 



Leaves petiolate, acute to short attenuate at their bases, covered with 

 sessile-stellate hairs above, sometimes sparsely pubescent; plants vines 

 or sometimes shrubs. 



Young branches densely covered with reddish hairs, the hairs mostly 

 with the central ray greatly exceeding the lateral ones; long, stipitate- 

 stellate (bristle-like) hairs absent; calyx lobes 5 mm. long or more, 

 lanceolate to lance-oblong, pubescent internally; plants shrubs. 



S. erythrotrichum. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 99 



Young branches sparsely to densely covered with yellowish or yellowish- 

 brown hairs, the hairs with the rays subequal; long, stipitate- 

 stellate (bristle-like) hairs often present on the branches; calyx lobes 

 3 mm. long or less, linear-oblong or caudate, glabrous internally; 

 plants vines. 



Leaves entire to shallowly angulately-lobed ; stems mostly densely 

 stellate-pubescent, often covered with long, stipitate-stellate 

 (bristle-like) hairs; anthers 6-11 mm. long; ovary and style 



pubescent S. lanceifolium. 



Leaves sinuately-lobed to deeply parted; stems glabrate, bearing a few 

 stellate hairs when young, long, stipitate-stellate (bristle-like) 

 hairs absent; anthers 3.5-4 mm. long; ovary and style glabrous. 



S. houstonii. 



Plants erect shrubs or small trees, sometimes herbs; branches armed with 

 straight, often long prickles, rarely a few prickles slightly recurved or the 

 branches sometimes unarmed. 

 Leaves with simple hairs above. 



Leaves with simple hairs above and below; fruit ovoid and abruptly con- 

 tracted at the apex into a short neck (mammillate) . . . . S. mammosum. 

 Leaves with at least some stellate hairs below; fruit globose. 



Stems and petioles with at least some of the long, eglandular hairs 1.5-2 



mm. in length; seeds with a prominent wing S. quinquangulare. 



Stems and petioles with short and mostly glandular hairs, the hairs 

 nearly always less than 1 mm. in length; seeds without a wing. 



S. globiferum. 

 Leaves with stellate hairs above. 



Plants herbs, 1 m. tall or less; calyx densely armed with prickles. 



Leaves shallowly lobed to deeply parted, densely pubescent above and 

 below; corolla purplish-white; stamens subequal in length; fruit 



not enclosed by the calyx S. campechiense. 



Leaves bipinnatifid, sparsely pubescent to nearly glabrous above; corolla 

 yellow; stamens unequal in length; fruit enclosed by the close fitting 



calyx S. cornutum. 



Plants shrubs or sometimes small trees, (0.5-) 1-5 m. tall; calyx unarmed. 

 Branches covered with reddish hairs. 



Branches sessile-stellate pubescent, the hairs mostly with the central 

 ray greatly exceeding the lateral ones; ovary densely stellate- 

 pubescent throughout S. erythrotrichum. 



Branches covered with long, stipitate-stellate (bristle-like) hairs, the 

 rays subequal; ovary sparsely stipitate-glandular pubescent 



apically S. hispidum. 



Branches covered with whitish, grayish or sometimes yellowish-brown 



hairs. 

 Fruit densely stellate-pubescent, the hairs very conspicuous; calyx 



lobes densely stellate-pubescent internally; anthers subsessile. 

 Branches covered with yellowish-brown hairs, at least some of the 

 hairs long stipitate-stellate; leaf lobes mostly 5-7; prickles 

 commonly with broad bases S. tequilense. 




100 FIELDIANA: BOTANY, VOLUME 24 



Branches covered with whitish or grayish, sessile-stellate hairs, at 

 least some of the hairs glandular; leaf lobes mostly 9 or more; 



prickles commonly with slender bases S. hirtum. 



Fruit glabrous; calyx lobes glabrous internally or sparsely pubescent 



apically; anthers with filaments 1 mm. long or more. 

 Corolla purplish; branches long, stipitate-stellate pubescent, the 



hairs whitish or grayish; montane species S. hartwegii. 



Corolla white; branches sessile-and/or short stipitate-stellate pubes- 

 cent, the hairs mostly yellowish-brown; lowland species. 

 Leaves entire; plants unarmed; pedicels and calyces eglandular; 

 fruit lustrous, 5-7.5 mm. in diameter; inflorescences consisting 



of bifurcate cymes S. blodgettii. 



Leaves sinuately lobed to deeply parted, sometimes subentire; 

 plants nearly always armed with prickles; pedicels and calyces 

 often glandular; fruit dull, 10-14 mm. in diameter; inflores- 

 cences consisting of mostly simple or furcate cymes . . S. torvum. 

 Anthers short, oblong or ellipsoid, the pores large; plants always unarmed. 

 Leaves simple. 



Pubescence of entirely simple hairs or the plants glabrous. 

 Plants herbaceous (rarely suffrutescent), not scandent. 



Corolla limb only shallowly lobed S. muricatum. 



Corolla limb parted to near the base. 



Pedicels articulate at or near the middle; pseudostipules present at the 



base of the petioles, rarely absent in S. clarum. 

 Anthers rounded and dorsally auriculate at their bases; filaments 

 often connate forming a column; plants nearly always epiphytic. 



S. morelliforme. 

 Anthers cordate or dorsally lobulate at their bases; filaments free or 



somewhat connate; plants terrestrial. 



Leaves truncate or cordate at their bases; anthers dorsally 3- to 

 5-lobulate at their bases; pseudostipular leaves ovate or ab- 

 sent S. clarum. 



Leaves shortly attenuate at their bases; anthers dorsally cordate 

 at their bases; pseudostipular leaves auriform. 



S. bulbocastanum. 

 Pedicels articulate at their bases; pseudostipules absent. 



Calyx densely hirsute, the calyx lobes much accrescent in fruit, 

 one-half to as long as the fruit; inflorescences sessile or nearly 



so; filaments and style glabrous S. deflexum. 



Calyx pilosulous, the calyx lobes only slightly accrescent in fruit; 



inflorescences long-pedunculate; filaments and style pubescent. 



Anthers 1.5-2 mm. long; corolla limb 5-7.5 mm. wide; sepals 



reflexed in fruit; lowland species, rarely found above 1,500 m. 



S, americanum. 



Anthers (2.5-)3-4 mm. long; corolla limb 6.5-10(-16) mm. wide; 

 sepals spreading to slightly ascending in fruit; ranging from 



1,500-3,900 m S. nigrescens. 



Plants shrubs or small trees, rarely scandent. 



Leaves with the hairs in tufts in the axils of the mid-vein below, some- 

 times the tufts of hairs inconspicuous S. nudum. 



Leaves glabrous or pubescent below but the hairs not in tufts in the axils 



of the mid-vein below. 



Young branches, leaves, peduncles, and pedicels densely covered 

 with a whitish or yellowish closely appressed tomentum, the 

 nature of the hairs discernable only with difficulty. . .S. nigricans. 

 Young branches and leaves glabrous or pubescent, not densely covered 

 with an appressed tomentum, the nature of the hairs easily 

 discernable. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 101 



Corolla violet, the limb 24-30 mm. wide; anthers 5-6 mm. long, 

 broad; pedicels articulate at their bases to just above the 



middle; scandent shrubs S. dulcamaroides. 



Corolla white, the limb 6-15 mm. wide; anthers 1.5-4 mm. long, 



slender; pedicels articulate at their bases; erect shrubs. 

 Filaments free or only slightly connate at their bases, less than 

 0.5 mm. long; leaves 3-9 cm. long, 1-2.5 cm. wide, the apex 



obtuse or sometimes acute S. diphyllum. 



Filaments connate one-half their entire length or more, 0.5-1 mm. 

 long; leaves mostly 10-27 cm. long, 2.5-12 cm. wide, the apex 

 acuminate or acute. 



Inflorescences inconspicuously puberulent throughout; pedicels 

 5-7 mm. long, puberulent; leaves broadest mostly above 

 the middle; principal lateral veins of the largest leaves 6-8 



in number; plants below 1,000 m S. rovirosanum. 



Inflorescences glabrous throughout; pedicels 10-15 mm. long, 

 glabrous; leaves broadest mostly at or below the middle; 

 principal lateral veins of the largest leaves 9-17 in number. 

 Corolla limb 15 mm. wide; anthers 3-4 mm. long; inflores- 

 cences mostly furcate; plants above 2,000 m... fontium. 

 Corolla limb 7-8 mm. wide; anthers 2-2.5 mm. long; in- 

 florescences simple; plants below 1,000 m. 



S. tuerckheimii. 



Pubescence of stellate (the rays radiating from a central point) or branched 

 (rays along their stalks or dichotomously branched) hairs; simple hairs 

 occasionally present in addition to the compound ones. 



Leaves with hairs in tufts in the axils of the mid-vein below or the hairs 

 confined to the edge of the mid-vein and the bases of the lateral veins, 

 sometimes the hairs generally dispersed but not so on the mid-vein, 

 sometimes sparsely pubescent. 



Leaves with hairs in mostly conspicuous tufts in the axils of the mid- 

 vein below; calyx lobes rounded and short-caudate; corolla limb 

 12.5-15 mm. wide S. aligerum. 



Leaves with the hairs mostly along the edge of the mid-vein below, the 

 hairs not in conspicuous tufts in the axils of the mid- vein; calyx 

 lobes triangular or oblong; corolla limb 8-12 mm. wide. 



S. cervantesti. 



Leaves evenly pubescent below and sometimes more densely so on the veins 

 but the hairs not confined to the edge of the mid-vein nor as tufts in 

 the axils of the mid-veins. 



Receptacle conspicuously swollen in bud and flower, often discernable in 



fruit S. hazenii. 



Receptacle not swollen. 



Leaf bases long attenuate, decurrent along the entire length of the 

 petiole or nearly so, sometimes decurrent only one-half the length 

 of the petioles. 



Stellate hairs of the young stems, petioles and peduncles at least 

 partly with many rays, usually greatly exceeding 12, the hairs 

 often with rays along their stalks; leaves mostly 8-15 cm. wide. 



Pedicels very short, rarely more than 1 mm. long; leaves at least 

 2 dm. long, 1 dm. wide; leaves with the apices acuminate. 



S. brevipedicellatum. 



Pedicels 2 mm. long, usually longer; leaves often smaller than above, 

 3-11 cm. wide; leaves with the apices acute or acuminate. 



S. atitlanum. 



Stellate hairs of the young stems, petioles and peduncles with few 

 rays, usually less than 12; leaves mostly 2-8 cm. wide. 




102 FIELDIANA: BOTANY, VOLUME 24 



Stems, peduncles, pedicels, and calyces with partly long stipitate- 

 stellate hairs; calyx visibly glandular at 10 X magnification. 



S. umbellatum. 



Stems, peduncles, pedicels, and calyces with sessile or short, 

 stipitate-stellate hairs; calyx not visibly glandular at 10 X 

 magnification. 



Leaves 2-3 cm. wide, the petioles 2-5 mm. long. . . .S. asperum. 

 Leaves 4.5-9 cm. wide, the petioles mostly 10-15 mm. long. 



S. rugosum. 

 Leaf bases rounded or obtuse to nearly truncate or attenuate, rarely 



decurrent to one-half the length of the petioles. 

 Young leaves, peduncles, and pedicels densely covered with whitish 

 or yellowish closely appressed, branched hairs, the branching of 



the hairs discernable only with difficulty S, nigricans. 



Young leaves, peduncles, and pedicels sparsely to densely covered 

 with stellate (rays radiating from a central point) or branched 

 (rays along their stalks) hairs, the nature of the hairs easily 

 discernable. 



Leaves densely covered with minute and closely appressed stellate- 

 like scales below, the nature of the hairs discernable only 

 with difficulty at 10 X magnification, the surface smooth to 



the touch S. lepidotum. 



Leaves covered with stellate hairs below, the nature of the hairs 

 easily discernable with the naked eye, the surface velvety to 

 slightly asperous to the touch. 



Pubescence almost entirely of branched hairs; principal lateral 

 veins of the larger leaves 20 or more; corolla purple to 



lavender S. muenscheri. 



Pubescence mostly of stellate hairs, sometimes a few branched 



hairs among the stellate ones; principal lateral veins of 



the larger leaves 5-8 (-11); corolla white, rarely lavender. 



Inflorescences subsessile, the peduncle 0.3 cm. long or the 



inflorescences sessile. 



Leaves with long acuminate apices, the upper surface with 

 at least some of the stellate hairs with the central ray 

 much longer than the lateral rays; calyx lobes much 

 accrescent in fruit, nearly as long as to slightly ex- 

 ceeding the mature fruit, the lobes stellate-pubescent 



internally S. cordovense. 



Leaves with acute apices, the upper surface with simple 

 and stellate hairs, the rays subequal in length; calyx 

 lobes slightly accrescent in fruit, only about one-third 

 the length of the mature fruit, the lobes glabrous 



internally S. huehuetecum. 



Inflorescences pedunculate, the peduncles 1.5-12 cm. long. 

 Petioles short, 0.2-0.5 cm. long; inflorescences lateral, 

 borne below the foliage on divergent peduncles. 



S. schlechtendalianum. 

 Petioles long, 1-10 cm. long; inflorescences pseudoterminal, 



borne above the foliage on erect peduncles. 

 Leaf bases rounded to nearly truncate or acute; flower 



buds turbinoid S. erianthum. 



Leaf bases long or short attenuate, the blade decurrent 

 on the petiole; flower buds subglobose to oblong. 



S. chiapasense. 

 Leaves pinnate or very rarely simple in part in S. seaforthianum. 



Inflorescences axillary S. trizygum. 



Inflorescences pseudoterminal, or lateral and internodal, or opposite the leaves. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 103 



Pedicels articulate at or near the middle. 



Terminal leaflets obovate or broadly elliptic, much larger than the lateral 



ones, the apices abruptly acuminate or acute; fruit globose or ovoid, 



rounded at the apex; plants often forming a rosette. . .S. demissum. 



Terminal leaflets lanceolate to narrowly elliptic or lanceolate-oblong, 



about the same size as the lateral ones, the apices long acuminate or 



narrowly acute; fruit ellipsoid or ovoid-cylindrical, acute at the 



apex, plants not forming rosettes. 



Leaves (at least some) with numerous and prominent interstitial 



leaflets S. agrimonifolium. 



Leaves without interstitial leaflets or rarely present, then few in num- 

 ber, minute and inconspicuous S. oxycarpum. 



Pedicels articulate at or near their bases. 

 Corolla limb scarcely lobed. 



Plants with spreading hairs; leaves without interstitial leaflets, the 

 terminal leaflet usually much larger than the lateral ones. 



S. fraxinifolium. 



Plants with appressed hairs; leaves with few interstitial leaflets, the 

 terminal leaflet usually not much larger than the lateral ones. 



S. suaveolens. 

 Corolla limb parted more than one-half way to the base. 



Leaflets always 3 S. phaseoloides. 



Leaflets 5-15, rarely a few of the leaves with 3 leaflets or the leaves 



simple in part. 

 Leaves with interstitial leaflets; stem herbaceous, not rooting at 



the nodes S. canense. 



Leaves without interstitial leaflets; stem woody, climbing and 

 rooting at the nodes, rarely without roots at the nodes as in 

 S. seaforthianum. 



Corolla limb 18-22 mm. wide, the flowers showy; stems not rooting 



at the nodes; pseudostipular leaves absent. .S. seaforthianum. 



Corolla limb 6-13 mm. wide, the flowers not showy; stems rooting 



at the nodes; pseudostipular leaves nearly always present. 

 Anthers apiculate to aristate and puberulent at their apices; 



leaflets nearly always 7-11 S. skutchii. 



Anthers rounded and retuse and glabrous at their apices; 



leaflets 5, rarely 3. 

 Style much exceeding the stamens. 



Filaments connate their entire length; young stems densely 



pubescent S. connatum. 



Filaments connate not more than one-half their length; 

 young stems mostly glabrous to sparsely pubescent. 



S. inscendens. 

 Style about equal to or shorter than the stamens. 



Peduncle and rachis sparsely pubescent to essentially 

 glabrous; pedicels with appressed hairs or glabrous. 



S. appendiculatum. 



Peduncle and rachis densely pubescent; pedicels with 

 spreading or subappressed hairs S. lacanense. 



Solatium agrimonifolium Rydb. Bull. Torrey Bot. Club 51: 

 154. 1924. Tisbotch; papa de marrana. 



Wet forests or cloud forests, alpine thickets, shaded barrancos or 

 stream banks, 1,900-3,400 m.; Quezaltenango ; San Marcos; Hue- 

 huetenango. Southern Mexico. 




104 FIELDIANA: BOTANY, VOLUME 24 



Plants herbaceous erect or ascending, 2. 5-12 dm. tall, nearly glabrous to densely 

 pubescent, the stem angular, simple or branched, often purple-tinged, non-tuber- 

 bearing or sometimes with small ellipsoid tubers; leaves odd-pinnate, 1-3.5 dm. 

 long; interstitial leaflets usually numerous, sessile; leaflets 9-1 5, elliptic-lanceolate 

 to narrowly lanceolate, 1.5-12 cm. long, 0.5-3.5 cm. wide, the apex narrowly acute to 

 long acuminate, rarely obtuse, the base oblique, rounded to cuneate, sessile or with 

 a short winged petiolule, the lowermost pair of leaflets much reduced; pseudo- 

 stipular leaves present; inflorescence pseudoterminal, becoming lateral and opposite 

 the leaves, few-to many-flowered, laxly cymose or sometimes racemose; peduncles 

 3-10.5 cm. long; pedicels 1-3.5 cm. long, articulate at or somewhat above the 

 middle; calyx 6-12 mm. long, glabrous or pubescent, parted to well below the 

 middle, the lobes ovate-lanceolate to lanceolate, acute or long acuminate; corolla 

 bright purple to lavender or rarely whitish, the limb mostly 2-2.5 cm. wide, very 

 shallowly lobed, the lobes 1.5-3 mm. long, puberulent; filaments 1-2 mm. long, 

 glabrous or sometimes pubescent; anthers 3.5-5.5 mm. long; style 7-8 mm. long, 

 exceeding the stamens; fruit ellipsoid-conic, acute, pale green, 2.5-4 cm. long, up 

 to 1.3 cm. in diameter near the base. 



Solanum aligerum Schlecht. Linnaea. 19: 301. 1845. Seconillo 

 (Quezaltenango). 



Wet cloud forests or in mixed forests or in moist sandy forests, 

 2,400-3,100 m.; Quiche"; Chimaltenango; Huehuetenango; Que- 

 zaltenango. Southern Mexico. El Salvador and Honduras. 



An unarmed shrub or tree, 1.5-10 m. tall, the branches glabrous; leaves 

 oblong-elliptic to narrowly elliptic, sometimes lanceolate, mostly 7.5-20 cm. long, 

 2-4 cm. wide, the apex acuminate or narrowly acute, the base attenuate or some- 

 times long attenuate, glabrous or sometimes pilosulous above, the hairs in tufts 

 in the axils of the mid-vein and dispersed along the edge of the mid-vein and 

 sometimes near the mid-vein below, the hairs branched, sometimes simple hairs 

 present; petioles 1-1.5 cm. long; inflorescence pseudoterminal, becoming lateral 

 and opposite the leaves, cymose, more or less elongate or broader at the base, 

 few-to many-flowered; peduncles short, 1-3 cm. long; pedicels 8-14 mm, long; 

 calyx 2-2.5 mm. long, sometimes cleft down one side, the lobes short, 0.5-1 mm. 

 long, rounded and short-caudate, ciliate and internally pilosulous and glandular; 

 corolla white, the limb 12.5-15 mm. wide, parted to below the middle, the lobes 

 5-6 mm. long, ciliate and apically pilosulous; filaments about 0.5 mm. long; 

 anthers 3-3.5 mm. long; style 6-6.5 mm. long, glabrous; ovary glabrous; fruit 

 globose, 7-10 mm. in diameter, black; seeds 3 mm. long. 



Solanum aligerum Schlecht. and S. cervantesii Lagasca are closely 

 related. They are distinguished by the pubescence on the adaxial 

 side of the leaves, calyx lobes, and, in general, by the inflorescence. 

 The corollas of S. aligerum are somewhat broader than in S. cer- 

 vantesii. 



Solanum americanum Miller, Gard. Diet. ed. 8. Solanum no. 

 5. 1768. S. nodiflorum Jacq. Icon. PI. Rar. 2: 288. 1786. Hierba 




GENTRY & STANDLEY: FLORA OF GUATEMALA 105 



mora (Chimaltenango; Jutiapa) ; macuy (Alta Verapaz); quilete (Santa 

 Rosa). 



Damp thickets or forests, open hillsides or field, a common weed 

 of cultivated and waste ground, 350-1,500 m., rarely higher; Pete"n; 

 Alta Verapaz; Zacapa; Baja Verapaz; Sacatepe"quez; Chimlatenango; 

 Huehuetenango; Jutiapa; Santa Rosa; Escuintla; Retalhuleu; San 

 Marcos. British Honduras. Western United States. Mexico to 

 Costa Rica. Panama and West Indies. South America. 



A perennial or annual herb, erect or decumbent, 1 m. tall or less, the young 

 stems pilosulous to nearly glabrous, the hairs incurved; leaves in pairs or solitary, 

 different in size, similar in shape, entire or sinuate-dentate, lanceolate to ovate, 

 the larger leaves 3.5-14 cm. long, 1.5-5.5 cm. wide, the apex narrowly acute or 

 acuminate, the base attenuate, sparsely to densely pilosulous or glabrous above 

 and below; petioles 5-30 mm. long; inflorescences lateral and internodal, sub- 

 umbelliform or racemiform, few-to several-flowered; peduncles 5-25 mm. long; 

 pedicels 5-10 mm. long, fruiting pedicels reflexed; calyx 1-2 mm. long, lobed to the 

 middle, the lobes ovate to oblong, acute or obtuse, reflexed in fruit; corolla white, 

 the limb 5-7.5 mm. wide, parted to near the base, the lobes 2-3 mm. long, externally 

 papillate; filaments 0.3-0.5 mm. long, ciliate; anthers 1.5-2 mm. long; style 

 2.5-3.5 mm. long, exceeding the stamens, the lower one-half densely pubescent; 

 ovary glabrous; fruit globose, 4-8 mm. in diameter, black at maturity; seeds about 

 1 mm. long. 



