UNIVt-RSn I OF
ILLINOIS LIBRARY
AT URBANA-CHAMPAIGN
21: 1984'
.
FLORA OF GUATEMALA
DOROTHY L. NASH
Cucurbitaceae contributed by:
JENNIE V. A. DIETERLE
7
FIELDIANA: BOTANY
Volume 24, Part XI, Number 4
Published by
Field Museum of Natural History October 20, 1976
The Library of the
JAN 06 1977
University of Illinois at Urbant-Champaign
FLORA OF GUATEMALA PART XI
FLORA OF GUATEMALA
DOROTHY L. NASH
Research Specialist, Department of Botany Field Museum of Natural History
Cucurbitaceae contributed by:
JENNIE V. A. DIETERLE
University of Michigan
FIELDIANA: BOTANY
Volume 24, Part XI, Number 4
Published by
Field Museum of Natural History October 20, 1976
LAURA M. SCHLIVRK
Axsiii-inti' Editor
Library of Congress Catalog Card Number: 48-3076 US ISSN 0015-0746
PRINTED IN THE UNITED STATES OF AMERICA
We acknowledge the assistance of the National Science Foundation, program for Systematic Biology, given to the principal investigator, Louis O. Williams, over a period of many years. This most welcome assistance made possible both the field work and the research necessary to complete this work.
The National Science Foundation, Office of Science Information Service granted the principal investigator funds which will permit the publication of the remaining parts of the "Flora of Guatemala." The use of these funds began with the publication of this part of the flora. The Museum and the principal investigator are most appreciative of this financial aid.
CONTENTS Families in Part XI, Number 4
Caprifoliaceae 276
Valerianaceae 296
Dipsacaceae 306
Cucurbitaceae 306
Campanulaceae 396
LIST OF ILLUSTRATIONS
Page
1. Lonicera japonica 277
2. Sambucus mexicana 281
3. Symphoricarpos microphyllus 283
4. Viburnum hartwegii 291
5. Valeriana clematitis 299
6. Ahzolia composita 313
7. Anguria diversifolia 317
8. Cayaponia attenuata 319
9. Cionosicyos macranthus 323
10. Citrullus lanatus 325
11. Cucumis anguria 327
12. Cucurbita lundelliana 332
13. Cyclanthera langaei 335
14. Doyerea emetocathartica 339
15. Echinopepon horridus 341
16. Feuillea cordifolia 344
17. Gurania makoyana 346
18. Hanburia parviflora 348
19. Lagenaria siceraria 350
20. Luffa cylindrica 352
21. Melot hriapendula 357
22. Microsechium helleri 359
23. Momordica charantia 362
24. Parasicyos maculatus 364
25. Polyclathra cucumerina 366
26. Posadaea sphaerocarpa 368
27. Pterosicyos laciniatus 370
28. Rytidostylis macrophyllus 373
29. Schizocarpum filiforme 375
30. Sechium edule 376
31. Sicana odorifera 379
32. Sicydium glabrum 381
33. Sicyos galeottii 384
34. Tecunumania quetzalteca 388
35. Ahzolia composita (POLLEN) 390
36. Cayaponia attenuata (POLLEN) 390
37. Citrullus lanatus (POLLEN) 391
38. Cucumis melo (POLLEN) 391
39. Cyclanthera pedata (POLLEN) 392
40. Echinopepon horridus (POLLEN) 392
VII
VIII
41. Echinopepon torquatus ( POLLEN) 393
42. Luffa operculata (POLLEN) 393
43. Melothria pendula (POLLEN) 394
44. Momordica charantia (POLLEN) 394
45. Polyclathra cucumerina (POLLEN) 395
46. Schizocarpujn filiforme ( POLLEN) 395
47. Sphenoclea zeylonica 399
48. Triodanis perfoliata andCampanula rapunculoides 401
49. Burmeistera virescens 404
50. Centropogon cordifolius 406
51. Diastatea micrantha 410
52. Heterotoma lobelioides 413
53. Hippobroma longiflora 415
54. Lobelia nana and Lobelia laxiflora 424
55. Pratia calochlamys 430
Flora of Guatemala — Part XI, Number 4
CAPRIFOLIACEAE. Honeysuckle Family
By DOROTHY L. NASH
References: Karl Fritsch, Caprifoliaceae in Engler & Prantl, Pflanzenf. IV (4): 156-169. 1897. Camillo K. Schneider, Caprifo- liaceae, in Illust. Handbuch der Laubholzkunde 2: 632-753. 1912. A. M. Wilkinson, Floral anatomy and morphology of some species of the tribe Lonicereae of the Caprifoliaceae, Amer. Journ. Bot. 35: 261-271. 1948; Floral anatomy and morphology of some species of the tribes Linnaceae and Sambuceae of the Caprifoliaceae, torn. cit. 365-371; Floral anatomy and morphology of the genus Viburnum of the Caprifoliaceae, torn. cit. 455-465; Floral anatomy and morphology of Triosetum and of the Caprifoliaceae in general, op. cit. 36: 481-489. 1949. I. K. Ferguson, The genera of Caprifoliaceae in the southeastern United States, Journ. Am. Arb. 47(1): 33-59. 1966.