Solanum americanum Miller and S. nigrescens Mart. & Gal. are 

 members of section Morella (Maurella Nees). The species in this 

 section are morphologically similar and very difficult to distinguish. 

 As a result, authors vary widely in their treatments of some members 

 of this section. 



All of the material included in this species has been identified as 

 either S. nodiflorum Jacq. or S. nigrum L. We are following Edmonds 

 (ne'e Gray) in placing S. nodiflorum in the synonymy of S. american- 

 um. Solanum nigrum L., sensu stricto, is not known to occur in 

 Guatemala, and probably not anywhere in Central America. Ed- 

 monds (1972) did not find any specimens in South America that could 

 be referred to S. nigrum L., sensu stricto. See Edmonds (J. Arnold 

 Arbor. 52: 635. 1972) for the differences separating S. americanum 

 from S. nigrum L., sensu stricto. 



In Guatemala the young foliage is used as one of the common 

 pot herbs and is consumed in large quantities. It is found in most 

 of the markets. 



Called bocano in British Honduras and mora in Honduras and 

 El Salvador. See comments under S. nigrescens Mart. & Gal. 




106 FIELDIANA: BOTANY, VOLUME 24 



Solatium appendiculatum H. & B. ex Dunal, Solan. Syn. 5. 

 1816. Mapix; tomatillo. 



Wet forests or cloud forests, in mixed or coniferous forests; 

 2,000-3,500 m., rarely lower; Alta Verapaz; Chiquimula; Jalapa; 

 Guatemala; Sacatepe'quez; Chimaltenango; Solola; Quezaltenango; 

 Totonicapan; Huehuetenango; Suchitepe"quez ; San Marcos. South- 

 central Mexico. 



A climbing or sometimes trailing woody vine, the stems rooting at the nodes, 

 non-tuber-bearing ; young branches densely pubescent to nearly glabrous, becoming 

 glabrous with age; leaves odd-pinnate, 4-10 (-15) cm. long, pubescent to essentially 

 glabrous above and below; petioles 1-3. 5 (-9) cm. long; interstitial leaflets absent; 

 leaflets 5, rarely 3, lanceolate to elliptic, rarely oblanceolate or ovate, mostly 

 1-5 cm. long, 1-2.5 cm. wide, the apex acute to acuminate, petiolule to 5 mm. long, 

 the lowermost pair of leaflets reduced, the terminal leaflet larger than the lateral 

 ones; pseudostipular leaves present; inflorescence pseudoterminal on leafy branches, 

 becoming lateral and opposite the leaves, cymose-paniculate; peduncle 5-20 mm. 

 long, sparsely pubescent to glabrous with age; pedicels 5-8 mm. long, glabrous to 

 sparsely pubescent with appressed hairs, articulate at their bases; calyx 2-2.5 mm. 

 long, pubescent to glabrous with age, divided to about the middle, the lobes 

 broadly ovate, acute or obtuse, apiculate, marginate; corolla white, rarely light 

 blue, the limb deeply lobed to near the base, the lobes 3-5 mm. long, elliptic- 

 lanceolate, obtuse to acute; filaments about 0.5 mm. long, pubescent; anthers 

 2-3 mm. long; style equal to or shorter than the stamens; fruit globose, red, about 

 1 cm. in diameter. 



Closely related to Solanum inscendens Rydb. 



We have followed Correll (1962) in recognizing five species of 

 Series Appendiculata as occurring in Guatemala, S. appendiculatum 

 H. & B. ex Dunal, S. connatum Correll, S. inscendens Rydb., S. 

 skutchii Correll, and S. tacanense Lundell. They are separated mostly 

 by minor morphological differences. Further collecting and study, 

 especially of S. connatum and S. tacanense, may result in a reap- 

 praisal of the present treatment. 



Solanum asperum L. C. Rich. Cat. PI. in Act. Soc. Hist. Nat. 

 Paris 1: 107. 1792. S. salmifolium Lam. Tabl. Encycl. Meth. 2: 14. 

 1793, non auct. 



Moist forests, in open places or in secondary growth, near sea 

 level; British Honduras. Panama. Trinidad, south to Brazil. 



An unarmed shrub or small tree 1-6 m. tall, the young branches ferruginous, 

 densely stellate-pubescent, the hairs sessile and short stipitate-stellate; leaves 

 narrowly elliptic or lanceolate, 10-17 cm. long, 2-3 cm. wide, the apex acuminate, 

 the base long attenuate, decurrent along entire length of the petiole or nearly so, 

 sparsely to moderately sessile-stellate pubescent and asperulous above, densely 

 pubescent below, the hairs ferruginous, sessile and short-stipitate stellate; petioles 

 short, 2-5 mm. long; axillary leaves absent; inflorescence cymose, pseudoterminal, 




GENTRY & STANDLEY: FLORA OF GUATEMALA 107 



becoming lateral, dichotomously branched, long pedunculate, the peduncle 4-13 

 cm. long; pedicels 2-6 mm. long, pubescent, the hairs sessile and short-stipitate 

 stellate; flower buds obovoid to ellipsoid; calyx lobed to the middle, the lobes 

 1.5-3 mm. long, glabrous or stellate-pubescent within, densely stellate-pubescent 

 externally; corolla white, the limb 1-1.2 cm. wide, parted to below the middle, 

 the lobes 4-5 mm. long; filaments 0.5-1.5 mm. long; anthers 2-3 mm. long; ovary 

 and lower half of style densely stellate-tomentose; fruit yellow, 8-10 mm. in 

 diameter, stellate-pubescent; seeds 1.5-2 mm. long. 



Solanum atitlanum Roe, Brittonia 19: 364. 1967. 



Dry thickets, edge of fields and barrancas, 1,000-2,000 m.; 

 Jalapa; Guatemala; Solola (type from 3 miles east of Panajachel, 

 Roe et al. 773); Sacatepe"quez ; Santa Rosa. Honduras. Nicaragua. 



An unarmed shrub or small tree, 1-10 m. tall, the young branches stellate- 

 pubescent, the hairs sessile and short stipitate-stellate, with many rays, often with 

 many rays along their stalks; leaves ovate to elliptic or obovate, mostly 8-30 long, 

 3-11 cm. long, the apex acute or acuminate, the base long attenuate nearly the en- 

 tire length of the petiole, asperulous to velutinous, the hairs mostly simple and 

 sessile or short stipitate-stellate and with few rays above, the hairs sessile or short 

 stipitate-stellate, rays few and many, whitish below; petioles stellate-pubescent, 

 the hairs sessile or short stipitate-stellate, the rays few; axillary leaves present 

 on recent growth of young plants; inflorescence cymose, pseudoterminal, becoming 

 lateral, dichotomously branched, long pedunculate, the peduncle 4-9 cm., the 

 hairs with few or many rays, some with rays along their stalks; pedicels 2-7 mm. 

 long, the hairs sessile and short stipitate-stellate, with few and many rays; flower 

 buds globose to ellipsoid; calyx lobed to the middle or slightly below, the lobes lance- 

 olate-ovate to ovate, 1.7-5 mm. long, stellate-pubescent within; corolla violet, the 

 limb 1-1.5 cm. wide, parted to below the middle, the lobes 4-6 mm. long; filaments 

 1 mm. long or less; anthers 2.5-3.7 mm. long; ovary stellate-tomentose; style stel- 

 late-tomentose; fruit 9-12 mm. in diameter, pubescence persistent; seeds 1.8-2.5 

 mm. long. 



Solanum blodgettii Chapm. Fl. South. U.S. 349. 1860. S. 

 decurtatum Standl. & Steyerm. Field Mus. Bot. 23: 20. 1943 (type 

 from Lago de Yaxha, Pete'n, Lundell 2012), syn. nov. 



Low forest; Pete'n. British Honduras. Southern Florida. Ba- 

 hamas. Southern Mexico. 



An unarmed shrub about 1 m. tall, the young branches, petioles, peduncles 

 and calyces densely stellate-pubescent, the hairs yellowish or grayish, sessile-to 

 short-stipitate; leaves entire, solitary, elliptic-oblong or oblong, 7-16.5 cm. long, 

 2.5-5.5 cm. wide, the apex acute or obtuse, sometimes acuminate, the base obtuse, 

 green and asperous, the hairs sessile or subsessile-stellate above, densely stellate- 

 tomentose and grayish below; petioles 1-2 cm. long; inflorescences pseudoterminal, 

 becoming lateral and internodal or opposite the leaves, dichotomously cymose, 

 several-to many-flowered; peduncles short, 1.5-3 cm. long; pedicels 4-6 mm. long; 

 calyx 2-2.5 mm. long, parted to the middle, the lobes ovate, narrowly acute or 




108 FIELDIANA: BOTANY, VOLUME 24 



mucronate; corolla white, the limb 12-16 mm. wide, divided to near the base, the 

 lobes 5-7.5 mm. long, densely stellate-pubescent externally; filaments 1 mm. long; 

 anthers 4.5-5.5 mm. long, linear, scarcely narrowed upward; style 6.5-8.5 mm. 

 long, glabrous; ovary glabrous; fruit globose, 5-7.5 mm. in diameter, red, lustrous; 

 seeds 2-2.5 mm. long. 



So Ian um brevipedicellatum Roe, Brittonia 19: 361. 1967. 

 Palito tabaco de montana (Quezaltenango) . 



Moist forests or in secondary growth thickets, 1,500-1,800 m.; 

 Chimaltenango; Solola; Quezaltenango (type from 4 km. north of 

 Santa Maria planta electrica, Roe et al. 741). Mexico (Chiapas). 



An unarmed shrub or small tree, 2-12 m. tall, the young branches stellate- 

 pubescent, the hairs sessile and short stipitate-stellate, mostly with many rays, an 

 occasional plant with some long stipitate-stellate hairs; leaves ovate to broadly 

 ovate, mostly 24-40 cm. long, 10-26 cm. wide, the apex acuminate, the base long 

 attenuate, asperulous and with simple, sessile or short stipitate-stellate hairs, some 

 of the hairs with many rays above, densely to sparsely pubescent below, the hairs 

 sessile and short stipitate-stellate; petioles 3.5-6.5 cm. long, the hairs with many 

 rays; axillary leaves usually present; inflorescence cymose, pseudoterminal, becom- 

 ing lateral, dichotomously branched, long pedunculate, the peduncles 5-13 cm. long, 

 the stellate hairs with many rays; pedicels very short, 1 mm. long or less, the 

 stellate hairs with many rays; flower buds elongated; calyx lobed to the middle or 

 nearly so, the lobes lanceolate-ovate, 1.2-2.5 mm. long, stellate-pubescent within; 

 corolla white or violet, the limb 1.5-2 cm. wide, parted to below the middle, the 

 lobes 6-8 mm. long; filaments 1.8-2 mm. long; anthers 3-4.5 mm. long; ovary 

 stellate-tomentose; style stellate-tomentose; fruit yellow, 10 mm. in diameter, 

 pubescence persistent; seeds 2-2.5 mm. long. 



Solatium bulbocastanum Dunal in Poir. Encycl. Suppl. 3: 749. 

 1814. 



Dry slopes and thickets, 1,350-2,300 m. 



Plants herbaceous slender or stout, usually grayish green, erect to spreading, 

 3-10 dm. tall, densely pubescent throughout with multicellular, erect-spreading hairs 

 to rarely subglabrous, stoloniferous and tuber-bearing, tubers globose to ellipsoid, 

 clear white to creamy white, 7 cm. long or less; stem simple or much branched; 

 leaves simple, broadly ovate to ovate-lanceolate or narrowly elliptic-lanceolate, 

 4-15 cm. long, 2-6 cm. wide, the apex obtuse to acute or shortly acuminate, the 

 base shortly attenuate; petioles 5 cm. long or less; pseudostipular leaves present; 

 inflorescence pseudoterminal, becoming lateral and opposite the leaves, cymose, 

 mostly few- to many-flowered; peduncles short, 3 cm. long or less; pedicels 5-10 

 mm. long, articulate at about or slightly below the middle; calyx 3-4.5 mm. long, 

 the lobes short, rarely to about 2.5 mm. long, obtuse to acute; corolla white to 

 light yellow or greenish-yellow, the limb 1.2-1.5 cm. wide, parted to well below 

 the middle, the lobes 5-7 mm. long, ovate-lanceolate or lanceolate, subobtuse or 

 acute, densely pilosulous at the margins; stamens equal, the filaments 1-2 mm. 

 long, often somewhat connate; anthers 4-6 mm. long, cordate at their bases; style 




GENTRY & STANDLEY: FLORA OF GUATEMALA 109 



7-11 mm. long, glabrous, exceeding the stamens, sometimes only slightly so; fruit 

 globose, greenish, about 1 cm. in diameter. 



Solanum bulbocastanum var. glabrum Correll, Agr. Monogr. 

 U.S.D.A. No. 11, 79, t. 53 & 5J>. 1952. 



Guatemala; Huehuetenango. South central Mexico. 



Variety glabrum differs from the typical variety in that the 

 pedicel above the articulation and the calyx are glabrous; the corolla 

 is usually less deeply lobed; and the corolla lobes are somewhat 

 broader. It is distinguished from variety partitum by the more shal- 

 lowly lobed corolla limb and the style evidently longer than the 

 stamens. 



Solanum bulbocastanum var. partitum Correll, Agr. Mono- 

 gr. U.S.D.A. No. 11, 83, t. 55. 1952. S. bulbocastanum subsp. partitum 

 Hawkes, Scottish Plant Breed. Sta. Rec. 95. 1963. Yerba mora. 



Baja Verapaz (type from Fatal, Tuerckheim 1 1 2316) ; Guatemala; 

 Solola; Huehuetenango. Mexico (Chiapas). 



Variety partitum is characterized by the deeply lobed corolla limb 

 cleft nearly to the base. Also, the pedicel above the articulation and 

 the calyx are glabrous. 



Solanum campechiense L. Sp. PL 187. 1753. S. guanicense 

 Urban, Symb. Ant. 1: 399. 1899. 



In damp thickets or on river banks just above sea level to 50 m., 

 Pete'n; Escuintla. British Honduras. Mexico (Chiapas) to Costa 

 Rica. West Indies. 



Plants annual, mostly 2-6 dm. tall, the stems and calyces armed with numer- 

 ous, long, straight, yellowish prickles, densely to sparsely stellate-pubescent 

 among the prickles, the hairs sessile and stipitate-stellate; leaves shallowly lobed 

 to deeply parted, the lobes angulate, shallowly lobate or undulate-dentate, the 

 leaves 4-24.5 cm. long, 2.5-14 cm. wide, the apex acute or obtuse, the base cordate 

 to truncate, sessile stellate-pubescent and prickly above and below; petioles 1-8 

 cm. long, stellate-pubescent and commonly with prickles; inflorescences lateral 

 and internodal, 1-3 flowered; peduncles 1-1.5 cm. long, prickly, stellate-pubescent; 

 pedicels 7-10 mm. long, stellate- pubescent and usually prickly; calyx 6.5-9 mm. 

 long, densely prickly and stellate-pubescent, deeply parted, the lobes lanceolate, 

 acute or acuminate; corolla purplish-white, the limb 20-25 mm. wide, shallowly 

 lobed, the lobes externally sessile stellate-pubescent; filaments 2-2.5 mm. long; 

 anthers 9.5-15.5 mm. long; style 13.5-18 mm. long, glabrous; ovary glabrous; 

 fruit globose, glabrous, 1-1.5 cm. in diameter, the accrescent calyx nearly as long 

 as the mature fruit; seeds 2 mm. long. 



Called huevos-de-gato in El Salvador. 




110 FIELDIANA: BOTANY, VOLUME 24 



Solanum canense Rydb. Bull. Torrey Bot. Club 51: 174. 1924. 



Usually along rivers and streams, near sea level to rarely up to 

 1,000 m.; Santa Rosa (Volcan de Tecuamburro, Standley 78562); 

 Honduras; Nicaragua. Panama. Western Venezuela and Colombia 

 to Ecuador. 



Plants herbaceous, weak and straggly, 4-15 dm. tall, the stems slender, some- 

 what angular, simple or branched, nearly glabrous to sparsely pilosulous, appar- 

 ently non-tuber-bearing; leaves odd-pinnate, 1-3.5 dm. long, nearly glabrous to 

 strigose; petioles 5 cm. long or less; interstitial leaflets numerous (rarely absent) 

 with short slender petiolules; leaflets 9-15, elliptic-lanceolate to narrowly lanceo- 

 late, 1.5-11 cm. long, 0.5-2.5 cm. wide, the apex acuminate to long-acuminate, 

 rounded to somewhat cordate at the oblique base, the lowermost pair of leaflets 

 much-reduced; petiolules slender, 2-12 mm. long; pseudostipular leaves present; 

 inflorescence pseudoterminal, becoming lateral and internodal, several-flowered, 

 bifurcate, cymose; peduncles slender, 4-13 cm. long, rarely 3-forked near the sum- 

 mit; pedicels slender, 5-10 mm. long, articulate at their bases; calyx 1.5-2.5 mm. 

 long, the lobes short rounded or apiculate, less than 1 mm. long; corolla white, the 

 limb 10-15 mm. wide, parted to about or just below the middle, the lobes ovate-lan- 

 ceolate to triangular-lanceolate, acute; filaments about 1 mm. long; anthers 2-3.5 

 mm. long, cordate at their bases; style 4-4.5 mm. long, slightly exceeding the sta- 

 mens; fruit globose to broadly ellipsoid, green, striped with pale or dark green, 

 1.5-2.5 cm. long. 



Solatium cervantesii Lagasca, Nov. Gen. & Sp. 10. 1816. 

 Fruta de schara (Chimaltenango) ; seconcillo and veneno (Quezal- 

 tenango). 



Moist forests or thickets, sometimes in dry thickets or open rocky 

 hillsides or oak forests, 2,000-3,150 m.; Baja Verapaz; Guatemala; 

 Chimaltenango; Quezaltenango ; Totonicapan ; Quiche"; Huehueten- 

 ango; San Marcos. Southern Mexico. El Salvador and Honduras. 



An unarmed shrub or small tree, 1-5 m. tall, often much branched, the branches 

 glabrous or puberulent; leaves lanceolate to oblong-elliptic or narrowly elliptic, 

 rarely oblanceolate, mostly 7-20 cm. long, 2-4 (-6. 5) cm. wide, the apex narrowly 

 acute or acuminate, the base long attenuate, sparsely pilosulous to glabrous above, 

 pubescent along the edges of the mid-vein and the lateral veins near the mid-vein, 

 the hairs branched or furcate, simple hairs often present below; petioles 1-3 cm. 

 long; inflorescences pseudoterminal becoming lateral and opposite the leaves, 

 dichotomously cymose, more or less flat-topped, several-flowered; peduncles long, 

 5-7 cm. long; pedicels slender, 5-8 mm. long; calyx 2-3 mm. long, the lobes 0.5-1 

 mm. long, triangular or oblong, pilosulous and glandular internally, acute and 

 apically pilosulous externally; corolla white, the limb 8-12 mm. wide, lobed to the 

 middle or just below, the lobes 2.5-5 mm. long, ciliate and, apically pilosulous; 

 filaments about 1 mm.; anthers 2-3 mm. long; style 4-5 mm. long, glabrous; ovary 

 glabrous; fruit globose, 5-10 mm. in diameter, black; seeds 3-3.5 mm. long. 



See comments under Solanum aligerum Schlecht. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 111 



Solatium chiapasense Roe, Brittonia 19: 367. 1967. 

 Thickets, 1,000-2,100 m.; Huehuetenango. Mexico (Chiapas) 

 type from 4 miles southwest of Las Rosas, Roe et al. 986. 



An unarmed shrub or small tree, 1-4 m. tall, the young branches stellate- 

 pubescent, the hairs sessile and short stipitate-stellate, with few or many rays; 

 leaves ovate, lanceolate-ovate or elliptic, mostly 12-30 cm. long, 5-15 cm. wide, the 

 apex acute to attenuate, the base attenuate, asperous to velutinous, the hairs 

 simple and sessile-stellate, with few rays above, densely stellate-pubescent, the 

 hairs sessile and short stipitate-stellate, with few or many rays below; petioles 2.5-8 

 cm. long; axillary leaves usually absent; inflorescence cymose, pseudoterminal, be- 

 coming lateral, dichotomously branched, long pedunculate, the peduncle 4-10 cm. 

 long, the hairs sessile or stipitate-stellate, with many or few rays; pedicels 2-5 mm. 

 long, pubescent, the stellate hairs with many or few rays; flower buds subglobose 

 to oblong; calyx lobed to the middle, the lobes triangular to ovate, 1.5-4.3 mm. long, 

 glabrous or occasionally pubescent within; corolla white or violet, the limb 12-20 

 mm. wide, parted to below the middle, the lobes 5-7.5 mm. long; filaments about 

 1.5 mm. long; anthers 3-4.5 mm. long; ovary pubescent; style glabrous or stellate- 

 pubescent; fruit yellow, 12-14 mm. in diameter, glabrescent; seeds 1.7-2.5 mm. 

 long. 



Solatium clarum Correll, Contr. Texas Res. Found. 1: 10, t. 

 4. 1950. 



Cloud forests and alpine areas, commonly under junipers and 

 pines, 3,100-3,800 m.; Totonicapan; Quezaltenango (type from Vol- 

 can Santa Maria, Skutch 858); Huehuetenango. Mexico (Chiapas). 



Plants erect, small, herbaceous, 1.5-2.5 dm. tall, sparsely pubescent through- 

 out, stoloniferous and tuber-bearing, the tubers small, ellipsoid, apparently pur- 

 plish, less than 2 cm. long; stem simple or branched; leaves simple, ovate, 1.5-8.5 

 cm. long, 1-3.5 cm. wide, the apex obtuse to acute, the base truncate to cordate; 

 petioles 5-45 mm. long; pseudostipular leaves present; inflorescence pseudotermi- 

 nal, becoming lateral and opposite the leaves, a short branched cyme, few-flowered; 

 peduncle 1-7 cm. long; pedicels 5-10 mm. long, articulate about or slightly 

 below the middle; calyx 3-4 mm. long, purple-tinged, the lobes short, about 1 mm. 

 long, triangular-ovate, obtuse to subacute; corolla purple at the tip of the lobes, 

 becoming white toward the base, parted to below the middle, the lobes 8-10 mm. 

 long, broadly lanceolate, acute to subobtuse; stamens equal, the filaments about 

 1 mm. long, glabrous, connate below the middle; anthers free, 3-4 mm. long, 

 dorsally minutely 3- to 5-lobulate at their bases; style puberulent, 7-8 mm. long, 

 exceeding the stamens; fruit (immature) broadly ovoid, 1.3 cm. long. 



Similar to S. morelliforme Bitter & Muench in general aspect but 

 distinguished from it by the anthers which are 3- to 5-lobulate 

 dorsally at their bases and the terrestrial habit. 



Solanum cobanense J. L. Gentry, Phytologia 26: 276. 1973. 

 Cyphomandm aculeata Donn.-Sm. Bot. Gaz. 57: 423. 1914 (type from 

 near Coban, Alta Verapaz, Lehmann 1334.). 




112 FIELDIANA: BOTANY, VOLUME 24 



Dense wet forests or wet thickets, 1,400-2,600 m.; Alta Verapaz; 

 Zacapa; Huehuetenango. Endemic. 



A woody vine, glabrous throughout, the branches armed with small recurved 

 prickles, the prickles sometimes numerous; leaves entire, subcoriaceous, solitary, 

 prickly below mostly on the mid-vein, lanceolate to lance-oblong or elliptic, 

 mostly 10-18 cm. long, 3-5.5 cm. wide, the apex acuminate or long acuminate, the 

 base obtuse to acute, glabrous above and below; petioles prickly, mostly 2.5-5 

 cm. long, glabrous; inflorescences pseudoterminal becoming lateral, few-flowered; 

 peduncles 4.5 cm. long, glabrous; pedicels 25-35 mm. long, glabrous; calyx un- 

 armed, subcoriaceous, glabrous, 3-3.5 mm. long, the lobes 1-1.5 mm. long, rounded 

 and mucronate; corolla dark purple, the limb about 23 mm. wide, parted nearly 

 to the base, the lobes about 10 mm. long, glabrous; filaments 2-2.5 mm. long; 

 anthers 6.5 mm. long; style glabrous, 6-6.5 mm. long, shorter than the stamens; 

 ovary glabrous; fruit glabrous, 7.5 cm. in diameter at the base, 6 cm. long, broadly 

 ovate; seeds 4.5-5 mm. broad, white. 



The treatment of this species has been confused. It was described 

 as a species of Cyphomandra and later placed in herbaria under 

 Solanum purulense Donn.-Sm., which proves to be a synonym of 

 S. lanceifolium Jacq. 



Solanum connatum Correll, Agr. Monogr. U.S.D.A. No. 11, 

 25, t. 11 & 12. 1952. 



On wooded slopes and epiphytic on trees, 2,500-3,000 m., ap- 

 parently endemic in San Marcos (type from between San Rafael at 

 northeast portion of Volcan Tacana and Guatemala-Mexico line, 

 Steyermark 36294). 



A woody, unarmed, non-tuber-bearing vine, the young stems and branches 

 densely pubescent, the hairs spreading; leaves odd-pinnate, 5.5-12 cm. long, 

 densely pilose above and below, glabrescent with age; petioles 1.5-4 cm. long; inter- 

 stitial leaflets absent; leaflets 5, elliptic to elliptic-oblanceolate or elliptic-obovate, 

 1-6.5 cm. long, 0.5-2.5 cm. wide, the apex acute to shortly and abruptly acuminate, 

 the petiolules subsessile to 1 cm. long, the lowermost pair of leaflets reduced, the 

 terminal leaflet larger than the lateral ones; pseudostipular leaves present or ab- 

 sent; inflorescence pseudoterminal, becoming lateral and opposite the leaves, 

 cymose; peduncle very short, 6 mm. long or less, densely pubescent with brownish- 

 yellow hairs; pedicels 5-7 mm. long, articulate at their bases, sparsely pubescent 

 to glabrescent; calyx 1.5-2 mm. long, the lobes short, triangular and acute; corolla 

 probably white, the limb about 8 mm. wide, parted to below the middle, the lobes 

 about 3 mm. long, broadly ovate, obtuse; filaments about 1 mm. long, pubescent, 

 united their entire length; anthers about 2 mm. long; style 5 mm. long, much 

 exceeding the stamens; fruit globose, apparently red, about 1 cm. in diameter. 



Solanum connatum Correll is closely related to S. tacanense 

 Lundell. The united filaments and the style exceeding the stamens 

 separate it from that species. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 113 



The following sterile collection, Steyermark 36834, as pointed 

 out by Correll (1962) probably represents a new species closely 

 related to S. connatum. The specimen has typically seven leaflets. 



Solatium cordovense Sess & Mocino Fl. Mex. ed. 2, 51. 1894. 

 S. lundettii Standl. Field Mus. Bot. 8: 42. 1930 (type from Roaring 

 Creek, British Honduras, Lundell 324), syn. nov. S. edwardsii Standl. 

 Trop. Woods 37: 31. 1934 (type from Temagua, Honduras, Edwards 

 P-639), syn. nov. 



Moist or wet thickets or forests, 200-800 m.; Alta Verapaz; 

 British Honduras. Southern Mexico. Honduras. Nicaragua. Costa 

 Rica. 



A slender, unarmed shrub, 1-5 m. tall, the young branches densely stellate- 

 pubescent, the hairs sessile or short stipitate-stellate, yellowish or brownish yellow, 

 sometimes glandular; leaves ovate-oblong to lance-oblong or elliptic-oblong, 6-14.5 

 cm. long, 2.5-5 cm. wide, the apex long acuminate, the base oblique, stellate-pubes- 

 cent, the hairs sessile, at least some with the central ray greatly exceeding the 

 others in length, sometimes giving the appearance of simple hairs above, the hairs 

 sessile or stipitate-stellate, velutinuous and more densely pubescent below; petioles 

 short, 2-7 mm. long; inflorescences lateral and internodal, cymose, sessile or short- 

 pedunculate; pedicels 1-2 cm. long, densely stellate-pubescent, often glandular; 

 calyx 5-6 mm. long, cleft nearly to the base, accrescent in fruit, densely sessile stel- 

 late-pubescent, at least some hairs with the central ray greatly exceeding the others 

 in length giving the appearance of simple hairs, the central ray often glandular, 

 the lobes oblong or elliptic, acute or obtuse, stellate-pubescent internally; corolla 

 white, the limb about 1.5 cm. wide, parted to near the base, the lobes about 6.5 mm. 

 long, a little longer than the calyx lobes, glabrous internally, stellate-pubescent ex- 

 ternally; filaments about 1 mm. long; anthers 2.5-3.5 mm. long; style 5-6.5 mm. 

 long, exceeding the stamens; fruit globose, 6-8 mm. in diameter, sparsely stellate- 

 pubescent or glabrous, the accrescent calyx nearly as long as to slightly exceeding 

 the mature fruit; seeds 2.5-3 mm. long. 



Solanum extensum Bitter, a species of Nicaragua, Costa Rica, 

 and Panama, is closely related to S. cordovense Sess & Mocino. 



Solanum cornutum Lam. Tab. Encycl. Meth. Bot. 2: 25. 1797. 

 In wet thickets or weedy fields, 900-1,450 m. ; Mexico. Honduras 

 (Dept. Morazan). To be expected in Guatemala. 



Plants herbaceous, 0.3-1 m. tall, the stems armed with numerous, long and 

 straight, yellowish prickles, covered with short, glandular and eglandular hairs 

 among the prickles, occasionally with a few stellate hairs intermixed, glabrate in 

 age; leaves bipinnatifid, the lobes entire to shallowly angulately-lobed, the leaves 

 5-20 cm. long, 2.5-10.5 cm. wide, prickly, glandular and sparsely sessile stellate- 

 pubescent, the stellate hairs with the central ray greatly exceeding the lateral 

 ones above, glandular and sessile stellate-pubescent below; petioles 1-7 cm. long, 




114 FIELDIANA: BOTANY, VOLUME 24 



prickly; inflorescences lateral and intern odal, 3-6 flowered; peduncles 1-2 cm. long, 

 prickly; pedicels 4-6 mm. long, prickles rarely present; calyx 5-7.5 mm. long, 

 receptacle and lower part of calyx covered with prickles of varying lengths and 

 sessile stellate-pubescent, the hairs with the central ray greatly exceeding the 

 lateral ones, parted to near the base, the lobes lanceolate to oblong-lanceolate, 

 somewhat unequal in length, stellate-pubescent, prickles absent; corolla yellowish, 

 slightly zygomorphic, the limb 2 cm. wide, shallowly lobed, externally sessile 

 stellate-pubescent; stamens unequal, the filaments of the short stamens 2-2.5 mm. 

 long, the fifth one 2-3 mm. long, the anthers of the short stamens 6-8 mm. long, 

 the fifth one much enlarged and incurved, 12-12.5 mm. long, sparsely villous; 

 style 13-15 mm. long; ovary glabrous; fruit globose, 7-9 mm. in diameter, enclosed 

 by the close fitting and prickly calyx, the calyx often adherent to the berry; seeds 

 2-2.5 mm. long. 



We have not seen specimens of Solanum cornutum Lam. from 

 Guatemala. It is included in the "Flora" because of its likely occur- 

 rence in the country. 



Solanum deflexum Greenm. Proc. Am. Acad. Sci. 32: 301. 1897. 

 In open forests or on brushy rocky slopes, 660-1,800 m. ; Chiqui- 

 mula; Jutiapa. Southern Mexico. Honduras to Costa Rica. 



A slender, erect or ascending herb, mostly 1-3 dm. tall, simple or much 

 branched, the stems with long spreading and short recurved hairs, the shorter 

 hairs inconspicuous and sometimes sparse; leaves solitary or in pairs, subequal in 

 size, similar in shape, the larger leaves ovate to lanceolate or elliptic-lanceolate, 

 2-8.5 cm. long, 0.5-3.5 cm. wide, the apex acute or sometimes acuminate, the 

 base rounded to short attenuate or sometimes subcordate, hirsute above and below, 

 the hairs sometimes subappressed ; petioles 5-25 mm. long; inflorescences lateral 

 and internodal, 1-4 flowered, umbelliform, sessile or the peduncles 2-3 mm. long; 

 pedicels hirsute, 7-14 mm. long, deflexed in fruit; calyx 1.5-2 mm. long, densely 

 hirsute, parted to near the base, the lobes lanceolate or oblong, acute or obtuse; 

 corolla white, the limb 7-8.5 mm. wide, shallowly lobed, the lobes hirsute and 

 papillate externally; filaments 0.5 mm. long; anthers 1.5-2 mm. long; style 2.5-3 

 mm. long, glabrous, exceeding the stamens; ovary glabrous; fruit globose, 5-8 mm. 

 in diameter; seeds 2.5-3 mm. long. 



Solanum deflexum Greenm. is distinguished from other species 

 in Guatemala by its anthers, simple leaves, habit, hirsute pubes- 

 cence, and the sessile or nearly sessile inflorescences. 



Solanum demissum Lindl. J. Roy. Hort. Soc. 3: 69,70. 1848. 

 S. alpicum Standl. & Steyerm. Field Mus. Bot. 23: 232. 1947 (type 

 from Cerro Chemal, Huehuetenango, Steyermark 50303}. Papa. 



Wet forests, alpine areas or on limestone outcrops, 2,250-3,700 m. ; 

 Sacatepe"quez ; Huehuetenango. South-central Mexico. 



Plants herbaceous, varying from a rosette to a much-branched spreading or as- 

 cending habit, commonly grayish-green, rarely up to 6 dm. tall, densely pubescent 




GENTRY & STANDLEY: FLORA OF GUATEMALA 115 



to essentially glabrous, tuber-bearing, the tubers 6 cm. long or less, usually ellipsoid 

 or panduriform, white or purple-tinged; leaves odd-pinnate 4.5-15(-30) cm. long, 

 interstitial leaflets present or absent; leaflets 5-9, usually 7, sessile or essentially so, 

 ovate to elliptic, the apex obtuse to acute, the base rounded to cuneate and slightly 

 oblique; lateral leaflets 1-6 cm. long, 0.5-2.5 cm. wide; terminal leaflet larger than 

 the lateral leaflets; inflorescence pseudoterminal, becoming lateral and opposite 

 the leaves, laxly cymose-paniculate; peduncle 5-30 mm. long; pedicels 1-2 cm. 

 long, articulate about or well above the middle; calyx 4-8 mm. long, lobed to 

 above or about the middle, the lobes ovate to triangular-lanceolate, acute; corolla 

 violet to purplish, the limb usually 20 mm. in diameter, shallowly lobed; filaments 

 about 1 mm. long; anthers 3-5 mm. long; style 9 mm. long or less, exceeding the 

 stamens; fruit globose to ovoid, green, sometimes with white specks, 1-2.5 cm. long. 



The fully ripened fruits have the odor of mango, papaya and, in 

 some, wild strawberry. 



Solanum demissum Lindl. is usually found at high elevations 

 where it is characteristically a rosette type of plant. At lower 

 elevations it often assumes an upright or bushy habit. 



This species is considered to be of potential value in the produc- 

 tion of blight and frost resistant strains in our cultivated potato. 



Solanum diphyllum L. Sp. PI. 184. 1753. 

 Damp thickets or forests, 325 m. or lower; Santa Rosa; Escuintla; 

 Suchitepe"quez; San Marcos. Mexico to Nicaragua. 



An unarmed shrub, 1-2 m. tall, glabrous throughout or nearly so, leaves solitary 

 or in pairs, unequal in size, different or similar in shape, the larger leaves narrowly 

 elliptic to oblanceolate or elliptic-oblong, sometimes ovate-lanceolate or lanceolate, 

 3-9 (-15. 5) cm. long, 1-2. 5 (-4) cm. wide, the apex obtuse or sometimes acute, equal 

 and attenuate at the base, glabrous above and below; petioles short, 2-5 mm. long, 

 sometimes winged to the base; inflorescences lateral and opposite the leaves, 

 cymose, simple, few-to several-flowered; peduncles short, 1-5 (-15) mm. long; calyx 

 1 mm. long, lobed to the middle, the lobes rounded and apiculate, obtuse or acute; 

 corolla white to greenish-white, the limb 6-7 mm. wide, parted to near the base, 

 the lobes 2.5-3 mm. long, inconspicuously ciliate; filaments about 0.3 mm. long; 

 anthers 1.5 mm. long; style 2.5-3 mm. long, glabrous, exceeding the stamens; ovary 

 glabrous; fruit globose, 6-8 mm. in diameter, black; seeds 3-3.5 mm. long. 



Apparently closely related to Solanum nudum H. B. K. and some- 

 times difficult to distinguish from it. Solanum nudum has the leaves 

 nearly always with a tuft of hairs in the axils of the mid-vein below 

 and the leaf apices acuminate or narrowly acute, whereas S. diphyl- 

 lum L. has glabrous leaves and the leaf apices mostly obtuse or acute. 



Solanum dulcamaroides Dunal in Poir. Encycl. Suppl. 3: 751. 

 1814. S. macrantherum Dunal, Solan. Syn. 16. 1816. 




116 FIELDIANA: BOTANY, VOLUME 24 



Huehuetenango (Sierra de los Cuchumatanes, probably about 

 3,200 m., Johnston 1978). Southern Mexico. 



A scandent, unarmed shrub, the branches pilosulous to glabrous; leaves ovate 

 to ovate-lanceolate, 5.5-14.5 cm. long, 2-5 cm. wide, the apex acuminate, the base 

 cuneate to rounded or sometimes subcordate, pilosulous above and below, especially 

 so on the veins to glabrous or essentially so; petioles long and slender, 2-4.5 cm. 

 long; inflorescences pseudoterminal, becoming lateral and opposite the leaves, 

 laxly cymose and widely paniculately disposed, several-flowered, pilosulous to 

 glabrous; peduncles 2-4 cm. long, pilosulous to glabrous; pedicels 1-2.5 cm. long, 

 articulate at their bases to just above the middle, pilosulous to glabrous; calyx 

 2-3 mm. long, broadly campanulate, sparsely pilosulous to glabrous, the lobes 

 apiculate, densely pilosulous; corolla violet, the limb 24-30 mm. wide, parted to 

 near the base, the lobes 10-13 mm. long, marginally pilosulous; filaments 2 mm. 

 long; anthers very broad, 5-6 mm. long; style greatly exceeding the stamens, 1-1.4 

 cm. long, glabrous; ovary glabrous; fruit globose, scarlet, about 10 mm. in diameter. 



Solanum dukamaroides Dunal is to be confused only with S. 

 seaforthianum Andr. in Guatemala and is distinguished from that 

 species by the broad anthers, simple leaves throughout, and the 

 pedicels articulate at their bases to slightly above the middle. 



Solanum erianthum D. Don, Prodr. Fl. Nepal. 96. 1825. S. 

 verbascifolium sensu auct., non L., 1753. Hediondilla (fide Aguilar); 

 tabaquillo. 



Moist or dry thickets, in fields or along river banks, often in 

 second growth, 5-2,000 m.; Peten, Izabal; Chiquimula; Jalapa; 

 Chimaltenango; Huehuetenango; Santa Rosa; Escuintla; San Marcos. 

 British Honduras. Southern Florida and Texas. Mexico south to 

 Costa Rica. Panama. West Indies. Galapogos Islands. Adventive 

 in the Old World. 



An unarmed shrub or small tree, 2-8 m. tall, the young branches densely 

 stellate-pubescent, the hairs sessile and short stipitate-stellate, with many rays 

 and often with many rays along their stalks; leaves entire, ovate to ovate-elliptic, 

 rarely lance-oblong, mostly 10-25 cm. long, 3-12 cm. wide, the apex acuminate, 

 the base rounded to nearly truncate or acute, velutinous and with simple and 

 sessile to short stipitate-stellate hairs above, simple hairs absent below; petioles 

 1-10 cm. long; axillary leaves absent; inflorescence cymose, pseudoterminal, be- 

 coming lateral, dichotomously branched, long pedunculate, the peduncle 3-12 cm. 

 long; pedicels 2-10 mm. long, densely stellate-pubescent, the hairs partly short 

 stipitate-stellate with many rays and often with many rays along their stalks; 

 flower buds turbinoid; calyx lobed to the middle, the lobes ovate, 2.5-4 mm. 

 long, stellate-pubescent within; corolla white, the limb 1-1.5 cm. wide, parted to 

 below the middle, the lobes 4-6 mm. long; filaments 1-1.5 mm. long; anthers 

 2.3-3 mm. long; ovary densely stellate-tomentose; style glabrous or sometimes 

 sparsely stellate-pubescent; fruit yellow, 1-1.2 cm. in diameter, stellate-pubescent; 

 seeds 1.5-2 mm. long. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 117 



Solanum erythrotrichum Fern. Proc. Am. Acad. Sci. 35: 561. 

 1900. Kaqi paxl (Quecchi) Coban; tomatillo (Alta Verapaz). 



Moist thickets or wet forests, sometimes along river banks, 

 1,000-1,450 m.; Pet6n; Alta Verapaz (type from Coban, Tuerckheim 

 1381}', Chimaltenango; British Honduras. Honduras. 



A shrub or small tree, 1.5-5 m. tall, the branches armed with few to many 

 prickles, these slightly recurved or spreading, mostly broad-based, the young 

 branches, peduncles and pedicels densely stellate-pubescent, the hairs ferruginous 

 and mostly with the central ray greatly exceeding the lateral ones; leaves solitary, 

 entire, ovate, sometimes elliptic or elliptic-oblong, 9-22 (-25) cm. long, 4. 5-10 (-14) 

 cm. wide, the apex acuminate or acute, the base rounded, subcordate or acute, 

 sometimes unequal, moderately covered with asperous, sessile-stellate hairs above, 

 the hairs grayish, densely sessile or subsessile stellate-pubescent below; petioles 

 2-4 cm. long, the hairs ferruginous, the central ray greatly exceeding the lateral 

 ones and with the rays subequal; inflorescences lateral and internodal, racemose, 

 few-flowered; peduncles 2-4.5 cm. long; pedicels 5-6 mm. long, erect in fruit, the 

 hairs with the central ray greatly exceeding the lateral ones and with the rays 

 subequal; calyx 6-7 mm. long, densely stellate-pubescent externally, moderately 

 so internally, parted nearly to the base, the lobes lanceolate or lance-oblong, 

 narrowly acute; corolla white, or sometimes lavender, the limb 20-23.5 mm. wide, 

 parted to near the base, the lobes 8-10 mm. long, densely sessile stellate-pubescent 

 externally; filaments 1-1.5 mm. long; anthers 5.5-7 mm. long; style 7-8.5 mm. 

 long, lower one-half sparsely stellate-pubescent; ovary densely stellate-pubescent, 

 the hairs sessile-stellate with the central ray greatly exceeding the lateral ones 

 and with the rays subequal; fruit globose, 1-1.5 cm. in diameter, stellate-pubes- 

 cent, sometimes sparsely so; seeds 4-4.5 mm. long. 



There are apparently two forms of Solanum erythrotrichum 

 Fernald. The common form in Guatemala has most of the hairs 

 with the central ray greatly exceeding the lateral ones on the young 

 branches, petioles, peduncles, and pedicels. A few specimens, Molina 

 253 and 298 and Standley 78365, have the rays of the hairs pre- 

 dominantly subequal. The form with the subequal rays is more 

 common in Honduras and may prove to be worthy of varietal rank. 



Solanum erythrotrichum is apparently closely related to S. ac- 

 crescens Standl. & Mort. and S. rubidum Standl. & Mort. of Costa 

 Rica. 



Called friega plato and lava platos in Honduras. 



Solanum fontium Standl. & Steyerm. Field Mus. Bot. 23: 234. 

 1944. Hediondilla (Huehuetenango) ; kaqi sakyol (Quecchi) Alta 

 Verapaz. 