Shrubs or trees, often scandent, rarely herbaceous or suffruticose plants, the branches mostly terete, often nodose; leaves opposite, simple or pinnately compound; usually estipular (stipules present in Sambucus); inflorescences various, usually cymose, the flowers perfect, actinimorphic or zygomorphic, often bracteate; calyx tube adnate to the ovary, usually more or less contracted below the limb, the limb 3 5-dentate or lobate, the segments equal or unequal; corolla gamopetalous, rotate, tubular, funnelform, or campanulate, the tube often gibbous at the base, the limb regular or bilabiate, the 5 lobes commonly imbricate; stamens usually 5, sometimes 4, alternate with the corolla lobes, the filaments inserted on the corolla tube, equal or unequal; anthers oblong or linear, dorsifixed, versatile, bithecous, longitudinally dehiscent, usually introrse (extrorse in Sambucus); disc epigynous or none; style one or absent, the stigma capitate or as many as the carpels, sometimes sessile on the ovary; ovary inferior (not completely so in Sambucus and Viburnum), syncarpous, 1- 5-carpellate, locules 2-5, placentation axile or parietal; ovules usually solitary or few in each locule, pendulous, anatropous; fruit drupaceous or baccate, 1-5-locular (most commonly 4, in ours), containing 1-several seeds or stones; endosperm copious, carnose; embryo usually minute, straight.
The family is closely allied to the Rubiaceae and perhaps should be united with it; apparently there is no one character consistently separating the two families. However, there is also some evidence of the affinity of Viburnum and Sambucus with the Cornaceae.
275
276 FIELDIANA: BOTANY, VOLUME 24
About 18 genera, mostly in the northern hemisphere, but in America several species extend into the mountains of South America. Four genera are found in Central America.
Leaves pinnate; anthers extrorse; fruit with 3-5 stones Sambucus.
Leaves simple; anthers introrse; fruit with only 1-2 stones, or a berry.
Plants scandeht (in ours); corolla bilabiate; fruit a fleshy berry with few-several
seeds; in Guatemala cultivated plants Lonicera.
Plants erect; corolla campanulate or rotate; fruit drupaceous with 1-2 stones;
native plants.
Flowers solitary in the leaf axils or in axillary or terminal clusters; corolla tubular-campanulate; fruits white (in ours), the drupe containing 2 stones.
Symphoricarpos.
Flowers disposed in compound cymes, the inflorescences often appearing paniculate or umbelliform; corolla rotate or nearly so; fruits purple-black or red, the drupe containing 1 stone Viburnum.