Wet forests, 2,100-2,850 m.; Chimaltenango; Huehuetenango; 

 Quezaltenango (type from Fuentes Georginas, western slope of Volcan 

 de Zunil, Standley 67473); San Marcos. Mexico (Chiapas). 




118 FIELDIANA: BOTANY, VOLUME 24 



An unarmed shrub, 1.5-3.5(-6.5) m. tall, nearly glabrous throughout; leaves 

 mostly in pairs, unequal in size but similar in shape, the larger leaves elliptic to 

 oblong-elliptic or oblanceolate or ovate-elliptic, mostly 10-26 cm. long, 2.5-8.5 cm. 

 wide, the apex acuminate or sometimes acute, the base equal or unequal and 

 cuneate or short attenuate, glabrous above, sometimes inconspicuously and sparsely 

 puberulent along the margin of the mid-vein below; petioles 1-3 cm. long; in- 

 florescences lateral, opposite the leaves or internodal, cymose, mostly furcate, 

 several-flowered, glabrous; peduncles short, 3.5-20 mm. long; pedicels 1-1.5 cm. 

 long; calyx 2-3 mm. long, the lobes short, 0.5-1 mm. long, rounded and short- 

 apiculate, internally glandular; corolla white, the limb 15 mm. wide, parted to 

 near the base, the lobes 6 mm. long, apically pilosulous; filaments connate their 

 entire length or nearly so, 1 mm. long; anthers 3-4 mm. long; style shorter than 

 the stamens, 2.5-3 mm. long, glabrous; ovary glabrous; fruit globose, about 10 mm. 

 in diameter, black. 



It seems probable that some earlier name will be found for this 

 species. 



Solatium fraxinifolium Dunal in DC. Prodr. 13, pt. 1: 39. 



1852. S. grossularia Bitter, Fedde Repert. Sp. Nov. 10: 537. 1912. 



In thickets 2,000-2,600 m.; Alta Verapaz; Zacapa. Nicaragua. 



Costa Rica. Panama. Northern Colombia and western Venezuela. 



Plants herbaceous vine-like, decumbent, trailing or climbing, apparently non- 

 tuber-bearing, mostly densely pilose throughout with spreading hairs, sometimes 

 merely puberulent, 15 dm. or more in length, the stems slender, somewhat angular, 

 branched or sometimes simple, the hairs unequal in length; leaves odd-pinnate, 7- 

 23 cm. long; petioles 1.5-5 cm. long; interstitial leaflets absent or rarely present; 

 leaflets 5-7; elliptic-lanceolate to ovate-lanceolate or elliptic, 1-9 cm. long, 0.5-4.5 

 cm. wide, the apex obtuse to long acuminate, rounded to subcordate at the oblique 

 base, subsessile or short petiolulate, the lowermost pair of leaflets much reduced, 

 terminal leaflet larger than the lateral ones; pseudostipular leaves present; inflor- 

 escence lateral and internodal or opposite the leaves, racemose, usually laxly 6-10 

 flowered; peduncles 3-7 cm. long; pedicels slender, 7-12 mm. long, articulate at or 

 near the base; calyx 4-6 mm. long, divided to below the middle, the lobes lanceo- 

 late or broadly rounded, abruptly acuminate to acute; corolla white, the limb 13- 

 20 mm. wide, shallowly lobed, pilosulous on the outside; filaments about 1 mm. 

 long, free or partially connate, glabrous or sparsely pilosulous; anthers 3-4 mm. 

 long; style 6-7 mm. long, exceeding the stamens; fruit ovoid to ovoid-ellipsoid, 

 somewhat compressed, green with darker green or purplish stripes, 1.5-2 cm. long, 

 1-1.5 cm. in diameter. 



Solanum globiferum Dunal in DC. Prodr. 13, pt. 1: 251. 1852. 

 Chichito de raton (Huehuetenango) . 



Dry or moist thickets or in pine-oak forests, sometimes in open 

 rocky places, 850-1, 650 m.;Petn; Baja Verapaz; Jalapa; Guatemala; 

 Chimaltenango; Huehuetenango; Jutiapa; Santa Rosa; Escuintla. 

 British Honduras. Southern Mexico to Nicaragua. Martinique. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 119 



Plants herbaceous or suffrutescent, 1.5 m. tall or less, the branches densely 

 pubescent, the hairs simple, short and mostly glandular, armed with a few, broad- 

 based, yellowish, somewhat recurved prickles and with numerous short, slender, 

 spreading or somewhat reflexed ones; leaves armed with prickles, broadly ovate, 

 sinuate to shallowly lobed or angulately-lobed, mostly 7.5-30 cm. long, 4.5-20 cm. 

 wide, the apex acute, the base cordate, densely pubescent with simple hairs, the 

 hairs long and soft, some of the hairs glandular above, densely pubescent with 

 simple, glandular and eglandular hairs and sessile-stellate hairs intermixed below; 

 petioles mostly 3-12 cm. long, densely pubescent with short glandular and eglandu- 

 lar hairs, the hairs sometimes almost entirely glandular, armed with prickles; 

 inflorescences lateral and internodal, umbelliform, sessile or subsessile, mostly 1-3 

 flowered; pedicels lax, 5-10 mm. long, armed, densely pubescent, the hairs short, 

 glandular and eglandular, the hairs sometimes almost entirely glandular; calyx 

 2-3 mm. long, densely pubescent, the hairs glandular and eglandular, often with 

 prickles, parted nearly to the base, the lobes lanceolate, sometimes pubescent 

 internally; corolla white, the limb 13.5-20 mm. wide, parted nearly to the base, 

 the lobes 5.5-8.5 mm. long, pubescent with long hairs externally or rarely glabrous; 

 filaments about 1 mm. long; anthers 6-7 mm. long; style slender, glabrous, 7-7.5 

 mm. long; ovary pubescent with simple glandular and eglandular hairs or only 

 glandular ones; fruit pale yellow, glabrous, 2-2.5 cm. in diameter, globose; seeds 

 2.5-3 mm. long, wing absent. 



Called huevo de gato and huistomate in El Salvador and chichiqua, 

 comida de culebra, manzanita, and tomate in Honduras. 



The correct name for this species may be Solanum chloropetalum 

 Schlecht. 



See comments under Solanum quinquangulare Roem. & Schult. 



Solanum hartwegii Benth. PI. Hartweg. 68. 1839. S. lauri- 

 folium sensu auct., non Miller 1768. Flor de pajalkish and limpia- 

 plato (Quezaltenango) ; friega-plato (Jalapa); huiz (Sacatepquez) ; 

 kakaquish and lava-plato (Huehuetenango) ; kakisacyol and kaqi-paxl 

 (Quecchi) Alta Verapaz. 



Moist or wet thickets, often on dry brushy hillsides, frequently 

 in oak or pine forests, 1,200-3,200 m., Alta Verapaz; Baja Verapaz; 

 Zacapa; Chiquimula; Jalapa; El Progreso; Guatemala; Sacatepe"quez ; 

 Chimaltenango; Solola; Quiche"; Totonicapan; Quezaltenango; Hue- 

 huetenango; Jutiapa; San Marcos. British Honduras. Mexico to 

 Costa Rica. Martinique. 



A shrub, 0.5-3.5 m. tall, the branches densely covered with a whitish or grayish 

 stellate-tomentum, the hairs weak, long stipitate-stellate, sparsely armed with 

 short, straight prickles or the prickles absent; leaves solitary, subentire to shallowly 

 lobed, ovate to ovate-elliptic or ovate-lanceolate, 12-25 cm. long, 8-15 cm. wide, 

 the apex narrowly acute to acute, the base rounded to cordate, unequal or equal, 

 sparsely to densely short stipitate stellate-pubescent or sometimes nearly glabrous, 

 especially in age, rarely with prickles above, densely stellate-tomentose, the hairs 




120 FIELDIANA: BOTANY, VOLUME 24 



whitish to grayish and stipitate-stellate, also with a few prickles on the mid-vein 

 below; petioles 2-3 (-5) cm. long, densely stellate-tomentose pubescent, the hairs 

 whitish to grayish and stipitate-stellate; inflorescences lateral and internodal, 

 cy'mose-corymbose, many-flowered, densely stellate-tomentose pubescent, the 

 hairs long stipitate-stellate and whitish to grayish; peduncles short, 5-20 mm. long; 

 pedicels short and stout, 5-10 mm. long; calyx (4.5-)6-8 mm. long, sessile and 

 long stipitate stellate-pubescent, the lobes (1.5-)4-5 mm. long, linear; corolla bright 

 to pale purple, the limb (25-)30-35 mm. wide, lobed about one-half to slightly 

 more than one-half way to the base, the lobes (7-)10-12 mm. long, ovate, apex 

 acuminate, densely sessile and short stipitate stellate-pubescent externally, the 

 hairs sessile-stellate on the mid-vein internally; filaments 2-2.5 mm. long; anthers 

 (5-)6-7.5 mm. long; style 9.5-10.5 mm. long, the base covered with stipitate- 

 glandular or sessile-stellate and stipitate-glandular hairs; ovary covered with 

 stipitate-glandular or sessile-stellate and stipitate-glandular hairs; fruit globose, 

 1-1.5 cm. in diameter; seeds 2-3 mm. broad. 



Solatium hartwegii is distinguished from other species in Guate- 

 mala by the anthers, habit, whitish to grayish, long stipitate-stel- 

 late hairs, and the purple corolla. 



The name Solatium laurifolium Miller has been applied to this 

 species. However, a recent examination of the type by W. G. D'Arcy 

 has shown that species to be a synonym of S. subinerme Jacq. 



I have included here a collection (from the Sierra de los Cuchu- 

 matanes, Huehuetenango, Steyermark 51626} which is densely stellate 

 pubescent throughout. The stem and inflorescences are covered 

 with relatively long, stipitate-stellate hairs. The hairs are reddish- 

 brown except for the grayish lower surface of the undulate to parted 

 leaves. 



Solarium hazenii Britton, Bull. Torrey Bot. Club 48: 338. 1922. 



Moist or dry thickets, 400-600 m. ; Zacapa; Chiquimula. Western 

 Mexico, south to Costa Rica. Panama. West Indies. Northern 

 South America. 



An unarmed shrub or small tree, 1-6 m. tall, the young branches densely 

 stellate-pubescent, the hairs sessile and short stipitate-stellate with many rays, 

 often with many rays along their stalks; leaves entire, ovate to ovate-elliptic or 

 elliptic, sometimes obovate, mostly 10-28 cm. long, 3-12 cm. wide, the apex acute 

 to acuminate, the bases short or long attenuate, asperous to finely veiutinuous, the 

 hairs simple and sessile or subsessile stellate-pubescent above, densely pubescent, 

 the hairs mostly subsessile to short stipitate-stellate below; petioles 1-7 cm. long; 

 axillary leaves present or absent; inflorescence cymose, pseudoterminal, becoming 

 lateral, dichotomously branched, long pedunculate, the peduncle 4-15 cm. long; 

 pedicels 3-8 mm. long, stellate-pubescent; flower buds orbicular, the base swollen; 

 calyx cleft to near the base, the lobes ovate, 2.5-4 mm. long, stellate-pubescent 

 internally; corolla white, the limb 10-15 mm. wide, parted to below the middle, the 




GENTRY & STANDLEY: FLORA OF GUATEMALA 121 



lobes 4-6 mm. long; filaments 1-1.3 mm. long; anthers 2-3 mm. long; ovary 

 glabrous or with occasional stellate hairs; style glabrous or sparsely stellate- 

 pubescent; fruit yellow, 8-10 mm. in diameter, glabrescent; seeds 1.5-2.2 mm. long. 



Solanum hirtum Vahl, Symb. Bot. 2: 40. 1791; Icon. PI. Am. 

 t.21. 1798. 



Wet thickets or forests, 200-600 m.; Pete"n; Zacapa; Chiquimula. 

 British Honduras. Mexico to Costa Rica. Panama. West Indies. 

 Northern South America. 



A shrub, 1-2 m. tall, densely armed with straight and commonly slender 

 prickles, the young branches, petioles, and pedicels densely covered with whitish 

 or grayish, sessile-stellate hairs, the central ray glandular (at least some) and 

 greatly exceeding the lateral ones, the rays subequal on some of the hairs; leaves 

 solitary, sinuate-lobate, broadly ovate, mostly 10-20 cm. long, 5.5-10 cm. wide, 

 the apex obtuse, the base cordate, soft to the touch, sessile stellate-pubescent, the 

 hairs often with the central ray greatly exceeding the lateral ones, a few simple 

 hairs usually present, the young leaves often conspicuously glandular above, 

 densely covered with grayish, sessile to short stipitate-stellate hairs below; petioles 

 armed with prickles, mostly 2.5-8 cm. long; inflorescences lateral and internodal, 

 umbelliform, subsessile; pedicels 10-13 mm. long; calyx 9-9.5 mm. long, densely 

 stellate-pubescent externally, the hairs sessile and short stipitate-stellate, parted 

 nearly to the base, the lobes reflexed in fruit, lanceolate or ovate, acuminate or 

 acute, densely sessile stellate-pubescent, the hairs with the central ray glandular 

 and much exceeding the lateral ones internally; corolla white, the limb 25-30 mm. 

 wide, parted nearly to the base, the lobes 10-12 mm. long, densely stellate-pubes- 

 cent, the hairs sessile and short stipitate-stellate externally, glabrous internally; 

 anthers subsessile, 7-7.5 mm. long; style 9-9.5 mm. long, glabrous; ovary densely 

 stellate-pubescent, the hairs with the central ray greatly exceeding the lateral ones, 

 glandular or eglandular; fruit globose, orange-red to red, 1.5-2 cm. in diameter, 

 densely stellate-pubescent; seeds 1.5-2.5 mm. long. 



Called shumpa in British Honduras, putbalam in Yucatan, and 

 huevo de gato in Honduras. 



Both Solanum hirtum Vahl and S. tequilense Gray are readily 

 distinguished from other members of the genus in Guatemala by 

 their large, densely stellate-pubescent, orange-red to red fruits. 



See comments under S. tequilense Gray. 



Solanum hispidum Pers. Syn. PL 1 : 228. 1805. Ruiz (Sacate- 

 pe"quez); kakisacyol, and kaqi paxl (Quecchi, Alta Verapaz); limpia- 

 plato (Quezaltenango). 



Moist or wet thickets, sometimes in oak or oak-pine forests or in 

 fields, 1,200-2,500 m.; Alta Verapaz; Izabal; Chiquimula; Baja 

 Verapaz; El Progreso; Sacatepe"quez ; Chimaltenango; Quiche"; Que- 




122 FIELDIANA: BOTANY, VOLUME 24 



zaltenango; Huehuetenango ; Santa Rosa; San Marcos. Southern 

 Mexico to Costa Rica. Panama. Peru. 



A shrub or small tree, 1-5 m. tall, armed with few prickles, rarely unarmed, 

 the prickles mostly stout and broad based, the young branches, petioles, peduncles 

 and pedicels densely covered with mostly stout, ferruginous, long stipitate- 

 stellate (bristle-like) hairs; leaves solitary, ovate to elliptic or oblong-elliptic, 

 sinuately-lobed to lobed or often pinnatifid, rarely entire, mostly 15-30 cm. long, 

 10-20 cm. wide, the apex narrowly acute to acute, the base unequal or equal, 

 rounded or obtuse, sometimes subcordate, asperous and sometimes bearing a few 

 prickles, covered with sessile-stellate hairs, the terminal ray exceeding the lateral 

 ones in at least some of the hairs above, sessile to stipitate stellate-pubescent below; 

 petioles 2-5 cm. long; inflorescences lateral and internodal, few-flowered, densely 

 stellate-pubescent; peduncles 3-15(-30) mm. long; pedicels 5-10 mm. long; calyx 

 5-9 mm. long, densely stellate-pubescent, parted nearly to the base, the lobes linear 

 or lanceolate and long attenuate; corolla white, the limb 20-28 mm. wide, divided 

 to well below the middle, the lobes 7.5-12 mm. long, densely stellate-pubescent 

 externally, sparsely stellate-pubescent along the mid-vein internally; filaments 2-3 

 mm. long; anthers 6.5-8 mm. long; style 9-13.5 mm. long, glabrous; ovary sparsely 

 pubescent apically, the hairs stipitate-glandular; fruit globose, 10-14 mm. in diam- 

 eter, glabrous; seeds 2-3 mm. long. 



Solatium hispidum Pers. is in most cases distinguished readily 

 from other members of the genus in Guatemala by the prominent 

 reddish or reddish-brown, long stipitate-stellate hairs covering most 

 parts of the plant. 



Solatium houstonii Dunal, Hist. Solan. 243. 1813. S. tampi- 

 cense Dunal in DC. Prodr. 13, pt. 1: 284. 1852. 



Damp thickets, swampy ground or open places, just above sea 

 level to 35 m.; Escuintla (near San Jose", Standley 64.257}; British 

 Honduras. Southern Mexico. West Indies. 



A small, woody vine, the stems glabrate, bearing a few minute, sessile-stellate 

 hairs when young, sparsely to densely armed with prickles, the prickles short and 

 recurved; leaves in pairs or solitary, sinuately-lobed to parted, the larger leaves 

 oblong to lance-oblong, 5.5-16 cm. long, 2.2-5.5 cm. wide, the apex acute, the 

 base attenuate, prickly below and often on the mid-vein above, sparsely pubescent 

 throughout, the hairs sessile-stellate and often with the central ray much longer 

 than the lateral ones above, the hairs sessile-stellate and with subequal rays below; 

 petioles glabrous or sessile stellate-pubescent, 5-15 mm. long, often armed with a 

 few long prickles; inflorescences lateral and opposite the leaves, cymose, sessile or 

 subsessile; pedicels glabrous or sparsely sessile stellate-pubescent, 5-8 mm. long, 

 recurved in age; calyx 1.5-2 mm. long, the lobes mucronate, shallowly lobed in age, 

 nearly always armed with prickles, glabrous or sometimes sparsely sessile stellate- 

 pubescent; corolla white to yellowish-white, the limb 12-14 mm. wide, parted 

 nearly to the base, the lobes 5-6 mm. long, ciliate and sparsely sessile stellate- 

 pubescent externally; filaments about 1 mm. long; anthers 3.5-4 mm. long; style 




GENTRY & STANDLEY: FLORA OF GUATEMALA 123 



5-6 mm. long, glabrous; ovary glabrous; fruit globose, red, 8-8.5 mm. in diameter; 

 seeds 2-2.5 mm. long. 



In Guatemala Solanum houstonii Dunal is to be confused only 

 with S. lanceifolium Jacq. 



Called sosumba in British Honduras. 



Solatium huehuetecum Standl. & Steyerm. Field Mus. Bot. 

 23: 234. 1947. Yerba mora. 



Known only from the type, Huehuetenango, above Democracia 

 on trail towards Jutal, 1,000 m., Steyermark 51039. Endemic. 



A small, unarmed, suffrutescent herb, about 3 dm. tall, the branches slender, 

 sparsely to moderately stellate-pubescent, the hairs sessile or short stipitate; 

 leaves ovate to lanceolate or lance-elliptic, 2.5-6 cm. long, 1-2 cm. wide, the apex 

 acute, the base short attenuate, the hairs short stipitate-stellate above and below, 

 simple hairs also present above; petioles slender, 5-15 mm. long; inflorescences 

 lateral and opposite the leaves or internodal, sessile or subsessile, cymose, few- 

 flowered; pedicels slender, about 2 cm. long, sparsely stellate-pubescent; calyx 

 about 2.5 mm. long, sparsely stellate-pubescent, parted to the middle, the lobes 

 ovate, acute; corolla white, 7-8 mm. long, lobed to the middle, the lobes apically 

 stellate-pubescent externally, otherwise glabrous; filaments connate to one-half or 

 almost their entire length, about 1 mm. long; anthers about 3 mm. long; style 

 3.5 mm, long, glabrous; ovary glabrous; fruit globose, about 6 mm. in diameter. 



A collection, Steyermark 51278, made near the type locality of 

 Solanum huehuetecum Standl. & Steyerm. has been included here. 

 The specimen is more densely pubescent, leaf bases rounded to 

 subcordate, and the filaments only connate at the base. It also 

 suggests S. lignescens Fern, of southern Mexico. 



Solanum inscendens Rydb. Bull. Torrey Bot. Club 51: 175. 

 1924. S. ovandense Lundell, Amer. Midi. Nat. 29: 490. 1943 (type 

 from Mount Ovando, Chiapas, Mexico, Matuda 4182). 



In mixed or pine-juniper forests, or cloud forests, commonly 

 climbing on trees, 2,400-3,768 m.; Baja Verapaz; Guatemala; Chi- 

 maltenango; Quezaltenango ; Huehuetenango; San Marcos. South- 

 central Mexico. 



A woody, non-tuber-bearing vine, up to 4 m. long or more, the stems rooting 

 at the nodes; young branches densely pubescent to essentially glabrous; leaves 

 odd-pinnate, 3-10 cm. long, pubescent to essentially glabrous above and below; 

 petioles 1-4 cm. long; interstitial leaflets absent; leaflets 5, ovate to elliptic, the 

 apex acute to acuminate, 1.5-5.5 cm. long, 0.5-1.5 cm. wide, shortly petiolulate 

 or subsessile, the lowermost pair of leaflets reduced or sometimes minute, the 

 terminal leaflet somewhat larger than the lateral ones; pseudostipular leaves 

 present; inflorescence pseudoterminal on leafy branches, becoming lateral and 




124 FIELDIANA: BOTANY, VOLUME 24 



internodal, cymose-paniculate; peduncle usually short, rarely 30 mm. long; 

 pedicels 5-10 mm. long, articulate at their bases, sparsely pubescent; calyx about 

 2 mm. long, shallowly lobed, the lobes short, triangular-ovate, acute to apiculate, 

 about 0.5 mm. long; corolla white, the limb 6-10 mm. wide, deeply lobed to near the 

 base, the lobes 2-4 mm. long, elliptic to lanceolate, obtuse to acute; filaments about 

 0.5 mm. long, pubescent, slightly connate below; anthers 2-2.5 mm. long; style 

 5-7 mm. long, conspicuously exceeding the stamens, curved with age, fruit globuse, 

 about 1 cm. in diameter. 



Solanum inscendens Rydb. is closely allied to S. appendiculatum 

 H. & B. ex Dunal. 



Solanum jamaicense Miller, Gard. Diet. ed. 8. Solanum no. 

 17. 1768. 



Moist or wet thickets or in pastures, just above sea level to 900 m. ; 

 Pete"n; Alta Verapaz; Izabal. British Honduras; Southern Mexico; 

 Honduras. Nicaragua. Costa Rica. Panama. West Indies. South 

 America. 



An erect shrub, 1-3.5 m. tall, the branches densely pubescent, covered with 

 short stipitate or sometimes subsessile-stellate hairs, these often yellowish or 

 yellowish-brown, armed with numerous short, recurved prickles; leaves sessile or 

 subsessile, in pairs or some solitary, angulately-lobed to subentire, rarely entire, 

 broadly ovate to elliptic or oblong-elliptic, often rhomboid, the subopposite leaves 

 similar in shape, different in size, the larger leaves 6-20 cm. long, 3.5-10 cm. wide, 

 the apex narrowly acute or long acuminate, the base broadly cuneate and de- 

 current, often bearing a few prickles, densely pubescent above and below, the hairs 

 short stipitate-stellate, usually more densely pubescent below; inflorescences 

 lateral and internodal, sessile or subsessile, umbelliform, several-flowered; pedicels 

 5-8 mm. long, stipitate stellate-pubescent, often with a few prickles; calyx 2.5-3 

 mm. long, stipitate stellate-pubescent, often with a few prickles; corolla white, 

 the limb 12-12.5 mm. wide, parted nearly to the base, the lobes 5-5.5 mm. long, 

 oblong-lanceolate to narrowly lanceolate, the hairs sessile and short stipitate- 

 stellate externally; filaments 0.5-1 mm. long; anthers 3.5-4.5 mm. long; style 

 5.5-6.5 mm. long, the hairs stipitate-glandular below, sometimes with a few sessile 

 stellate hairs in addition to the glandular ones; ovary stipitate-glandular pubescent, 

 sometimes with a few sessile stellate hairs in addition to the glandular ones; fruit 

 globose, red, 7.5-8.5 mm. in diameter, glabrous; seeds 1.5 mm. broad. 



Solanum jamaicense Miller shows little variation throughout its 

 range. It is distinguished from other species of the genus in Guate- 

 mala by its anthers, strongly recurved prickles, sessile or subsessile 

 and broadly cuneate leaves, lateral and internodal and sessile in- 

 florescences and the stipitate-stellate hairs on the leaves. 



A common weedy shrub of the Caribbean coast region. 

 Called friega plato and tomate del diablo in Honduras. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 125 



Solatium lanceifolium Jacq. Coll. Bot. 2: 286. 1789. S. 

 scabrum Vahl, Eclog. Am. 1: 22. 1796, non S. scabrum Miller. S. 

 enoplocalyx Dunal in DC. Prodr. 13, pt. 1: 222. 1852, syn. nov. S. 

 donnell-smithii Coult. Bot. Gaz. 16: 144. 1891 (type from Concepcion, 

 Escuintla, Donn.-Sm. 2261), syn. nov. S. purulense Donn.-Sm. Bot. 

 Gaz. 52: 52. 1911, syn. nov. Kishtan del monte (San Marcos); man- 

 zanilla de montana (Suchitepe"quez) ; tomatillo (Jutiapa) . 



Wet or moist thickets or forests, sometimes in dry thickets, 

 50-1,650 m.; Pete'n; Alta Verapaz; Izabal; El Progreso; Solola; 

 Guatemala; Quezaltenango ; Jutiapa; Santa Rosa; Escuintla; Suchi- 

 tepe"quez; Retalhuleu; San Marcos. British Honduras. Southern 

 Mexico to Costa Rica. Panama. West Indies. Northern South 

 America. 



A woody vine, trailing over shrubs or small trees, the branches armed with 

 short, stout, broad-based, recurved prickles, also densely covered with sessile to 

 short stipitate-stellate hairs or long, stout, stipitate-stellate (bristle-like) hairs, 

 these unequal in length, sometimes sparsely stellate-pubescent; leaves shallowly 

 angulately-lobed to entire, in pairs, similar in shape, slightly different in size, 

 ovate or elliptic to oblong, sometimes obovate, 10-22 cm. long, 4.5-10 cm. wide, 

 the apex acuminate to narrowly acute or acute, the base acute to short attenuate, 

 stellate-pubescent, the hairs sessile-stellate, the central ray much exceeding the 

 lateral ones, occasionally with a few prickles above, stellate-pubescent, the hairs 

 sessile-stellate, the rays nearly subequal, prickly along the mid- vein below; 

 petioles 1.5-3.5 cm. long, sessile and short stipitate stellate-pubescent, armed with 

 prickles; inflorescences lateral and internodal, cymose; peduncles unarmed, 1-2.5 

 cm. long, the hairs long stipitate-stellate, unequal in length or short stipitate to 

 sessile-stellate; pedicels 5-8 mm. long, the hairs unequal, long stipitate-stellate or 

 short to sessile-stellate, often with a few prickles; calyx 3-4 mm. long, densely 

 armed with prickles, densely stellate-pubescent, the hairs long stipitate-stellate or 

 sessile and short stipitate-stellate, the lobes linear-oblong or caudate, 2-3 mm. long, 

 the calyx splitting in age; corolla pale purple or white, the limb 22-27 mm. wide, 

 parted nearly to the base, the lobes 10-12,5 mm. long, lance-oblong, densely sessile 

 stellate-pubescent externally, the hairs sessile-stellate on the mid-vein internally; 

 filaments 1 mm. long; anthers 6-11 mm. long; style 9-11.5 mm. long, lower one-half 

 to three-fourths sessile stellate-pubescent, the central ray exceeding the lateral 

 ones, or rarely with stipitate-glandular hairs; ovary apically sessile stellate- 

 pubescent or rarely with stipitate-glandular hairs; fruit globose, orange, 12-15 mm. 

 in diameter; seeds 3 mm. broad. 



Called huevo de gato and tomate del diablo in Honduras and 

 tomatillo in British Honduras. 



Solanum lanceifolium Jacq. is characterized by its high degree of 

 variability, especially as to pubescence. Part of the material in- 

 cluded here may prove to be Solanum subinerme Jacq. 




126 FIELDIANA: BOTANY, VOLUME 24 



Solanum lepidotum H. & B. ex Dunal, Solan. Syst. 17. 1816. 

 Hoja bianco, (San Marcos) ; hoja huaco (Quezaltenango) . 



Wet forests, 150-1,000 m., sometimes higher; Alta Verapaz; 

 Suchitepe"quez; Huehuetenango ; Quezaltenango; San Marcos. British 

 Honduras. Southern Mexico. Costa Rica. Panama to northern 

 South America. 



A shrub or small tree, unarmed, 1-6 m. tall, the young branches, petioles and 

 peduncles, grayish, densely covered with minute, closely appressed stellate- 

 like scales and stellate hairs; leaves mostly solitary, sometimes in pairs and 

 then very unequal in size but similar in shape, the larger leaves lance-oblong to 

 elliptic-oblong, sometimes elliptic, mostly 7-15 cm. long,. 3-7 cm. wide, the apex 

 acuminate or long acuminate sometimes abruptly so, the base obtuse or rounded, 

 rarely short attenuate, essentially glabrous or with a few scattered, minute, 

 stellate hairs, smooth to the touch above, densely covered with minute and 

 closely appressed whitish or grayish stellate-like scales below; petioles 5-10 (-15) 

 mm. long; inflorescences pseudoterminal, becoming lateral and opposite the leaves, 

 dichotomously cymose, many-flowered; peduncles 2.5-8.5 cm. long; pedicels 5 mm. 

 long, densely stellate-tomentose; calyx 2-2.5 mm. long, densely stellate-tomentose, 

 parted to the middle, the lobes ovate, acute; corolla white, the limb 7 mm. wide, 

 parted to near the base, the lobes 3 mm. long, densely stellate-tomentose externally; 

 filaments short, about 0.5 mm. long; anthers 1.5-2 mm. long; style 3 mm. long, with 

 a few scattered stellate hairs; ovary glabrous; fruit globose, 7-8 mm. in diameter, 

 black, glabrous; seeds 3 mm. long. 



Solanum lepidotum is most likely to be confused with S. schlecht- 

 endalianum but can be distinguished by the glabrous ovary and the 

 closely appressed stellate-like scales on the lower surface of the leaves. 



Solanum mammosum L. Sp. PL 187. 1753. Cantu (Quecchi, 

 signifying yellow breast); chichigua (Chiquimula) ; chichita (Pete"n); 

 tetereta (Jutiapa). 



Dry or moist thickets or fields, often in waste ground, 150-1,500 

 m., but in cultivation at much higher elevations; Pete"n; Alta Verapaz; 

 Chiquimula; Jalapa; Huehuetenango; Jutiapa; Santa Rosa. British 

 Honduras. Southern Mexico. El Salvador. Honduras. Nicaragua. 

 Costa Rica. Panama. West Indies. South America. 



Plants herbaceous or suffrutescent, 1.5 m. tall or less, armed with numerous 

 slender or broad-based, yellowish, straight or partly recurved prickles, the branches 

 stout, densely pubescent with long, simple hairs, the hairs soft and multicellular 

 and stipitate-glandular; leaves armed with prickles, ovate to broadly ovate, shal- 

 lowly lobed, mostly 10-25 cm. long, 8-18 cm. wide, the apex acute, the base 

 cordate to subcordate or nearly truncate, densely pilose above and below, the hairs 

 simple; petioles mostly 4-10 cm. long, densely long pilose and with stipitate- 

 glandular hairs, armed with prickles; inflorescences lateral and internodal, cymose, 

 sessile or subsessile, mostly 1-4 flowered; pedicels 9-14.5 mm. long, densely pilose 

 and with stipitate-glandular hairs; calyx 4-5 mm. long, unarmed, densely pilose 




GENTRY & STANDLEY: FLORA OF GUATEMALA 127 



and with stipitate-glandular hairs, parted nearly to the base, the lobes narrowly 

 lanceolate to linear; corolla violaceous, the limb 24-28 mm. wide, parted nearly to 

 the base, the lobes pilose externally; filaments 1-1.5 mm. long; anthers 8.5-10 mm. 

 long; style stout, about as long as the stamens, apparently developing late in some 

 flowers, the lower part glandular; ovary glandular; fruit bright yellow, glabrous, 

 2.5-4.5 cm. broad, 3-6.5 cm. long, ovoid and abruptly contracted at the apex into 

 a short neck (mammillate); seeds 3.5-4 mm. long. 



Called chichihua in Honduras and Nicaragua. 



This is one of the best known plants in Guatemala because of its 

 connection with the pilgrimage each January to the Sanctuary of 

 Esquipulas, most famous church of Central America. The church is 

 visited each year by many tens of thousands of romeros or pilgrims 

 from Mexico and Central America and even from more distant 

 regions. One meets many pilgrims on the roads before and after the 

 celebration, most of whom travel on foot, thus acquiring greater 

 merit. They are easily recognized because the women wear decora- 

 tions of these chichita fruits. The men have their hats decorated 

 with Spanish moss (Tillandsia usneoides L.). The word chichihua 

 and its variations, used generally in Mexico and Central America 

 for women's breasts, is derived from a Nahuatl word having the 

 same significance, and alludes to the shape of the fruit. Since a 

 great many of the women who go to Esquipulas do so because of a 

 belief that it will result in bearing a child, the connection between 

 the Solatium fruit and pilgrimage is rather obvious. The custom is 

 probably a very old one. All over Guatemala one sees the dry fruits 

 used as decorations in the humblest dwellings, a souvenir of an earlier 

 pilgramage. The seeds are often planted about dwellings far above 

 the elevations at which the plants grow naturally, as a further indi- 

 cation that people of that home have made the pilgrimage. It may 

 be noted that pilgrims to another famous church, at San Felipe, near 

 Antigua, carry branches of the pepper tree, Schinus molle L. Branch- 

 es of Solanum mammosum L. with the brilliant yellow fruits are often 

 used about Guatemala City as Christmas decorations. The fruits 

 sometimes are used in domestic medicine as a remedy for colds, but 

 their use is rather dangerous as there is a belief, probably well based, 

 that they are poisonous. 



Solanum melongena L. Sp. PL 186. 1753. Berenjena; egg- 

 plant. 



A native of Asia, now cultivated in moist warm or warm-tem- 

 perate countries for its edible fruits. Planted in various parts of 

 Guatemala at middle and lower elevations. The eggplant, so widely 




128 FIELDIANA: BOTANY, VOLUME 24 



used as a vegetable in the United States, is not very popular in 

 Central America. The fruits are usually smaller than those produced 

 in the United States. 



Solarium molinarum J. L. Gentry, Phytologia 26: 276. 1973. 



Brushy rocky hillside or shaded quebrada, 350-1, 480 m.; Zacapa; 

 Chiquimula; Jalapa. Honduras (type from edge of Humuya River 

 bank, vicinity El Ede*n, Comayagua Valley, Comayagua, A. & 

 A. A. Molina). 



A small vine, the branches slender, puberulent, armed with numerous, short 

 and recurved prickles; leaves pinnate, 8-20 cm. long, pubescent with very short 

 hairs above and below, armed with prickles; petioles 1.5-4 cm. long; leaflets 5-7, 

 lanceolate to lance-ovate, 1.5-8 cm. long, the lowermost pair reduced, the terminal 

 leaflet larger than the lateral ones, the apex acuminate; inflorescence cymose, 

 lateral and internodal, paniculately disposed, few-flowered, rachis puberulent; 

 peduncle 3-5.5 cm. long, puberulent and armed with prickles; pedicels 7-8.5 mm. 

 long, sparsely and inconspicuously pubescent and unarmed; calyx unarmed; 

 2-2.5 mm. long, lobed about one-half way to the base, the lobes rounded, glabrous; 

 corolla white, the limb 12-16 mm. wide, lobed to below the middle, the lobes 

 5.5-7 mm. long, acute; stamens unequal, four of the filaments 1.5 mm. long, the 

 fifth one 2.5 mm. long ; the anthers 3.5 mm. long; style glabrous, 5 mm. long; ovary 

 glabrous; fruit globose, mottled with light and dark green. 2.2-2.8 cm. in diameter; 

 seeds 3.5-4.5 mm. long. 



Solatium morelliforme Bitter & Muench, Fedde Repert. Sp. 

 Nov. 12: 154, t. 2. 1913. 



Usually epiphytic on Quercus spp., sometimes on walls or boul- 

 ders, rarely terrestrial, 1,600-2,800 m.; Chimaltenango; Quezalten- 

 ango; Huehuetenango. East-central Mexico (type from Chiapas, 

 Gueytepec, Muench s.n.). 



Plants herbaceous, nearly always epiphytic, laxly ascending or pendent, usually 

 about 3 dm. tall, glabrous or essentially so, stoloniferous and tuber-bearing, the 

 tubers small, often numerous, globose or ellipsoid to obovoid, commonly purplish, 

 1-3 cm. long, 1-1.5 cm. in diameter; stem slender, purplish, simple or branched, 

 fistulose; leaves simple, ovate-lanceolate to narrowly lanceolate, 2.5-14 cm. long, 

 1-4 cm. wide, the apex acute to acuminate, often attenuate, the base oblique, 

 shortly attenuate to somewhat rounded; petioles 5-40 mm. long; pseudostipular 

 leaves present or absent; inflorescence pseudoterminal, becoming lateral and oppo- 

 site the leaves, laxly cymose, mostly 15-flowered; peduncle slender, 2-4.5 cm. long; 

 pedicels slender, 9-17 mm. long, articulate at about the middle; calyx 1.5-2 mm. 

 long, the lobes short, toothlike and acute; corolla white, the limb about 10 mm. 

 wide, parted to near the base, the lobes about 6 mm. long, lanceolate, acuminate, 

 ciliate; stamens equal, the filaments 1-1.5 mm. long, glabrous, often united to form 

 a column; anthers coherent, 3-5 mm. long, forming a column about the style, 




GENTRY & STANDLEY: FLORA OF GUATEMALA 129 



rounded and somewhat dorsally auriculate at their bases; style about as long as the 

 stamens; fruit globose, green or yellowish green, apiculate, 5-7 mm. in diameter. 



Solatium morelliforme Bitter & Muench is most likely to be 

 confused with S. clarum Correll from which it is distinguished by the 

 epiphytic habit, and the rounded and dorsally auriculate bases of 

 the anthers. 



Solanum muenscheri Standl. & Steyerm. Field Mus. Bot. 22: 

 275. 1940. 



Damp thickets or open rocky hillsides, sometimes in Juniperus 

 forests or in mixed forests, 2,500-3,500 m.; Chimaltenango; Solola 

 (type from near Santa Maria, Muenscher 12360)', Totonicapan ; 

 Huehuetenango ; San Marcos. Endemic. 



An unarmed shrub, 1-3 m. tall, the young branches, petioles, peduncles, 

 pedicels, and calyces densely pubescent, the hairs yellowish, mostly with rays 

 along their stalks (branched); leaves lanceolate to narrowly lanceolate, rarely 

 narrow oblong-lanceolate or narrowly elliptic, 6-15(-20) cm. long, l-2.5(-4) cm. 

 wide, the apex narrowly acute, the base rounded or obtuse, pubescent above, more 

 densely so below, velutinuous, the hairs branched and minute, the lateral veins 

 conspicuous; petioles 1-2. 5 (-4) cm. long; inflorescences pseudoterminal, becoming 

 lateral and opposite the leaves, cymose and paniculately disposed or racemose, 

 much shorter than the leaves, several-to many-flowered, short pedunculate to 

 nearly sessile; pedicels 8-10 mm. long; calyx 3.5-4 mm. long, parted to the middle, 

 the lobes lanceolate or ovate, acute; corolla purple to lavender, the limb 12.5-15 

 mm. wide, densely pubescent externally, parted to just below the middle, the 

 lobes 3.5-5 mm. long; filaments about 1 mm. long; anthers 2.5-3 mm. long; style 

 4.5-6 mm. long, the lower one-half densely covered with branched hairs; ovary 

 apically pubescent; fruit globose, 7-10 mm. in diameter, black, lustrous, glabrous 

 or sometimes sparsely pubescent apically; seeds 2-3 mm. broad. 



Solatium muricatum Ait. Hort. Kew. 1: 250. 1789. S. guate- 

 malense Hort., Bitter, Fedde Repert. Sp. Nov. 12 : 444. 1913. Pepino ; 

 petito. 



Cultivated in temperate and cool regions for its edible fruit, 

 occasionally escaping cultivation, 1,000-2,700 m.; Alta Verapaz; 

 Jalapa; Guatemala; Sacatepe"quez ; Quezaltenango. Mexico to Costa 

 Rica. Panama. Peru, Bolivia, Colombia, Ecuador, and Chile. 



An erect or ascending herb, sometimes with a woody base, up to 1 m. tall or 

 more, non-tuber-bearing; stem glabrous or nearly so, branched or sometimes sim- 

 ple; leaves simple (in ours), lanceolate to elliptic-lanceolate, mostly 8-18 cm. long, 

 1.5-3 cm. wide, the apex obtuse or acute to acuminate, the base rounded or shortly 

 attenuate, with appressed or subappressed hairs; petioles long, 3-7 cm. long; 

 inflorescences pseudoterminal, becoming lateral and internodal, few-to many- 

 flowered; peduncle forked once or simple, 5-8 cm. long; pedicels 5-15 mm. long, 




130 FIELDIANA: BOTANY, VOLUME 24 



articulate at or near the base; calyx 4-7 mm. long, divided to about or below the 

 middle, the lobes irregular and unequal, lanceolate, acute; corolla violet-purple 

 or white marked with purple, the limb 40 mm. wide or less, shallowly lobed, the lobes 

 broadly rounded, apiculate at the apex, externally strigose; stamens equal, the 

 filaments about 1.5 mm. long, glabrous; anthers 4.5-6 mm. long; style papillose, 

 5-9 mm. long, slightly exceeding the stamens; fruit ovoid to ellipsoid, pendent, 

 white or pale green and purple-variegated, up to about 10 cm. long and 6 cm. 

 in diameter. 



Solanum muricatum Ait. is believed to be a native of the Andean 

 region, but now is probably known only in cultivation or some- 

 times as an escape from cultivation. 



The green fruits are cooked and eaten, the ripe ones eaten raw. 

 Fruits of the cultivated plants are usually seedless. 



Distinguished from the other simple leaved species in our area 

 with short, oblong anthers and large apical pores, by the simple 

 hairs, habit, and the shallowly lobed corolla limb. 



Solanum nigrescens Mart. & Gal. Bull. Acad. Brux. 12, pt.l: 

 140. 1845. Hierba mora (Quezaltenango) ; macuy (Sacatepe'quez) . 



Wet thickets or forests or in mixed forests or on dry brushy 

 hillsides, sometimes a weed of cultivated fields, 1,500-3,900 m.; 

 Chiquimula; El Progreso; Sacatepe'quez; Solola; Quezaltenango ; Hue- 

 huetenango; Escuintla; San Marcos. Southern Mexico. Costa Rica. 



A sprawling or erect herb, 1-1.5 m. tall, sometimes 3.5 m., the young stems 

 pilosulous, sometimes sparsely so, the hairs spreading or incurved; leaves in pairs 

 or solitary, different in size, similar in shape, entire or sinuate-dentate, ovate to 

 ovate-lanceolate, rarely broadly ovate, the larger leaves 3-15(-18) cm. long, 

 1.5-6.5(-10.5) cm. wide, the apex acuminate to narrowly acute, the base attenuate, 

 pilosulous or villosulous above and below, sometimes sparsely so with age; petioles 

 5-35 mm. long; inflorescences lateral and internodal, racemiform or subumbelliform, 

 few-to several-flowered; peduncles 1-3 cm. long; pedicels 6-10 mm. long, fruiting 

 pedicels reflexed; calyx 1-1.5 mm. long, slightly accrescent in fruit, shallowly 

 lobate, the lobes ovate and acute or obtuse; corolla white or purplish with a dark 

 spot at the base of each corolla lobe, the limb 6. 5-10 (-16) mm. wide, parted to 

 near the base, the lobes (2.5-)3.5-4(-7) mm. long, externally pilosulous; filaments 

 0.5-l(-2) mm. long, ciliate; anthers (2.5-)3-4 mm. long; style (4-)5-5.5 mm. long, 

 exceeding the stamens, the lower one-half densely pubescent; ovary glabrous; 

 fruit globose, 4.5-7 mm. in diameter; seeds 1-1.5 mm. long. 