LONICERA Linnaeus. Honeysuckle
Erect or scandent shrubs, glabrous or pubescent, often glaucous; leaves opposite or rarely ternate, sessile or short-petiolate, frequently connate at the base, entire or sometimes pinnate-lobate on young plants; inflorescence a 2-3-flowered cyme, usually axillary, bracteate, the flowers perfect, pentamerous, usually bracteolate; calyx tube adnate to the ovary, the limb short, 5-dentate or rarely truncate, deciduous or persistent; corolla tubular, funnelform, or campanulate, the tube short or elongated, sometimes gibbous at the base, the limb commonly oblique or bilabiate, sometimes obscurely bilabiate and appearing nearly regular, the lobes short or elongated, imbricate; stamens 5, usually exserted, the filaments inserted near the top of the corolla tube, the anthers oblong or linear, bithecous, introrse; ovary 2-3-locular (rarely 5-locular), the placentation axile or sometimes parietal in the upper part of the ovary; style filiform, the stigma capitate; ovules several in each locule; fruit baccate, usually red or black, rarely yellow or white, fleshy, the few seeds ovoid; endosperm carnose.
About 180 species in temperate and tropical areas, almost confined to the northern hemisphere. In America there are perhaps 25 species, mostly in the United States. Only one species, L. japonica Thunb., a native of eastern Asia, is found in Guatemala, but because L. pilosa (HBK.) Sprengel is found in nearby Chiapas, Mexico, it is included here.
Leaves directly below the inflorescence connate into a disc; corolla orange or reddish, 4-5.5 cm. long, the tube gibbous at the base, the limb obscurely bilabiate and appearing nearly regular; berries red L. pilosa.
Leaves directly below the inflorescence not connate; corolla white or pinkish, changing to yellow in age, the tube slender, not gibbous at the base, the limb distinctly bilabiate; berries black L. japonica.
NASH: FLORA OF GUATEMALA
277
FIG. 1. Lonicera japonica. A, habit, x Vfe; B, buds terminating a branchlet, x l'/2; C, flowers in natural position, X 1; D, flower dissected, X \l/i. (C and D after Botanical Register 1: t. 70.)
Lonicera japonica Thunb. Fl. Jap. 89. 1784. Madreselva; xian; raxcam (Quecchi, Coban, Alta Verapaz); Japanese honeysuckle.
Planted commonly for ornament from sea level up to 2,500 m. and perhaps higher; naturalized rather abundantly in Alta Verapaz in some localities about Coban and Tactic. Native of eastern Asia, but planted generally in temperate and tropical regions; extensively naturalized in the southeastern United States.
Small or large woody vines, the younger stems pubescent to pilose with spreading hairs; leaves short-petiolate, ovate to oblong, mostly 2-7 cm. long, acute, rounded or broadly cuneate at the base, entire, glabrous above, usually pubescent beneath, at least on the costae; inflorescences arising in the upper leaf axils, short-pedunculate, the 2 flowers subtended at the base by leaflike, ciliate bracts; calyx tube glabrous, the 5 teeth narrowly triangular to almost linear, ciliate, usually long-ciliate at the apex; corolla usually white, sometimes pinkish, changing to yellow in age, 2-4 cm. long, the tube slender, the limb bilabiate, somewhat shorter than the tube; stamens exserted; berries black.
These ornamental vines with very fragrant flowers are very common in many parts of Central America. About Coban there are dense tangles of vines in thickets and hedges, where they have escaped. In and about San Marcos, there are handsome, dense hedges of considerable length that have been planted. In the southeastern and eastern United States this species has become a great pest, spreading through woods over wide areas and driving out other vegetation.
278 FIELDIANA: BOTANY, VOLUME 24
Lonicera pilosa (HBK.) Sprengel, Syst. Veg. 1: 758. 1825. Caprifolium pilosum HBK. Nov. Gen. & Sp. 3: 427, t. 298. 1819. L. tubulosa Benth. PI. Hartweg. 37. 1840. L. pilosa f. schaffneri Rehder, Rep. Mo. Bot. Card. 14: 172. 1903. L. pilosa f. tubulosa Rehder, I.e.
A Mexican species, not reported from Guatemala, but as it has been twice collected in Chiapas (Ghiesbreght 699 and Laughlin 1092) as well as in northern Mexico, it may be expected in Guatemala. In Mexico, it is often found in openings in oak and pine forests, 1,200-2,800 m.