The material included in this species was determined by Heiser 

 (1955) as S. amethystinum (Kuntze) Heiser and S. leonii Heiser. He 

 also suggested at this time that S. amethystinum may prove to be 

 only a subspecies of S. douglasii Dunal. In a recent study of Ed- 

 monds (1972), S. douglasii is treated as a synonym of S. nigrescens. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 131 



A more intensive study might reveal the presence of a third 

 species, S. macrotonum Bitter, in Guatemala. 



See comments under S. americanum. 



Solatium nigricans Mart. & Gal. Bull. Acad. Brux. 12, pt. 1: 

 134. 1845. S. vernicinitens Bitter, Fedde Repert. Sp. Nov. 18: 58. 

 1922 (type from Coban, Alta Verapaz; Tuerckheim 7.45), syn. nov. 

 Aurella (Quezaltenango) . 



Wet thickets or dense wet forests, often in Abies or Cupressus 

 forests, sometimes in open pine-oak forests, 1,200-2,700 m.; Alta 

 Verapaz; Zacapa; Baja Verapaz; Jalapa; Guatemala; Chimaltenango; 

 Solola; Quiche" ; Totonicapan; Quezaltenango; Huehuetenango ; Suchi- 

 tepe"quez; San Marcos. Southern Mexico. Honduras. 



An unarmed shrub or small tree, 1-6 m. tall, the young branches, leaves, 

 peduncles, and pedicels densely covered with whitish or yellowish, closely ap- 

 pressed, branched hairs, the branching of the hairs discernible only with difficulty, 

 soon glabrescent; leaves solitary, firm, the lateral veins prominent, elliptic to 

 narrowly elliptic or elliptic-ovate, rarely ovate or obovate, (3-)6-15 cm. long, 

 (l-)2-5.5 cm. wide, the apex acuminate, the base short attenuate or cuneate; 

 petioles 5-15 mm. long; inflorescences lateral and opposite the leaves, cymose, 

 several-flowered; peduncles very short, 2-5 (-10) mm. long, sparsely pubescent or 

 glabrescent; pedicels 1-2 cm. long, sparsely pubescent or glabrescent; calyx densely 

 pubescent to glabrous, 1-1.5 mm. long, the lobes rounded and apiculate, obtuse; 

 corolla white, the limb 12.5-14 mm. wide, parted to near the base, the lobes 5-6 

 mm. long, papillate marginally and apically; filaments 0.5 mm. long; anthers 

 3-3.5 mm. long; style exceeding the stamens, 5.5-6 mm. long; ovary glabrous; 

 fruit globose, black, 1-1.5 cm. in diameter; seeds 3.5-5 mm. long. 



A common shrub of the western Highlands, sometimes forming 

 dense thickets. 



Solanum nigricans Mart. & Gal. has the younger stages of the 

 branches, leaves, and inflorescences densely covered with whitish or 

 yellowish, appressed hairs. The branches of the hairs are discernable 

 only with difficulty. 



Solanum nudum H.B.K. Nov. Gen. & Sp. PI. 3: 33. 1818. S. 

 parcebarbatum var. minorifrons Bitter, Fedde Repert. Sp. Nov. 18: 

 52. 1922 (type from near Finca Sepacuite, Alta Verapaz, Cook & 

 Griggs 14.6). Hediondilla (Quezaltenango); sakyol (Alta Verapaz). 



Damp or wet forests or thickets, sometimes in oak-pine forests 

 or wet pine forests, 1,000-2,500 m. or sometimes lower; Pet&i; Alta 

 Verapaz; Izabal; Baja Verapaz; Chiquimula; El Progreso; Guate- 

 mala; Sacatepe"quez; Chimaltenango; Quiche"; Quezaltenango; Hue- 




132 FIELDIANA: BOTANY, VOLUME 24 



huetenango; Santa Rosa; Escuintla; Suchitepe"quez ; San Marcos. 

 British Honduras. Southern Mexico. Honduras to Costa Rica. 

 Panama. 



An unarmed shrub, 1-3.5 m. tall, sometimes a tree of 6 m., the branches 

 sparsely pilosulous or glabrous; leaves solitary or in pairs, unequal in size, different 

 or similar in shape, the larger leaves narrowly elliptic to elliptic or ovate-elliptic, 

 sometimes ovate or rarely elliptic-oblong, 8-15(-20) cm. long, (2.5-)4-7 cm. wide, 

 the apex mostly acuminate or narrowly acute, the base short attenuate, glabrous 

 above, the hairs in tufts in the axils of the mid-vein, sometimes sparsely and 

 inconspicuously so, sometimes hairs along the margins of the mid-vein, rarely 

 glabrous below; petioles 1-1.5 cm. long; inflorescences lateral and opposite the 

 leaves, cymose, simple, few-to several-flowered; peduncles short, 1.5-2.5 (-10) 

 mm. long; calyx 1-2 mm. long, lobed to the middle, the lobes ovate or rounded, 

 and apiculate or shallowly lobate, glabrous or pilosulous apically, rarely sparsely 

 pilosulous; corolla white, the limb 8.5-10 mm. wide, parted to near the base, the 

 lobes 3-4 mm. long, inconspicuously ciliate with papillae; filaments about 0.5 mm. 

 long, sometimes connate one-half to nearly their entire length; anthers 2-2.5 mm. 

 long; style 4-5 mm. long, glabrous, exceeding the stamens; ovary glabrous; fruit 

 globose, 7-10 mm. in diameter, black; seeds 3-3.5 mm. long. 



Called yerba de barrer in British Honduras and chillo and cor- 

 doncillo in Honduras. 



This is a very common shrub in many parts of Guatemala being 

 most plentiful at middle elevations. The leaves are used in Alta 

 Verapaz by the Indians to treat sores. 



See comments under S. diphyllum L. 



Solarium oxycarpum Schiede in Schlecht. Hort. Hal. 1: 5, t. 

 3. 1841. 



Usually in rain or cloud forests, 2,500-3,000 m.; El Progreso 

 (Steyermark 43672}. Southern Mexico. Costa Rica. Panama. 



Plants herbaceous, ascending or nearly erect, 1.5-9 dm. tall, the stem glabrous 

 or sparsely pubescent, non-tuber-bearing or occasionally with tubers up to 6 cm. 

 long, 2 cm. in diameter; leaves odd-pinnate 6-26 (-32) cm. long; interstitial leaflets 

 absent or rarely with several minute ones; leaflets 7-10, rarely 5 or 13, ovate- 

 lanceolate to narrowly lanceolate or elliptic-lanceolate. 2-9 cm. long, 1-3.5 cm. 

 wide, the apex acute to long acuminate or rarely obtuse, the base broadly rounded 

 to cuneate, sessile or with a short-winged petiolule, the lowermost pair of leaflets 

 often much reduced; pseudostipular leaves present; inflorescence pseudoterminal, 

 becoming lateral and opposite the leaves, cymose, few-to many-flowered; peduncle 

 4-10 cm. long; pedicels slender, 1.5-2.5 cm. long, articulate at or somewhat above 

 the middle (rarely below the middle); calyx 3-7 mm. long, sparsely pubescent to 

 glabrous, lobed to about the middle, the lobes ovate to elliptic or narrowly lanceo- 

 late, acute to acuminate, rarely nearly obtuse; corolla pale or deep purple, occasion- 

 ally white, the limb 15-22 (-30) mm. wide, shallowly lobed, the lobes 2.5-4 mm. 

 long, densely pubescent; filaments 1-1.5 mm. long, glabrous or rarely puberulent; 




GENTRY & STANDLEY: FLORA OF GUATEMALA 133 



anthers 3-5 mm. long; style 5-8 mm. long, exceeding the stamens, puberulent 

 below; fruit ovoid-cylindric, acute, usually about 2 cm. long, 0.7 cm. in diameter. 



The inclusion of S. oxycarpum Schiede in the "Flora" is based 

 upon a collection (Steyermark 4-3672) from El Progreso. The speci- 

 men is in fruit and difficult to place but appears to be a form of this 

 species, Correll (1962). 



Solanum phaseoloides Polak. Linnaea 41: 585. 1877. S, 

 olivaeforme Donn.-Sm. Bot. Gaz. 14: 28. 1889. (type from Panasa- 

 mala, Alta Verapaz, Donn.-Sm. 1226). 



Wet thickets or moist forests, sometimes on dry slopes, 280- 

 1,500 m.; Alta Verapaz; Izabal; Quezaltenango ; Suchitepe"quez ; 

 Retalhuleu; San Marcos. British Honduras. Mexico (Chiapas). 

 Costa Rica. 



Plants herbaceous, creeping, rooting at the nodes, glabrous throughout or 

 essentially so; leaves trifoliolate and pinnate, long petiolate; leaflets 3, short 

 petiolulate, ovate to lanceolate or elliptic, 2-6.5 cm. long, 1-3.5 cm. wide, the apex 

 acute or obtuse, the base unequal, short-petiolulate, sparsely pilosulous, more 

 densely so on the veins or essentially glabrous above, glabrous below; pseud ostipular 

 leaves absent; inflorescences lateral and internodal or opposite the leaves, cymose, 

 3-7 flowered; peduncle (l-)2-4(-10) cm. long; pedicels slender, 1-2.5 cm. long; 

 calyx 1.5-2 mm. long, glabrous or pilosulous, parted to about the middle, the lobes 

 rounded or triangular; corolla white, the limb 10-16 mm. wide, parted to near the 

 base, the lobes 4-7.5 mm. long, ciliate and apically pilosulous, otherwise glabrous; 

 filaments 1 mm. long; anthers 2-2.5 mm. long; style 2.5-3.5 mm. long, equalling 

 to exceeding the stamens; fruit ovoid-conic, 2-3 cm. long. 



Solanum quinquangulare Willd. ex Roem. & Schult. Syst. 

 Veg. 4: 699. 1819. S. ciliatum sensu auct., non Lam. S. aculeatis- 

 simum sensu auct., non Jacq. Ixtu (Quecchi) Alta Verapaz. 



Moist thickets or wet forests, sometimes in pine forests or in 

 pastures, 1,260-1,800 m.; Pete"n; Alta Verapaz; Baja Verapaz; 

 Chimaltenango. Southern Mexico. Honduras. Nicaragua. Costa 

 Rica. 



Plants herbaceous or somewhat suffrutescent, 0.5-1.5 m. tall, the branches 

 pubescent with simple hairs, the hairs long and eglandular, and intermixed with 

 short glandular ones, armed with numerous, straight or somewhat reflexed prickles; 

 leaves armed with prickles, broadly ovate or round-ovate, shallowly lobed to 

 parted, mostly 10-30 cm. long, 7-20 cm. wide, the apex acute, the base cordate to 

 subcordate or nearly truncate, pubescent above with simple hairs, the hairs long, 

 moderately spaced, pubescent below with sessile-stellate and simple hairs, the 

 simple hairs commonly confined to the veins; petioles mostly 6-10 cm. long, the 

 hairs long and short glandular, armed with prickles; inflorescences lateral and 




134 FIELDIANA: BOTANY, VOLUME 24 



internodal, umbelliform, mostly 3-5 flowered; peduncles 2.5-20 (-35) mm. long, 

 sometimes subsessile, commonly armed with prickles, the hairs long and short 

 glandular; pedicels 6-10 mm. long, commonly armed with prickles, the hairs long 

 and short glandular; calyx 2-3 mm. long, sparsely pilose, prickly or unarmed, 

 parted about one-half way to the base, the lobes ovate; corolla white, the limb 

 11.5-15 mm. wide, parted nearly to the base, the lobes 4.5-6 mm. long, pubescent 

 with long hairs externally; filaments about 1 mm. long; anthers 4.5-5 mm. long; 

 style slender and glabrous, about 6.5 mm. long; ovary pubescent with simple 

 glandular and eglandular hairs; fruit red, globose, 1-1.5 cm. in diameter; seeds 

 compressed, 3-3.5 mm. wide, broadly winged. 



Solatium quinquangulare Roem. & Schult. and S. globiferum Dunal 

 can be easily separated by their very different seed morphology and 

 pubescence of their stems and petioles. The leaves of S. globi/erum 

 are much more densely pubescent and usually not as deeply lobed 

 as those of S. quinquangulare. 



Solanum ciliatum Lam., a species often confused with S. quin- 

 quangulare, has glabrous leaves below, large seeds and fruits, and 

 the stems only pilose, whereas in S. quinquangulare the leaves are 

 pubescent below with sessile-stellate and simple hairs, smaller seeds 

 and fruits and the stems hirsute or pilose and nearly always with 

 short glandular hairs. 



The specimens placed in this species have been identified in 

 herbaria as S. aculeatissimum and S. ciliatum. 



Solanum rovirosanum Donn.-Sm. Bot. Gaz. 48: 297. 1909. 

 S. nacoticosmum Bitter, Fedde Repert. Sp. Nov. 18: 55. 1922 (type 

 from above Jacaltenango, Huehuetenango, C. & E. Seler 2627), syn. 

 nov. S. schippii Standl. Publ. Carnegie Inst. Wash. 461: 85. 1935 

 (type from Machaca, British Honduras, Schipp 5584), syn. nov. 



Open forests, 15-350 m. ; Pete"n; Alta Verapaz. British Honduras. 

 Southern Mexico. 



An unarmed shrub, 1 m. tall, mostly glabrous throughout; leaves elliptic- 

 obovate to obovate or oblanceolate, rarely elliptic, mostly 10-27 cm. long, 4.5-12 

 cm. wide, the apex acute or sometimes acuminate, the base equal and short at- 

 tenuate, glabrous above and below; petioles 5-10 (-15) mm. long; inflorescences 

 opposite the leaves or internodal, cymose, furcate, several-flowered, inconspicuously 

 puberulent throughout; peduncles stout, 1-3 cm. long; pedicels short, 5-7 mm. 

 long, puberulent; calyx 2-2.5 mm. long, puberulent, the lobes short, 0.5 mm. long, 

 rounded and short-apiculate, obtuse; corolla white, the limb 12 mm. wide, parted 

 to near the base, the lobes 5 mm. long, pilosulous apically; filaments connate their 

 entire length, 1 mm. long; anthers 2.5-3 mm. long; style exceeding the stamens, 

 5-5.5 mm. long, sparsely puberulent near the base; ovary puberulent; fruit globose, 

 about 8 mm. in diameter, inconspicuously and sparsely puberulent. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 135 



Solatium rovirosanum Donn.-Sm. is closely related to S. fontium 

 Standl. & Steyerm. It is a plant of the Atlantic lowlands. 



Solanum nacoticosmum Bitter is tentatively treated here as a 

 synonym. 



Solanum rugosum Dunal in DC. Prodr. 13, pt. 1: 108. 1852. 

 Huevo de paloma. 



Moist or wet forests, in forest clearings and along river banks, 

 just above sea level to 165 m.; Izabal. British Honduras. Honduras 

 to Costa Rica. West Indies. Panama to Brazil. 



An unarmed shrub or small tree, 1-9 m. tall, the young branches sparsely 

 sessile stellate-pubescent; leaves entire, elliptic, obovate or oblong, 10-30 cm. 

 long, 4.5-9 cm. wide, the apex acuminate, the base long attenuate, decurrent along 

 the entire length of petiole or nearly so, asperous and sessile stellate-pubescent 

 above and below; petioles mostly 1-1.5 cm. long; axillary leaves absent; inflores- 

 cence cymose, pseudoterminal, becoming lateral, dichotomously branched, long 

 pedunculate, the peduncle 5-15 cm. long; pedicels 2-10 mm. long, the hairs sessile 

 stellate-pubescent; flower buds globose to nearly ellipsoid; calyx lobed to the mid- 

 dle, the lobes 1-2 mm. long, sparsely to densely stellate-pubescent internally, 

 densely stellate-pubescent externally; corolla white, the limb about 15 mm. wide, 

 parted to below the middle, the lobes about 6 mm. long; filaments 1-1.5 mm. long; 

 anthers 2.5-3 mm. long; ovary glabrous to sparsely stellate-pubescent; style 

 glabrous or sparsely stellate-pubescent; fruit yellow, about 1 cm. in diameter, 

 glabrescent or sparsely stellate-pubescent; seeds 1.5-2 mm. long. 



Solatium schlechtendalianum Walp. Repert. Bot. 3: 61. 

 1844-45. S. gentlei Lundell, Contr. Univ. Mich. Herb. 8: 84. 1942 

 (type from Swasey Branch, Monkey River, Toledo District, British 

 Honduras, Gentle 394^). Matej (Coban, Quecchi). 



Wet thickets or forests, sometimes in pine forests, also collected 

 in Manicaria Swamps (Izabal), just above sea level to 1,650 m.; Alta 

 Verapaz; Izabal; Baja Verapaz; Huehuetenango ; Santa Rosa. Brit- 

 ish Honduras. Southern Mexico. Honduras to Costa Rica. Pan- 

 ama. Cuba. 



A shrub or small tree, unarmed, 1.5-5 m. tall, the young branches, petioles, 

 peduncles, pedicels and calyces densely stellate-pubescent, the hairs grayish or 

 sometimes yellowish-brown, sessile and short stipitate-stellate; leaves solitary or in 

 pairs, then very unequal in size and different in shape, the larger leaves ovate to 

 ovate-elliptic or ovate oblong, mostly 7-16 cm. long, 3-7 cm. wide, the apex acumi- 

 nate, the base rounded or obtuse, rarely subcordate, sparsely to moderately sessile 

 stellate-pubescent and often asperulous above, densely stellate-tomentose and 

 grayish with sessile and short-stipitate hairs of two different lengths and velutinous 

 below; petioles short, 2-5 mm. long, rarely to 15 mm. long; smaller leaves broadly 

 elliptic to suborbicular, 1-3 cm. long, 1-2.5 cm. wide, obtuse; petioles 2-3 mm. 

 long; inflorescences pseudoterminal becoming lateral and opposite the leaves, 




136 FIELDIANA: BOTANY, VOLUME 24 



dichotomously cymose, sometimes simple, several-flowered; peduncles short, 

 1.5-3.5 cm. long; pedicels 4-8 mm. long; calyx 2-2.5 mm. long, parted to the middle 

 or scarcely lobed, the lobes rounded and short-caudate; corolla white, the limb 

 8.5-14 mm. wide, parted to just below the middle or divided to near the base, the 

 lobes 3-6 mm. long, densely stellate-tomentose externally; filaments short, about 

 0.5 mm. long; anthers 2-2.5 mm. long; style 3.5-6 mm. long, glabrous; ovary 

 stellate-tomentose; fruit globose, 7-8 mm. in diameter, black, sparsely stellate- 

 pubescent or essentially glabrous; seeds 2-2.5 mm. broad. 



Solatium seaforthianum Andr. Bot. Repos. 8, t. 504. 1808. 

 S. seaforthianum var. disjunctum Schulz in Urban, Symb. Ant. 6: 

 169. 1909. Colacidn (Santa Rosa); eolation blanca (Guatemala); 

 Idgrimas de la virgen (Peten). 



Cultivated as an ornamental in Guatemalan gardens and becom- 

 ing naturalized in wet or damp thickets at 850-1,400 m.; Peten; 

 Jalapa; Quezaltenango ; Huehuetenango ; Jutiapa; Santa Rosa; Re- 

 talhuleu. Southern Mexico. British Honduras. Honduras to Costa 

 Rica. Panama and West Indies. Venezuela, Colombia and Peru. 



An unarmed, woody vine, essentially glabrous throughout; leaves long petiolate, 

 odd-pinnate or pinnatifid, sometimes simple and entire, 5-12 cm. long; interstitial 

 leaflets absent; leaflets 5-7, lanceolate to ovate or elliptic, 1.5-6 cm. long, 0.5-2 cm. 

 wide, the apex acute or acuminate, sessile or short-petiolulate; pseudostipular 

 leaves absent; inflorescence pseudoterminal, becoming lateral and internodal or 

 opposite the leaves, laxly paniculate, many-flowered; pedicels 1-1.5 cm. long; 

 calyx 1 mm. long, glabrous, inconspicuously lobed, the lobes minute; corolla purple 

 or white, the limb 18-22 mm. wide, parted to near the base, the lobes 8-10 mm. 

 long, ciliate and pilosulous apically, otherwise glabrous; filaments 1.5-3 mm. 

 long, one slightly longer than the others; anthers 3-3.5 mm. long; style 8-10 mm. 

 long, much exceeding the stamens; fruit red, globose, 5-8 mm. in diameter; seeds 

 3-3.5 mm. long. 



Solanum seaforthianum Andr. is one of the common ornamental 

 vines of Central America. 



Called Josefina in El Salvador. 



Solanum skutchii Correll, Contr. Texas Res. Found. 1: 4, t. 1. 

 1950. Chumuchumutella (San Marcos). 



Wet forests or cloud forests or mixed forests, sometimes on stony 

 banks or along streams, 915-2,850 m.; Alta Verapaz; Jalapa; Guate- 

 mala; Chimaltenango; Quich6 (type from Nebaj, Skutch 1772); 

 Solola; Quezaltenango; San Marcos. Northeastern Mexico to Ni- 

 caragua. 