Scandent or subscandent, woody plants, when young sometimes appearing more shrubby than scandent but later climbing, the young stems glabrous or somewhat pilosulous; leaves opposite or rarely some or all of them ternate, short-petiolate or sessile, at least those directly below the inflorescence connate into a disc, the blades of other leaves ovate, ovate-oblong, or oblong-oval, mostly 3-7 cm. long, acute or obtuse and apiculate, rounded or subacute at the base, the margins entire, ciliate or not, glabrous above, more or less pilose or glabrous beneath or sometimes villous or pilose only along the costae; inflorescences terminal, pedunculate, spicate or headlike, the flowers disposed in sessile, 1-3-flowered cymes, the bracteoles about half as long as the ovaries; calyx tube glabrous or somewhat pubescent especially near the apex, sometimes more or less glandular, the 5 teeth short, triangular to ovate, glabrous or rather long-hirsute and/or long-ciliate; corolla orange or reddish outside, orange or yellow within, 4-6 cm. long, the tube gibbous at the base, sparsely or rather densely pilose outside or almost glabrous, the limb obscurely bilabiate, the lobes less than 1 cm. long; stamens and style short-exserted; berries red.
The leaves of all specimens examined are uniformly opposite with three exceptions: Laughlin 1092 from Chiapas, Municipio of Ixtapa, has ternate leaves; Nelson 4471 from Tamaulipas, Mexico (identified previously as L. ciliosa Poir.) and Waterfall & Wallis 13639 from Durango, Mexico, both with essentially glabrous corollas, have both opposite and ternate leaves. However, as these specimens match in all other respects with L. pilosa f. schaffneri Rehder, I do not feel justified in naming a new form or variety.
The form with obtuse leaf blades and hirsute calyx segments is apparently most common in Chiapas.
SAMBUCUS Linnaeus
Reference: F. von Schwerin, Monographic der Gattung Sam- bucus, Mitt. Deutsch. Dendrol. Ges. 18: 1-56. 1909; Revisio generis Sambucus, ibid. 29: 194-231. 1920.
Shrubs or trees (rarely herbaceous), the branches stout, with large pith; leaves opposite, odd-pinnate, with or without stipules, the leaflets serrate, with or without
NASH: FLORA OF GUATEMALA 279
stipels; inflorescence a terminal, compound cyme, becoming corymbiform or paniculate; calyx adnate to the ovary, the lobes 5, minute or nearly obsolete; corolla rotate, the 5 lobes suborbicular, ovate, or oblanceolate; stamens about equalling the corolla lobes (sometimes slightly shorter or longer than the lobes), the anthers oblong, the thecae distinct, extrorse; style very short, the stigma 3-5-lobate; ovary inferior, 3- 5-locular (most often 4-locular), the ovules pendulous in each locule, the placentation axile; fruits berrylike, juicy, globose or nearly so, containing 3-4 stones.
A genus of temperate zones and mountainous regions of the tropics of both hemispheres. More than 40 species have been named; many are very closely related and it is doubtful that even half that number are valid. There is practically no variation in floral characters, most species being distinguished by leaf shape, size, and/or indument. Some authors have used the size of the inflorescence and the length of the stamens in relation to the corolla lobes as distinguishing characters, but neither is reliable, both often varying considerably on the same plant. Only two species are found in Guatemala, and there are forms appearing intermediate between these.
United States authors have referred to Sambucus as "deci- duous," and in the colder regions this is strictly correct. However, the leaves do not color in autumn as do those of most deciduous trees, but remain green until killed by frost. Dr. Standley has noted in manuscript that in southern New Mexico where the winters are mild, Sambucus plants retain their leaves through the cold season and in the tropics they have leaves throughout the year.
The wood of the genus is light but not durable. When the large pith is removed from pieces of stem, the remaining tubes are often used by children for making flutes, whistles, and pop guns. The fruits of both of the following species are eaten by birds and are used in many localities to make wine and jelly.
Leaflets mostly 3.5-9 cm. long with 5-7 pairs of lateral veins, usually cuneate or acute at the base, the serrate margins commonly with 5-30 teeth on each side, the lowest leaflets usually trifoliate with the lateral ones reduced S. mexicana.
Leaflets mostly 6-19 cm. long, with 7-13 pairs of lateral veins, usually obtuse or rounded at the base and often unequal or oblique, the margins closely serrate, commonly with 30-90 teeth on each side, the lowest leaflets occasionally trifoliate S. canadensis.
Sambucus canadensis L. Sp. PI. 269. 1753; Torrey & Gray, Fl. N. Am. 2: 13. 1843; Fernald, Gray's Manual of Botany, 8th ed.: 1342. 1950. S. oreopola Donn.-Sm., Bot. Gaz. 25: 146. 1898. S. canadensis var. oreopola Rehder in Sargent, Trees & Shrubs 2 : 188.
280 FIELDIANA: BOTANY, VOLUME 24
1911. S. canadensis var. submollis Rehder, I.e. Sauco; sauco bianco; tlacha (Quezaltenango), tunalj (Sacatepequez).
Damp thickets and forest or in open places, abundant in hedges and along roads, 1,300-3,700 m.; Alta Verapaz; Chimaltenango; Chiquimula; Huehuetenango; Jalapa; El Progreso; Quezaltenango; El Quiche; Sacatepequez; San Marcos; Solola; Totonicapan. Nova Scotia to Manitoba, Canada, south through middle and eastern United States to Florida, west to Oklahoma and Texas; Mexico; El Salvador; Costa Rica; Panama.
Large shrubs (to about 4 m. in north temperate zones) or often small trees, 5-12 m. tall (in the mountains of tropical and subtropical regions), the trunk short and thick, the bark light brown, shallowly furrowed, often streaked with green, the wood yellow, the pith white; leaflets 5-11 (most commonly 7-9), sessile or short-petiolulate, usually oblong- lanceolate, sometimes lance-ovate or elliptic, mostly 6-19 cm. long, with 7-13 pairs of lateral veins, acuminate or long-acuminate, usually obtuse or rounded at base and often unequal or oblique, the margins closely serrate, commonly with 30-90 teeth on each side (occasionally only about 25), the lowest leaflets sometimes trifoliate, glabrous above, glabrous or hirtellous to densely short-hirsute or canescent beneath; inflorescences corymbiform, flat, mostly 10-30 cm. in diameter, the flowers fragrant, the pedicels glabrous or pubescent; corolla white or cream- colored, 5-7 mm. broad, the lobes rounded; mature fruits purple-black (in ours), 5-8 mm. in diameter.
Highly variable as to leaf form and size, this species is apparently closely related to the European S. nigra L. In the north temperate regions, forms sometimes occur with fruits red or orange or, rarely, greenish.
When these plants become arborescent, the branches are often large and heavy so that they spread horizontally or even rest on the ground. They are often planted for hedges.
Sambucus mexicana Presl ex A. DC. in DC. Prodr. 4: 322. 1830. S. bipinnata Schlecht. & Cham. Linnaea 5: 171. 1830. S. mexicana var. bipinnata Schwerin, Mitt. Deutsch. Dendr. Ges. 1909: 34, 328. 1909. S. simpsonii Rehder in Sargent, Trees and Shrubs 2: 187, t. 175. 1911. Aralia sololensis Donn.-Sm., Bot. Gaz. 56: 58. 1913. Sauco; sauco Colorado; sauco extran/ero (Ostuncalco, Quezaltenango); sacatsun (Quecchi, Alta Verapaz); bahman (Soloma, Huehuetenango, fide Standley); tzolokquen (Suchitepe- quez, fide Standley); tzoloj, tzolojche, tzolojque (El Quiche, fide Standley).
Planted commonly near houses, in hedgerows, and in gardens, from sea level to 3,000 m.; native of central and southern Mexico (or perhaps known only in cultivation?); probably not native
NASH: FLORA OF GUATEMALA
281
FIG. 2. Sambucus mexicana. A, habit, X Vz; B, part of fruiting inflorescence, X '/2j C, fruits, one dissected, X 2Vfc; D, flowers, one with corolla removed, X 6; E, stamen, x 7V4.
anywhere in Central America but widely planted throughout the area and in South America; also found in the Dominican Republic and in low, wet places in Georgia, Louisiana, Florida, and Texas.