A climbing or trailing, non-tuber-bearing, woody vine, up to 7.5 m. long or 

 more, the stems rooting at the nodes, the young branches pubescent becoming 

 glabrous with age; leaves odd-pinnate, 2. 5-12 (-22) cm. long, pubescent to essen- 




GENTRY & STANDLEY: FLORA OF GUATEMALA 137 



tially glabrous above and below; petioles mostly 1-6 cm. long; interstitial leaflets 

 absent; leaflets 7-11, very rarely 5, elliptic to elliptic-oblong, elliptic-oblanceolate 

 or elliptic-obovate, sometimes lanceolate to elliptic-lanceolate, 1-9 cm. long, 0.5- 

 2 (-3) cm. wide, the apex acute to shortly acuminate, petiolule to 5 mm. long, the 

 lowermost pair of leaflets reduced, the terminal leaflet generally somewhat larger 

 than the lateral ones; pseudostipular leaves present; inflorescence almost always 

 axillary or pseudoterminal on leafy branches, racemose or cymose -paniculate; 

 peduncle 5 mm. long or less, rarely to 3.5 cm. long; pedicels (5-) 10-25 mm. long, 

 articulate at their bases, glabrous; calyx 2.5-4 mm. long, nearly truncate to shal- 

 lowly lobed, the lobes ovate, rounded and apiculate, conspicuously marginate; 

 corolla white, the limb 10-13 mm. wide, parted to near the base, the lobes 4-5.5 

 mm. long, elliptic-lanceolate, prominently reflexed at maturity, acute; filaments 

 about 1 mm. long, sparsely pubescent or glabrous, united for more than half their 

 length; anthers 3-4 mm. long, apiculate to aristate and puberulent at the apex; 

 style 8 mm. long, much exceeding the stamens; fruit ovoid to globose, orange-red to 

 orange, about 1.5 cm. in diameter. 



S. skutchii Correll is related to S. inscendens Rydb. 

 It is reported that the leaves are boiled and eaten as a pot-herb 

 in San Marcos. 



Solatium suaveolens Kunth & Bouche", Ind. Sem. Hort. Berol. 

 14. 1848. S. belizense Lundell, Mich. Univ. Herb. Contr. 7: 48. 1942 

 (type from near Vaca, El Cayo, British Honduras, Gentle 2319). 



In thickets and forested areas or in milpa clearings, usually below 

 1,000 m.; Pete"n. British Honduras. Southcentral Mexico. Costa 

 Rica. North and northwestern South America. 



Plants lax and straggly, usually about 4 dm. tall, more or less strigose through- 

 out, the stems weak, somewhat angular, apparently non-tuber-bearing ; leaves odd- 

 pinnate, 5-17 cm. long, densely strigose above and below; petioles 2-4 cm. long; 

 interstitial leaflets few or absent, subsessile to shortly petiolate; leaflets 7-13, 

 usually 9, lanceolate to elliptic-lanceolate, 2-9.5 cm. long, 0.5-2 cm. wide, the 

 apex acute to long acuminate, the base oblique, rounded to broadly cuneate, 

 sessile or with a slender petiolule, the lowermost pair of leaflets much reduced, 

 1 cm. long or less; pseudostipular leaves present; inflorescence lateral and inter- 

 nodal, racemose, several-flowered; peduncles simple, 4.5-10.5 cm. long; pedicels 

 filiform, 5-10 mm. long, articulate at their bases; calyx 2.5-3.5 mm. long, strigose 

 to nearly glabrous, divided to about the middle, the lobes ovate to ovate-lanceolate 

 or narrowly lanceolate, acute to acuminate; corolla white or yellowish, fragrant, the 

 corolla limb mostly 8-12 mm. wide, shallowly lobed; filaments 0.5-1 mm. long, 

 sometimes slightly connate; anthers about 3 mm. long; style about 3.5 mm. long, 

 slightly exceeding the stamens, puberulent; fruit ovoid to globose, green striped 

 with dark purplish green, about 2 cm. long, 1.5 cm. in diameter. 



Solatium suaveolens Kunth & Bouche" is readily distinguished 

 from S. fraxinifolium Dunal by the appressed hairs on the stem, 

 peduncle, pedicels, and calyx. The pubescence of S. fraxinifolium 

 is spreading. 




138 FIELDIANA: BOTANY, VOLUME 24 



Solanum tacanense Lundell, Amer. Midi. Nat. 29: 490. 1943. 



Creeping on tree trunks, 2,100-2,400 m.; San Marcos (Standley 

 68611}. Mexico (type from Volcan de Tacana, Chiapas, Matuda 

 2958). Apparently endemic in the mountains of southern Mexico 

 and western Guatemala. 



A woody or rarely somewhat herbaceous, unarmed vine, the stems rooting at 

 the nodes, non-tuber-bearing, the young branches densely pubescent, the hairs 

 brownish-yellow; leaves odd-pinnate, 2-14 cm. long, generally sparsely pubescent 

 above and below, more densely so on the veins; rachis densely pubescent with 

 brownish-yellow hairs; petioles 4.5 cm. long or less; interstitial leaflets absent; leaf- 

 lets 5, elliptic-lanceolate to elliptic or obovate-lanceolate, 1-6.5 cm. long, 0.5-2.5 

 cm. wide, the apex nearly obtuse to short acuminate, the petiolules subsessile to 1 

 cm. long; terminal leaflets larger than the lateral leaflets; pseudostipular leaves 

 present; inflorescence pseudoterminal on leafy branches, cymose-paniculate; pe- 

 duncle 3.5 cm. long or less, densely pubescent with brownish-yellow hairs; pedicels 

 5-7 mm. long, articulate at their bases, sparsely pubescent, the hairs spreading or 

 subappressed ; calyx about 2 mm. long, shallowly lobed, the lobes triangular, obtuse 

 and apiculate, about 0.5 mm. long; corolla probably white, the limb about 9 mm. 

 wide, parted to below the middle, the lobes about 3.5 mm. long, ovate to oblong- 

 lanceolate, obtuse to nearly acute; filaments about 1 mm. long, pubescent; anthers 

 2-2.5 mm. long; style about 2 mm. long, shorter than the stamens; fruit globose, 

 red, about 1 cm. in diameter. 



Solanum tacanense Lundell is closely related to S. connatum 

 Correll. It can be separated from that species by the shorter style 

 and the filaments only slightly connate at their bases. 



Solanum tequilense Gray, Proc. Am. Acad. Sci. 22: 441. 1887. 

 S. hirtum sensu auth., non Vahl 1798. Coyol de chucho; coyol de gato 

 (Retalhuleu), lava de platos (Santa Rosa). 



Wet thickets, sometimes on dry brushy plains or rocky hillsides, 

 near sea level to 900 m.; Pete"n; Izabal; Chimaltenango; Jutiapa; 

 Santa Rosa; Escuintla; Retalhuleu. British Honduras. Mexico to 

 Costa Rica. Panama. Ecuador. 



An herb or shrub, 1-1.5 m. tall, densely armed with straight and commonly 

 broad-based prickles, the young yellowish-brown branches, petioles, pedicels, and 

 calyces densely stellate-pubescent, the hairs sessile and short stipitate-stellate, some 

 hairs long stipitate-stellate; leaves solitary, sinuate -lobate to shallowly lobed, 

 broadly ovate to oblong or elliptic, 15-45 cm. long, 4-15 cm. wide, the apex acute or 

 obtuse, the base cordate to subcordate or obtuse, soft to the touch, the hairs sessile- 

 stellate, often with the central ray greatly exceeding the lateral ones above, densely 

 covered with yellowish or grayish, sessile to short stipitate-stellate hairs below; 

 petioles 4-15 cm. long, armed with prickles; inflorescences lateral and internodal, 

 subsessile, umbelliform; pedicels 8-10 mm. long; calyx about 7.5 mm. long, parted 

 nearly to the base, the lobes ovate, acute, sparsely sessile stellate-pubescent inter- 




GENTRY & STANDLEY: FLORA OF GUATEMALA 139 



nally, some hairs with the central ray greatly exceeding the lateral ones; corolla 

 white, the limb about 30 mm. wide, parted nearly to the base, the lobes 12 mm. 

 long, densely stellate-pubescent externally, some hairs long stipitate-stellate; an- 

 thers subsessile, about 8.5 mm. long; style 9 mm. long, glabrous; ovary densely 

 stellate-pubescent, the hairs with the central ray greatly exceeding the lateral ones; 

 fruit globose, orange-red to red, 2-2.5 cm. in diameter, densely stellate-pubescent; 

 seeds 3-3.5 mm. long. 



Called "wild tomato" in British Honduras and huevo de goto in 

 El Salvador. 



Solanum tequilense Gray is closely related to S. hirtum Vahl. It 

 has at least some of the hairs long stipitate-stellate on the young 

 branches, petioles, and pedicels, the young branches yellowish, 

 brown, the leaf lobes mostly 5-7 and the plants commonly with 

 broad-based prickles. In comparison, in S. hirtum Vahl the young 

 branches, petioles, and pedicels have sessile-stellate hairs with the 

 central ray glandular (on at least some of the hairs) and greatly ex- 

 ceeding the lateral ones, the young branches whitish or grayish, the 

 leaf lobes mostly 9 or more and the plants commonly with slender- 

 based prickles. There are a few intermediate specimens that are 

 difficult to place. 



See comments under S. hirtum Vahl. 



Solanum torvum Swartz, Prodr. Veg. Ind. Occ. 47. 1788. S. 

 ferrugineum Jacq. Hort. Schoenb. 3: 46, t. 334. 1798, syn. nov. 

 S. verapazense Standl. & Steyerm. Field Mus. Bot. 22: 276. 1940 

 (type from Santa Rosa, Baja Verapaz, Standley 69756), syn. nov. 

 S. mayanum Lundell, Contrib. Univ. Mich. Herb. 8: 85. 1942 (type 

 from ridge near Cowpen, Swasey Branch, Toledo District, British 

 Honduras, Gentle 3976), syn. nov. Huiz (Guatemala, Quezaltenango, 

 Sacatepe"quez) ; huevo de gato (Jalapa); lava platos (Huehuetenango, 

 Suchitepe"quez) ; pajch (Alta Verapaz). 



Dry or wet thickets, sometimes in pine-oak forests, often in 

 second growth, just above sea level to 1,500 m., rarely higher; Pete"n; 

 Alta Verapaz; Izabal; Zacapa; Chiquimula; Jalapa; El Progresso; 

 Baja Verapaz; Guatemala; Sacatepe"quez ; Chimaltenango; Solola; 

 Quiche" ; Quezaltenango; Huehuetenango; Jutiapa; Santa Rosa; Es- 

 cuintla; Suchitepe'quez ; Retalhuleu; San Marcos. British Honduras. 

 Mexico to Costa Rica. Panama and West Indies. South America. 



A shrub, 1-5 m. tall, the branches short stipitate-stellate and/or sessile stellate- 

 pubescent, the hairs yellowish-brown or sometimes reddish-brown, armed with 

 straight prickles, sometimes with a few slightly recurved ones, rarely unarmed; 

 leaves subentire to deeply parted, solitary or sometimes in pairs, similar in shape, 




140 FIELDIANA: BOTANY, VOLUME 24 



different in size, ovate to oblong or elliptic, 6-25 cm. long, 4.5-18 cm. wide, the 

 apex acuminate or acute, the base mostly unequal, rounded or cuneate, sometimes 

 truncate or subcordate, sessile stellate-pubescent above, the hairs short stipitate- 

 stellate and sessile-stellate below, sometimes prickly above and below; petioles 

 1.5-6(-7.5) cm. long; inflorescences lateral and internodal; peduncles l-1.5(-3) cm. 

 long, the hairs short stipitate-stellate and sessile-stellate; pedicels 6-12 mm. long, 

 the hairs mostly simple and sessile-stellate, often with the central ray of the 

 stellate hairs exceeding the lateral ones, at least some of the hairs glandular or 

 completely eglandular; calyx 3-6 mm. long, pubescence like the pedicels, the lobes 

 1-2.5 mm. long, linear or subulate, the calyx splitting in age, the lobes 2-5 mm. 

 long, lanceolate to ovate-lanceolate, the apex mucronate; corolla white, the limb 

 18-30 mm. wide, lobed to the middle or just below, the lobes 5-12 mm. long, 

 densely stellate-pubescent externally; filaments 1-1.5 mm. long; anthers 5.5-7.5 

 mm. long; style 8-11 mm. long, glabrous or sparsely glandular at the base; ovary 

 sparsely glandular; fruit globose, 10-14 mm. in diameter, the fruiting pedicels 

 erect or sometimes recurved; seeds 2-2.5 mm. broad, inconspicuously reticulate. 



This species is a very common weedy shrub of the Pacific lowlands 

 of Guatemala, as well as in other parts of the country. 



The Solanum torvum complex has proven to be one of the most 

 difficult groups in the preparation of this genus for the "Flora." We 

 have treated it as a single polymorphic species. The proper elucida- 

 tion of the "taxa" in this complex will require a more detailed 

 study. A study of this kind will probably necessitate some changes 

 in the present treatment. W. D'Arcy (personal communication) has 

 treated the elements of this complex, sensu stricto. Unrecognized 

 juvenile forms may be part of the problem in this complex (see K. 

 Roe, Sida 2(5): 381-385. 1966). 



The names, Solanum ochraceo-ferrugineum (Dunal)Fern. and S. 

 diversifolium Schlecht. non H. & B., have been applied to the 

 eglandular material placed in this species. 



Called susumba in British Honduras. 



Solanum trizygum Bitter, Fedde Repert. Sp. Nov. 11: 470. 

 1912. Candelaria (Quezaltenango; San Marcos); palo de agua (Hue- 

 huetenango); tzikin-ita (Huehuetenango). 



Moist or wet dense forests, sometimes along streams, 900-2,000 

 m.; Alta Verapaz; Baja Verapaz; Quezaltenango; Huehuetenango; 

 San Marcos. Costa Rica. Colombia and Venezuela. 



Plants herbaceous, erect, 1 m. tall or less, simple or sparsely branched, the 

 stem somewhat succulent, glabrous or nearly so; leaves odd-pinnate, 1.5-3 dm. 

 long; interstitial leaflets absent; leaflets 7-11, lanceolate to oblong-lanceolate or 

 elliptic, (5-)8-15 cm. long, 1.5-3.5 cm. wide, the apex long acuminate, the base 

 rounded to nearly cordate and unequal, short-petiolulate, sparsely pilosulous above, 




B 



D 



FIG. 19. Solanum torvum. A, habit and leaves, X Yi\ B, corolla dissected, 

 X 2; C, calyx and pistil, X 2; D, mature fruit, X 2. 



141 




142 FIELDIANA: BOTANY, VOLUME 24 



especially on the veins, glabrous below; pseud ostipular leaves absent; inflorescences 

 axillary, cymose, often in pairs, 7-9 flowered; peduncle 1 cm. long or less; pedicels 

 5-10 mm. long, articulate at their bases; calyx 1.5-2 mm. long, glabrous, parted to 

 the middle, the lobes rounded and short apiculate; corolla pale green, greenish- 

 yellow or white, parted to near the base, 10-12 mm. wide, the lobes 4-5 mm. long, 

 internally glabrous, ciliate and apically pilosulous; filaments about 1 mm. long; 

 anthers 2 mm. long; style 4-5 mm. long, exceeding the stamens; fruit ovoid-conic, 

 1.5-2.5 cm. long. 



Solanum trizygum Bitter can be separated from all other Solanums 

 with compound leaves in Guatemala by its axillary inflorescences, 

 ovoid-conic fruits, and lack of pseud ostipular leaves. 



Solanum tuberosum L. Sp. PI. 185. 1753, sensu cult. Caxlan; 

 papa; paps (Quecchi) ; puntas de papa; sis. 



The cultivated potato is a native of South America, probably 

 from the Andes of Bolivia and Peru. It was probably introduced 

 into Central America after the Spanish conquest. The potato is 

 grown in the mountains in Guatemala. A large part of the Guate- 

 malan potatoes are grown by the Indians, whose methods of cultiva- 

 tion are often considerably different from those practiced in the 

 United States. For instance, the plants are sometimes left for two 

 or three years, the tubers being removed as they form, without 

 destroying the plant. The quality of the potatoes varies greatly, but 

 in general they are small, and of poor quality when placed on the 

 table. Medium-sized potatoes can be purchased in some of the 

 markets, but the majority of the tubers are about as large as walnuts, 

 and many of them no larger than cherries. One can imagine the 

 large amount of time required to peel such small potatoes, for all of 

 them are peeled, not scraped. It is rarely, if ever, that they are 

 cooked with skins. 



Potatoes are one of the important food crops of Guatemala. 

 Costa Rica is the only other Central American country in which 

 they are grown in quantity. They are carried on men's backs from 

 the producing regions, especially by the Quiches of Totonicapan and 

 San Francisco El Alto, to remote parts of the country, such as 

 Guatemala, Coban, and Zacapa. Potatoes are said to be the only 

 crop grown in the highest inhabited parts of the Cuchumatanes. 



Starch extracted from potatoes is sold in small white balls or 

 cubes and is often mistaken for marshmallows by the tourists, to the 

 amusement of the market people. 



In Guatemala it is a common practice to gather the tender 

 sprouts of puntas de papa and use them as a pot herb. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 143 



Solanum tuerckheimii Greenm. in Donn.-Sm. Bot. Gaz. 37: 

 212. 1904. 



Wet forests, 200-600 m.; Alta Verapaz (type from Cubilqiiitz, 

 Tuerckheim 84.92) ; Huehuetenango. Endemic. 



An unarmed shrub, 2.5-3.5 m. tall, glabrous throughout; leaves solitary or 

 sometimes in pairs, unequal in size but similar in shape, the larger leaves narrowly 

 elliptic to elliptic-oblong, 10-20 cm. long, 2.5-7 cm. wide, the apex acuminate or 

 acute, the base unequal or equal, cuneate or short attenuate; petioles 1-2 cm. long; 

 inflorescences lateral and opposite the leaves or internodal, cymose, simple, 

 several-flowered; peduncles short and slender, 5 mm. long; pedicels slender, 

 1-1.5 cm. long; calyx 1.5-2 mm. long, lobed to the middle, the lobes rounded and 

 apiculate, obtuse; corolla probably white, the limb 7-8 mm. wide, parted to near 

 the base, the lobes 2.5 mm. long; filaments connate one-half their length or more, 

 0.5 mm. long; anthers 2-2.5 mm. long; style exceeding the stamens, 3.5 mm. long, 

 glabrous; ovary glabrous; immature fruit globose. 



Solatium umbellatum Miller, Gard. Diet. ed. 8. Solanum No. 

 27. 1768. Lava platos; mayte (Quecchi); tobaco de montana; tabaquillo. 



Dry or moist thickets or in fields or along river banks, 75-2,000 

 m.; Pete"n; Alta Verapaz; Izabal; Chiquimula; Jutiapa; Quezalte- 

 nango; Huehuetenango; Santa Rosa; Escuintla; Suchitepe"quez ; Re- 

 talhuleu; San Marcos. British Honduras. Mexico to Costa Rica. 

 Panama. West Indies. Northern South America. 



An unarmed shrub or small tree, 1-6 m. tall, the young branches covered with 

 sessile and subsessile-stellate hairs, some long stipitate-stellate hairs also present; 

 leaves entire, elliptic, lanceolate or oblanceolate, mostly 7-25 cm. long, 2.5-8 cm. 

 wide, the apex acuminate, the base long attenuate, decurrent along the entire 

 length of the petiole or nearly so, asperous and sessile stellate-pubescent above, 

 densely sessile and stipitate stellate-pubescent below; petioles short; axillary 

 leaves present or absent; inflorescence cymose, pseudoterminal, becoming lateral, 

 dichotomously branched, long pedunculate, the peduncle 3-12 cm. long; pedicels 

 3-6 mm. long, some of the hairs long stipitate-stellate; flower buds ellipsoid or 

 globose; calyx lobed to the middle or slightly below, the lobes lanceolate to oblong, 

 2-3 mm. long, glabrous or with a few stipitate-glandular hairs internally, partly 

 long stipitate stellate-pubescent externally; corolla white, the limb 10-15 mm. wide, 

 parted to below the middle, the lobes 4-6 mm. long; filaments 0.5-1 mm. long; 

 anthers 2-3.5 mm. long; ovary glabrous or sparsely stellate-pubescent; style 

 glabrous; fruit yellow, 8-11 mm. in diameter, glabrous; seeds 1.2-2 mm. long. 



Called friega-plato and palo de chincho in Honduras. 



Solanum wendlandii Hook. f. Bot. Mag. 113, t. 6914. 1887. 

 S. mazatenangense Coult. & Donn.-Sm. Bot. Gaz. 37: 421. 1904 (based 

 on Guatemalan material, 6 syntypes cited), syn. nov. S. unquis-cati 




144 FIELDIANA: BOTANY, VOLUME 24 



Standl. Field Mus. Bot. 4: 320. 1929 (type from Honduras), syn. nov. 

 Ixt&n (Pete"n, fide Lundell) Santa Rosa and Retalhuleu; Quixtdn 

 (Guatemala) ; Quishta (San Marcos). 



Wet or moist forests or thickets, sometimes in dry thickets, 

 200-2,850 m.; often cultivated for ornament; Pete"n; Zacapa; Chiqui- 

 mula; Guatemala; Sacatepe"quez; Quezaltenango; Jutiapa; Santa 

 Rosa; Retalhuleu; Suchitepe"quez; San Marcos. Southern Mexico to 

 Costa Rica. West Indies. Northern South America. 



A woody vine, the branches glabrous or rarely inconspicuously puberulent, 

 sparsely to densely armed with short, recurved prickles; leaves glabrous or rarely 

 inconspicuously puberulent above and below, mostly armed with prickles on the 

 mid-vein below, the upper leaves entire, ovate or elliptic, or 3-lobate, the apex 

 acuminate, the base rounded to short attenuate or the upper and lower leaves 

 pinnatifid or rarely pinnate, mostly 9-22.5 cm. long, 4.5-9.5 cm. wide, the petioles 

 long, mostly 3-5 cm. long, commonly armed with short, recurved prickles; in- 

 florescences pseudoterminal, becoming lateral and opposite the leaves, corymbose- 

 paniculate, several-flowered; peduncle (1.5-) 4-11 cm. long; pedicels 8.5-30 mm. 

 long, occasionally with one to few prickles; calyx unarmed, 3.5-9 mm. long, shal- 

 lowly lobed or deeply parted, the lobes rounded and mucronate or oblong, rounded 

 and mucronate at the apex, puberulent apically; corolla large and showy, lavender 

 to blue, the limb 35-55 mm. wide, very shallowly lobed to nearly entire, the lobes 

 abruptly acuminate, glabrous or sometimes puberulent externally; stamens 

 unequal, four of the filaments 1-1.5 mm. long, the fifth 2.5-4 mm. long, the four 

 shorter stamens with the anthers 6.5-9 mm. long, the longer stamen with the 

 anther 7.5-9.5 mm. long; style glabrous, 9.5-12 mm. long; ovary glabrous; fruit 

 ovoid, yellowish, 3.5-4 cm. in diameter. 