Shrubs or small trees, mostly 3-5 m. tall, essentially glabrous throughout, or with some pubescence usually confined to leaf petioles and costae; the bark light brownish gray, furrowed and somewhat scaly; leaflets usually 5-7, sessile or short-petiolulate, the lowest ones frequently trifoliate with the lateral ones reduced, ovate, ovate- lanceolate, oblanceolate, or elliptic-oblong, mostly 3.5-9 cm. long with 5-7 pairs of
282 FIELDIANA: BOTANY, VOLUME 24
lateral veins, short-acuminate, usually cuneate or acute at the base, the margins serrate commonly with 5-30 teeth on each side; inflorescence corymbiform, convex or flat, 6-20 cm. in diameter; corolla white, 5-7 mm. in diameter, the lobes rounded; mature fruits purple-black, 5-8 mm. in diameter.
Widely planted in Guatemala, where an infusion or decoction of the flowers is a popular remedy for colds.
The department of Solola derives its name from that of S. mexicana, according to the account in Dice. Geogr. de Gautemala 2: 300. 1962, as follows: "....desde tiempos inmemorables, esta conocida con las designaciones de Tzoloj-ja en Quiche y Tzoloj-ya en Cakchiquel y Zutujil, que de ambos modos significa agua de saiico (Sambucus mexicana} deja o ya = agua; tzoloj = sauco."
SYMPHORICARPOS Duhamel
Reference: G. N. Jones, A monograph of the genus Sym- phoricarpos, Journ. Arn. Arb. 21: 201-252. 1940.
Shrubs, often densely branched; leaves opposite, short-petiolate, the blades small but broad, entire or sometimes sinuately dentate or lobate; stipules none; flowers small, perfect, solitary or in axillary or terminal clusters; calyx small, shallowly cupuliform, 4-5-dentate; corolla campanulate or tubular-funnelform, 4-5-lobate, regular or nearly so; stamens 5 or 4, the filaments inserted on the corolla tube, included or short-exserted, the anthers oblong, the thecae distinct, introrse; style one, slender, the stigma capitate or shallowly bilobate; ovary inferior, with 2 fertile, uniovulate locules and 2 sterile, several-ovulate ones, the placentation axile; fruit a berrylike drupe containing 2 stones; seeds with copious endosperm, the embryo minute.
Sixteen species in North America and one in China. Only the following reaches Central America. Some United States species are cultivated because of their ornamental fruits.
Symphoricarpos microphyllus HBK. Nov. Gen. & Sp. 3: 424. 1820; Jones, Journ. Arn. Arb. 21: 233. 1940. Anisanthus micro- phyllus Willd. in Roem. & Schult. Syst. Veg. 5: 223. 1819. Symphoricarpos glaucescens HBK. Nov. Gen. & Sp. 3: 424. t. 295. 1820. S. montanus HBK. torn. cit. 425. t. 296. Symphoria microphylla Sprengel, Syst. Veg. 1: 757. 1825. S. glaucescens Sprengel, I.e. S. montana Sprengel, I.e. Descliaea leucocarpa Sesse & Mocino ex DC. Prodr. 4: 483. 1830, pro. syn. Margaris barbigera. Margaris nudiflora DC. I.e. M. barbigera DC. I.e. Chiococca axillaris Sesse & Mocino, PI. Nov. Hisp. 36. 1887. Ac-cul, chipilin de monte (Huehuetenango).
FIG. 3. Symphoricarpos microphytlus. A, a branch, x 1; b, tip of flowering branch, X 2; C, flower with subtending bract, X 4'/2; D, flower dissected, corolla with style, calyx and hypanthium, subtending bract, X 4V6; E, fruits, one dissected, X 3.
283
284 FIELDIANA: BOTANY, VOLUME 24
Usually in dense, mixed forest, sometimes in oak, pine, or Juniperus forest, or on open, rocky, brushy slopes, 2,100-3,800 m.; Chimaltenango; Huehuetenango; El Progreso; Quezaltenango; Sacatepequez; San Marcos; Totonicapan. New Mexico; Mexico; Honduras.
Erect shrubs, 1-3 m. tall, usually densely branched, the young twigs puberulent to almost tomentulose; leaves usually short-petiolate, sometimes subsessile, the blades ovate to broadly elliptical or almost orbicular, mostly 5-20 mm. long, obtuse or rounded and apiculate at the apex, rounded or acute at the base, glabrous or puberulent above, pubescent to short-pilose or glabrate beneath or the indument confined to costae and veins, very finely reticulate-veined, the lateral veins (3-) 5-7 pairs; flowers solitary or paired, bibracteate; calyx teeth 5, broad, subacute or obtuse, sometimes ciliate; corolla narrowly tubular-campanulate, 6-10 mm. long, glabrous outside, white or tinged with pink or purple, white and pubescent within, the 5 lobes about equal, one-fourth to one-third the length of the tube; stamens 5, slightly exserted; style glabrous; fruit globose, juicy, white, about 8 mm. in diameter, containing 2 nutlets about 3 mm. long.
Common shrubs at many places in the higher mountains, especially in Juniperus forests of the Cuchumatanes. In the open the bushes are low and often very dense, in the forest taller and more open. They are often browsed by stock, probably goats and sheep.
VIBURNUM Linnaeus
Reference: C. V. Morton, The Mexican and Central American species of Viburnum, Contr. U. S. Nat. Herb. 26: 339-366. 1933.
Trees or shrubs, glabrous or pubescent; leaves opposite or rarely ternate, petiolate, the blades pinnately or rarely palmately nerved, the margins entire or dentate; stipules usually none; inflorescence a compound, bracteate cyme, usually pedunculate and appearing umbelliform (in ours), the flowers small; calyx tube often glandular or pubescent, the lobes 5, usually persistent in fruit; corolla white or cream-colored, campanulate or rotate, the 5 lobes imbricate in bud; stamens 5, the filaments inserted on the base of the corolla tube, the anthers versatile, longitudinally dehiscent; ovary inferior, appearing unilocular (one large fertile locule and 2 uppermost, abortive locules), the placentation parietal, the fertile ovule anatropous, pendulous from near the apex of the locule; style thick and short, the 3 stigmas capitate; fruit a red or black drupe, the stone often grooved.
About 140 species, widely distributed, chiefly in temperate regions; in the tropics confined to the mountains. In addition to the 13 species treated here, a few others are found in southern Central America.
As the result of study of a large series of specimens, some of the characters of pubescence of calyx, corolla, and foliage often relied
NASH: FLORA OF GUATEMALA 285
upon by Morton in the above-cited reference were found not to be dependable. As more material is collected, it is quite possible that some species now recognized will have to be reduced to synonymy.
Probably the fruit of all Guatemalan species is edible, although we have no information regarding any use made of it. In the United States the fruits are sometimes used to make wine, jams, and jellies, and are a favorite food of many birds. The wood of the genus is pale brown to yellow, usually straight-grained. Apparently, no use is made of it in Guatemala, unless for firewood, and the largest trees are too small to have commercial importance.
Leaf blades broadly rhombic to very broadly ovate or suborbicular, the margins conspicuously and usually rather coarsely dentate, the teeth mostly 10-26 on each side.
Lower leaf surfaces more or less stellate-pubescent to stellate-tomentose. Indument appearing whitish, the stellae about 0.3 mm. in diameter.
V. euryphyllum. Indument appearing brownish or yellowish, the stellae 0.5-0.9 mm. in diameter.
V. jucundum. Lower leaf surfaces glabrous or nearly so, appearing greenish, the hairs mostly
simple or paired, confined to costae and veins V. jucundum var. detractum.
Leaf blades ovate, lance-ovate, or oblong-ovate to lance-oblong or elliptic, the
margins entire or sometimes remotely glandular-dentate, the teeth when present
usually less than 10 on each side (rarely some leaves minutely serrulate in V.
elatum).
Lower leaf surfaces commonly densely stellate-tomentose or stellate-hirsute (rarely
only sparsely so).