Solatium wendlandii can be distinguished from other members of 

 the genus in Guatemala by its anther shape, habit, broad and shal- 

 lowly lobed to nearly entire corolla limb, unequal stamens and nearly 

 always glabrous throughout. 



Solanum wendlandii Hook. f. is said to be introduced in most 

 Central American countries, being native only in Costa Rica. It 

 was, in fact, described from plants grown in England from seed sent 

 from Costa Rica. In Guatemala the plant is frequent in the central 

 and western mountain forests, but it is likely to be overlooked 

 because the flowers are among the branches of the taller trees. It is 

 common in cultivation in almost all parts of the country, and often 

 is found in hedges, where it has escaped from cultivation. The plant 

 is rather attractive because of the large and showy corollas that are 

 commonly produced in abundance. The large fruits are eaten and 

 the young shoots are sometimes cooked in stews. The local name, 

 Ixtan, is used as the name of a village in Retalhuleu. 



Called Elisa in El Salvador. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 145 



WITHERINGIA L'Heritier 

 Sicklera Sendtner, Flora 29: 178(194). 1846. 



Reference: A. T. Hunziker, Estudios sobre Solanaceae. V. Contri- 

 bution al conocimiento de Capsicum y generos afines (Witheringia, 

 Acnistus, Athenaea, etc.). Kurtziana 5: 101-179. 1969. 



Herbs or shrubs; leaves alternate, in pairs or sometimes solitary, entire or 

 sometimes angulately lobed, petiolate; inflorescences consisting of axillary fascicles 

 or sometimes in long-pedunculate, few-flowered clusters, the flowers pedicellate; 

 calyx campanulate, truncate, or 5-dentate, or 5-lobed, not at all to much accrescent 

 in fruit; corolla subrotate, narrowly campanulate, or funnelform, yellow or greenish- 

 yellow, or sometimes purple, the mouth of the corolla tube mostly pubescent, the 

 limb 5-parted, the lobes valvate or sometimes induplicate-valvate in bud; stamens 

 (4-) 5; filaments filiform or broad, exserted or included, mostly pubescent; anthers 

 oblong, longitudinally dehiscent, mostly apiculate; ovary bilocular, the ovules 

 numerous, the disc inconspicuous; style slender or sometimes broad, exserted or 

 included; stigma entire or emarginate; fruit a berry, globose or oblong, the fruiting 

 calyx little or not at all accrescent to much accrescent and surpassing the fruit; 

 seeds laterally compressed; embryo curved. 



A genus of some 15 species all in tropical America. Most of the 

 species are found in Central America. The work by Hunziker (1969) 

 greatly improved the generic limits of Athenaea and Capsicum. We 

 have accepted Hunziker's work for the most part. 



Inflorescences consisting of only 2-3 pedicellate flowers or the inflorescences long- 

 pedunculate; corolla throat glabrous, the filaments sometimes glabrous. 



Branches densely pubescent; inflorescences consisting of 2-3 pedicellate flowers, 

 peduncle absent W. physocalycia. 



Branches glabrous; inflorescences long-pedunculate W. cuneata. 



Inflorescences consisting of axillary fascicles; corolla throat and filaments densely 

 pubescent. 



Calyx truncate, the margin entire, scarcely or not at all accrescent in fruit. 



Plants pubescent, generally so throughout; larger leaves broad, mostly 1.5-2 

 times as long as wide; corolla mostly tetramerous W. solanacea. 



Plants glabrous throughout; larger leaves narrow, mostly 2.5 to 3 times as 

 long as wide; corolla pentamerous W. meiantha. 



Calyx 5-lobed or 5-dentate, little to much accrescent in fruit. 

 Leaves acute or obtuse to nearly truncate at their bases. . . . W. stramonifolia. 

 Leaves cordate at their bases. 



Pubescence of long, glandular hairs, at least some of the hairs 1 mm. long 

 or more; corolla marked with purple W. nelsonii. 



Pubescence of short, eglandular hairs, the hairs less than 1 mm. long; 

 corolla not marked with purple W. affinis. 



Witheringia affinis (Morton) A. T. Hunziker, Kurtziana 5: 

 162. 1969. Athenaea affinis Morton, Contr. Univ. Mich. Herb. 4: 

 24. 1940. Capsicum lundellii Morton, Contr. Univ. Mich. Herb. 4: 




146 FIELDIANA: BOTANY, VOLUME 24 



25. 1940 (type from Cohune Ridge, El Cayo District, British Hon- 

 duras, Lundell 6453}. 



In secondary forest, British Honduras (type from Cohune Ridge, 

 El Cayo District, British Honduras, Lundell 64-52}. 



A shrub, 2-6 m. tall, villousulous throughout with eglandular, multicellular 

 hairs; leaves in pairs, similar in shape, different in size, villousulous above and 

 below, the hairs more dense on the veins, the larger leaves broadly ovate, 17-20 

 cm. long, 9.5-11.5 cm. wide, the apex acuminate, the base cordate, the petioles 

 2-7 cm. long, villousulous; inflorescences consisting of 7-10 flowers, the pedicels 

 1.2-2 cm. long, villousulous; calyx about 3 mm. long, villousulous, shallowly lobed, 

 much accrescent in fruit; corolla pentamerous, narrowly campanulate, yellow, 

 8-12 mm. long, villousulous at the mouth of the corolla tube, the corolla limb 

 parted to near the base; stamens inserted below the middle of the corolla tube; 

 filaments villousulous at their bases, about 0.5 mm. long; anthers about 2 mm. 

 long, apiculate; berry globose, red, about 5 mm. in diameter, nearly included in 

 the accrescent calyx. 



Witheringia affinis is a poorly known species being represented 

 by only two collections. One of the collections represents the type 

 of Capsicum lundellii. The flowers of both W. affinis and C. lundellii 

 are very similar but the fruiting calyces are quite different. In W. 

 affinis the fruiting calyx is enlarged and encloses the fruit except at 

 the top, whereas in C. lundellii the fruiting calyx is not as strongly 

 accrescent. Capsicum lundellii has been placed in this species until 

 further collections are available for study. 



Witheringia cuneata (Standl.) A. T. Hunziker, Kurtziana 5: 

 118. 1969. Lycianthes cuneata Standl. Field Mus. Bot. 4: 260. 1929. 



In moist thickets, 30-150 m.; known in Guatemala from the 

 following collection, Rio Bonito, Cerro San Gil, Izabal, Steyermark 

 41657. Nicaragua, Costa Rica. Panama. Northern South America. 



A shrub, 2.5 m. tall, the branches glabrous; leaves entire, in pairs, glabrous 

 above, puberulent on the veins below, the paired leaves very different in size and 

 shape, the larger leaves elliptic to elliptic-oblong, 19-23 cm. long, 8.5-16 cm. wide, 

 the apex long acuminate, the base acute or rounded, the petiole short, 0.5 cm. long, 

 puberulent; smaller leaves elliptic, 7-14.5 mm. long, 8.5-16 mm. wide, subsessile; 

 inflorescence pedunculate, the peduncle 12 mm. long, glabrous, with 2 flowers, the 

 pedicels 7.5 mm. long, glabrous; calyx 1.5 mm. long, truncate, coriaceous, glabrous 

 except for the sparsely ciliolate margin; corolla pentamerous, funnelform or nar- 

 rowly campanulate, creamy white, 8.5 mm. long, the mouth of the corolla tube 

 glabrous, the corolla limb parted to near the base, glabrous externally; stamens 

 inserted at the mouth of the corolla tube; filaments glabrous, 0.5 mm. long; anthers 

 2.5 mm. long, minutely apiculate; style 5 mm. long; berry globose, dark violet, 

 about 8 mm. in diameter; seeds 3-3.5 mm. long. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 147 



Witheringia cuneata differs from the other Guatemalan species by 

 its pedunculate inflorescence, the paired leaves, very different in size 

 and shape and the glabrous corolla throat and glabrous filaments. 



The description was taken from only the cited Guatemalan col- 

 lection with the exception of the fruit and seeds. This collection 

 differs from the southern element of the species by the smaller leaves 

 of a pair being different in shape and wider than long, the leaves 

 puberulent on the veins below and the shorter corolla lobes. It is 

 tentatively placed in W. cuneata until further collections are available 

 for study. 



Witheringia meiantha (Donn.-Sm.) A. T. Hunziker, Kurtziana 

 5: 147. 1969. Brachistus meianthus Donn.-Sm. Bot. Gaz. 57: 424. 

 1914. Capsicum meianthum Standl. & Steyerm. Field Mus. Bot. 

 22: 273. 1940. Capsicum silvigaudens Standl. & L. Wms. Ceiba 3: 

 57. 1952. Witheringia solanacea L'Herit. var. silvigaudens A. T. 

 Hunziker, Kurtziana 5: 146. 1969, syn. nov. 



Moist or wet thickets or forests, sometimes in wet mixed forests, 

 1,200-1, 650m.; Alta Verapaz (syntype Pansamala, TuerckheimllSJi); 

 Baja Verapaz (syntype near Panzal, Tuerckheim 3936)', Quezalte- 

 nango; San Marcos. Honduras. Nicaragua. Costa Rica. Panama. 



An herb or shrub, 1-5 m. tall, glabrous throughout or nearly so; leaves entire, 

 in pairs, rarely solitary, glabrous above and below, the paired leaves different in 

 size and shape, the larger leaves narrowly elliptic to elliptic, 10.5-22 cm. long, 

 3. 5-6. 5 (-9) cm. wide, the apex acuminate, the base short-attenuate or obtuse, the 

 petioles 1-4 cm. long; smaller leaves ovate to ovate-elliptic, 4.5-8 cm. long, 

 2.5-3.5 cm. wide, the petioles 0.5-1 cm. long; inflorescences consisting of (2-)4-12 

 flowers, the pedicels 8-18 mm. long, glabrous; calyx (1.5-) 1.8-2 mm. long, truncate, 

 glabrous or sometimes ciliolate; corolla pentamerous, narrowly campanulate or 

 subrotate, greenish-yellow or yellow, 5-10 mm. long, densely villousulous at the 

 mouth of the corolla tube, the corolla limb parted to near the base, papillate- 

 puberulent externally; stamens inserted just below the mouth of the corolla tube; 

 filaments villousulous, 1 mm. long; anthers 2-2.5 mm. long, apiculate; style 5-7 

 mm. long; berry globose, red, 5-5.5 mm. in diameter; seeds 1.5-2 mm. long. 



Witheringia nelsonii (Fern.) A. T. Hunziker, Kurtziana 5: 

 160. 1969. Athenaea nelsonii Fern. Proc. Am. Acad. Sci. 35: 567. 

 1900. Capsicum viscidum Standl. Carnegie Inst. Publ. 461: 84. 1935 

 (type from Oaxactum, Pete"n, Bartlett 12690}. A. viscida (Standl.) 

 Standl. & Steyerm. Field Mus. Bot. 23: 18. 1943. Yerba moro 

 macho (Huehuetenango). 




148 FIELDIANA: BOTANY, VOLUME 24 



In wet forests, 500-2,000 m.; Pete"n; Alta Verapaz; Huehuete- 

 nango. Southern Mexico (type from between Tumbala and El 

 Salto, Chiapas, Nelson 3395). 



A shrub, 2.5-3.5 m. tall, densely villous throughout with glandular, multi- 

 cellular hairs; leaves entire, undulate or slightly sinuate lobed, mostly in pairs, 

 similar in shape, different in size, glandular-villous above and below, the larger 

 leaves broadly ovate, 14-28 cm. long, 7-14 cm. wide, the apex acuminate, the base 

 cordate, the petioles long, 4-9 cm. long, glandular-villous; inflorescences consisting 

 of (3-)5-8(-15) flowers, the pedicels 1.5-2 cm. long, glandular-villous; calyx 4.5-5 

 mm. long, glandular-villous, shallowly lobed, the lobes triangular, 1.5-2 mm. long, 

 acute or obtuse, the calyx much accrescent in fruit; corolla narrowly campanulate, 

 yellow, 10-16 mm. long, marked with purple, villousulous at the mouth of the 

 tube, the corolla limb parted to near the base; stamens inserted about the middle 

 of the corolla tube; filaments villousulous on the lower one-half, about 1.5 mm. 

 long; anthers 2-2.5 mm. long ; apiculate; style 5.5-6.5 mm. long; berry globose, 

 red, 6-8 mm. in diameter, included in the accrescent calyx; seeds 1-1.4 mm. long. 



A fruiting collection, Tun 24-6, from Pete"n has been tentatively 

 placed here until more material is available for study. It is very 

 similar to Witheringia nelsonii except that the fruiting calyx is not 

 strongly accrescent. 



The plants have a fetid odor. They are used in domestic medicine 

 in Huehuetenango. 



Witheringia physocalycia (Donn.-Sm.) J. L. Gentry, Phytolo- 

 gia 26:278. 1973. Brachistus physocalycius Donn.-Sm. Bot. Gaz. 40: 

 8. 1905. Athenaea physocalycia Standl. & Steyerm. Field Mus. Bot. 

 23: 18. 1943. 



Wet forests, 50-500 m.; Peteri; Alta Verapaz (type from Cubil- 

 qiiitz, Tuerckheim 8553). Endemic. 



A shrub, 1-1.5 m. tall, the branches densely pubescent, the hairs hirsutulous, 

 incurved or slightly so, at least some of the hairs greatly swollen at their bases; 

 leaves in pairs, very unequal in size and different in shape, glabrous above, hir- 

 sutulous below, the larger leaves inaequilateral, lanceolate to lance-oblong or 

 narrowly elliptic, 13.5-22 cm. long, 4-6.5 cm. wide, the apex narrowly long- 

 acuminate, the base acute; petioles 0.5-1.2 cm. long, hirsutulous; smaller leaves 

 nearly rotund to broadly ovate or elliptic-oblong, 1.5-6.5 cm. long, 1-3.5 cm. wide, 

 the apex obtuse; inflorescences consisting of 2-3 flowers, the pedicels 4-6 mm. long, 

 glabrous or sparsely hirsutulous; calyx 3 mm. long, much accrescent in fruit, very 

 sparsely hirsutulous, the lobes denticulate or the calyx nearly truncate; corolla 

 pentamerous, funnelform, white, 12 mm. long, glabrous, the lobes 3 mm. long; 

 filaments 3 mm. long, villousulous on the lower one- half or glabrous; anthers 2 mm. 

 long, apiculate; style 9 mm. long; berry globose, 7 mm. in diameter, nearly included 

 in the accrescent calyx; seeds 2 mm. long. 




GENTRY & STANDLEY: FLORA OF GUATEMALA 149 



Witheringia physocalycia is readily distinguished from the other 

 species in Guatemala by the characteristic pubescence on the 

 branches. 



This species was placed in Acnistus by A. T. Hunziker, in herb., 

 in 1962, but we feel that the correct placement is in Witheringia. 

 The generic limits of Acnistus require further study. 



Witheringia solanacea L'Herit. Sert. Angl. 33, t. 1. 1788. 

 Witheringia macrophylla H.B.K. ex Dunal, Solan. Syn. 1. 1816. 

 Sicklera solanacea Sendt. Flora 29: 179(195). 1846. Brachistus 

 solanaceus Hemsley, Biol. Centr.-Amer. Bot. 2: 424. 1882. Bassovia 

 macrophylla Coulter, Bot. Gaz. 16: 145. 1891. Solanum peckii Blake, 

 Contr. Gray Herb. 52: 87. 1917 (type from Monkey River, British 

 Honduras, Peck 585). Capsicum macrophyllum Standl. J. Wash. 

 Acad. Sci. 17: 16. 1927. Tomatillo (Quezaltenango) . 



Wet or moist thickets or moist forests, 1,400 m. and below; 

 Pete"n; Alta Verapaz; Izabal; Sacatepe"quez ; Chimaltenango; Que- 

 zaltenango ;Escuintla;Suchitepe'quez;Retalhuleu. Southern Mexico 

 to Costa Rica. Panama and West Indies. South America. 



Herb or shrub, 0.5-4 m. tall, the branches pubescent with simple, short multi- 

 cellular hairs (in ours) or glabrescent, rarely a few bifid hairs intermixed; leaves 

 entire, in pairs, rarely solitary, pubescent with short, multicellular hairs above and 

 below, sometimes sparsely pubescent, more dense on the veins below, the paired 

 leaves different in size, similar or slightly different in shape, the larger leaves 

 elliptic to ovate-elliptic or ovate, 12-32 cm. long, 5.5-12 cm. wide, the apex 

 acuminate, the base acute or obtuse, sometimes rounded, the petioles 2-7 cm. 

 long; smaller leaves 5.5-12 cm. long, 2.5-6.5 cm. wide, the petioles 0.5-1.5 cm. 

 long; inflorescences consisting of 10-40 flowers, the pedicels 8-12 mm. long, pubes- 

 cent with short, multicellular hairs; calyx 0.5-1.5(-2) mm. long, truncate, pubescent 

 with short, multicellular hairs; corolla mostly tetramerous or pentamerous, 

 narrowly campanulate or subrotate, yellow or greenish-yellow, 5-6 mm. long, 

 densely villousulous at the mouth of the corolla tube, the corolla limb parted to 

 near the base, papilla te-puberulent externally; stamens inserted above the middle 

 of the corolla tube; filaments villousulous, 1-1.5 mm. long; anthers 2-2.5(-3) mm. 

 long, apiculate; style 4.5-6 mm. long; berry globose, red, 4.5-6.5 mm. diameter; 

 seeds 1-1. 5 (-2) mm. long. 



This is the most wide-ranging species in Witheringia. Three 

 collections, two from Izabal, Steyermark 38209 and Jones & Facey 

 3259, and one from (Chiapas) Mexico, Clarke 303, are suggestive of 

 W. asterotricha. However, these specimens lack the branched hairs 

 of that species. 



Witheringia stramonifolia H.B.K. Nov. Gen. Sp. 3: 13. 1818. 

 Brachistus stramonifolius Miers, Ann. Mag. Nat. Ser. II, 3(16): 263. 




FIG. 20. Witheringia stramonifolia. A, habit, X K; B, fruiting inflorescence, 

 X 1^2; C, flower, X 3; D, part of dissected corolla showing stamens, X 4; E, calyx 

 and style, X 3. 



150 




GENTRY & STANDLEY: FLORA OF GUATEMALA 151 



1849. Capsicum stramonifolium Kuntze, Rev. Gen. 2: 450. 1891. 

 Bassovia donnell-smithii Coulter, Bot. Gaz. 16: 145. 1891 (syntypes, 

 Guatemala, Dept. Guatemala, Donn.-Sm. 2270; Duenas, Sacatepe"- 

 quez, Donn.-Sm. 2258). B. stramonifolia Standl. Contr. U.S. Natl. 

 Herb. 23: 1303. 1924. Athenaea locuples Standl. & Steyerm. Field 

 Mus. Bot. 22: 375. 1940 (type from above Finca El Porvenir, San 

 Marcos, Steyermark 37953). Chile cimarrdn (Chimaltenango) ; tolocic 

 Quecchi, (Alta Verapaz); tomatillo (Sacatep^quez). 



In moist or wet mixed forests or thickets, sometimes in cafetales, 

 300-2,000 m.; Pete"n; Alta Verapaz; El Progreso; Guatemala; Saca- 

 tepe"quez; Chimaltenango; Solola; Quiche 1 ; Quezaltenango ; Huehue- 

 tenango; Santa Rosa; Escuintla. Mexico. El Salvador. Honduras. 

 Costa Rica. Panama. 



Herb or shrub, 1-6 (-10) m. tall, the branches densely to sparsely covered with 

 short, multicellular hairs, the hairs erect or subappressed, rarely glandular; leaves 

 entire or often repand to shallowly angulately lobed, solitary or sometimes in pairs, 

 then similar in shape, different in size, the leaves lanceolate to broadly ovate, 

 7.5-30 cm. long, 2-15 cm. wide, sparsely to densely pubescent with short, multi- 

 cellular hairs, apex acuminate, base acute or obtuse to nearly truncate, the petioles 

 long, 2.5-9 cm., pubescent with short, multicellular hairs; inflorescences consisting 

 of 5-15 flowers, sometimes more, the pedicels 1-2.5 cm. long, pubescent with short, 

 multicellular hairs, glandular hairs rarely present; calyx 1.5-3 (-4) cm. long, 

 pubescent with short, multicellular hairs, glandular hairs rarely present, rarely 

 parted more than one-half way to the base, the lobes denticulate to broadly 

 deltoid, 0.2-1.5(-2.5) mm. long, little to much accrescent in fruit; corolla pentamer- 

 ous, narrowly campanulate to subrotate, yellow or sometimes greenish-yellow, 

 7-10 mm. long, densely villousulous at the mouth of the tube, the corolla limb 

 parted to near the base, the lobes papillate-puberulent internally, densely pubescent 

 with short, multicellular hairs externally; stamens inserted about the middle of the 

 corolla tube; filaments villousulous on the lower one-half, 1-1.5 mm. long; anthers 

 2.5-3 mm. long, generally not apiculate; style 5.5-6.5 mm. long; berry globose, 

 red, 5-5.5 mm. in diameter, the accrescent calyx rarely covering more than one- 

 half of the fruit; seeds 1-1.5 mm. long. 



Witheringia stramonifolia is a variable species, with the leaf 

 margin, calyx lobes, and the accrescent calyx showing the greatest 

 variability. It is often weedy, growing in cafetales and thickets. 

 The young leaves are sometimes cooked and eaten as a vegetable. 








.. 



Publication 1184 









UNIVERSITY OF ILLINOIS-URBANA