Leaf blades mostly ovate to obovate, sometimes elliptic, mostly 3-8 (-10) cm. long, the margins commonly sparsely but conspicuously irregularly gland- ular-dentate (rarely entire), the indument of lower surfaces usually whitish.
V. discolor.
Leaf blades mostly lance-ovate to lance-oblong, sometimes oblong-oval, mostly 7-25 cm. long, the margins commonly entire (rarely remotely and inconspicuously glandular-dentate), the indument of lower surfaces pale yellowish or brownish.
Lower leaf surfaces long-hirsute with a mixture of paired and stellate hairs, the stellae 1-2 mm. in diameter, the paired hairs at least 1 mm. long.
V. mortonianum.
Lower leaf surfaces short-stellate-tomentose, the stellae mostly about 0.5 mm. in diameter, a few rarely to 1 mm.
Calyx tube densely stellate-tomentose V. disjunction
Calyx tube appearing green and more or less visicid, with relatively few hairs,
these both simple and stellate V. disjunctum var. mendax.
Lower leaf surfaces essentially glabrous or the indument confined to costae, veins, and/or vein-axils, the hairs appearing simple but usually a mixture of simple, paired, and some fasciculate hairs.
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Leaf margins entire or minutely serrulate, the lower surfaces more or less glandular-punctate, the punctae dark red to black; inflorescences sessile, peduncle none; fruits conspicuously flattened, to 1.5 cm. long, the stone not
grooved V. elatum.
Leaf margins entire or remotely glandular-dentate but never minutely serrulate, the lower surfaces not glandular-punctate; inflorescences usually conspicuously pedunculate (rarely subsessile); fruits little if any flattened, 0.7-1 cm. long, the stone grooved. Leaf blades commonly 2.5-5 (-6) cm. long.
Peduncles and rays of cymes conspicuously long-hiruste with spreading, white hairs about 1 mm. long, or if glabrous or glabrate, with at least some long, spreading, white hairs about the nodes; pedicels and calyx
tubes eglandular V. lautum.
Peduncles and rays of cymes more or less pubescent or puberulent with short hairs (rarely glabrate); pedicels and calyx tubes more or less red- glandular. Leaf apex obtuse; peduncles and rays densely puberulent with a mixture
of minute, simple and falsely stellate hairs V. obtusatum.
Leaf apex apiculate, acute, or sharply acuminate; peduncles and rays pubescent with appressed or subappressed hairs or essentially glabrous. Leaf blades mostly obovate, almost rounded and apiculate at the apex
or abruptly short-acuminate V. tacanense.
Leaf blades mostly lanceolate to elliptic, acuminate to long-acuminate
at the apex V. siltepecanum.
Leaf blades commonly 5-15 cm. long.
Leaf blades mostly 5-9 cm. long, acute to sharply long-acuminate at apex, the margins entire or with a few remote, glandular teeth on each side; caducous bracts filiform or linear to narrowly elliptic; cymes commonly 3-6 cm. in diameter.
Rays of cymes glabrous; calyx lobes not or scarcely ciliate V. blandum.
Rays of cymes pubescent with subappressed hairs; calyx lobes ciliate.
V. amatenangense.
Leaf blades mostly 6-15 cm. long, obtusely acuminate at the apex, often mucronulate, the margins usually completely entire, only rarely with 1-2 glandular teeth mostly near the base; caducous bracts leaflike; cymes commonly 6-14 cm. in diameter V. hartwegii.
Viburnum amatenangense Lundell, Wrightia 3: 170. 1966.
Not reported from Guatemala; known only from the type collection from Chiapas, Mexico, Amatenango del Valle, 1,835 m., Matuda 15854.
Small tree to 3 m. tall, the branches hirsute with both stellate and simple hairs; leaves on short, hirsute petioles 3-7 mm. long, the blades ovate, oval-ovate, elliptic, or lanceolate, mostly 5-7 cm. long, 2.2-4.3 cm. wide, more or less bullate, acute to short- acuminate, rounded to subcordate at the base, the margins entire or with 1-2 glandular teeth near the base, glabrous above or nearly so, more or less pubescent on costae and veins beneath; peduncles 1.5-3 cm. long, pubescent with subappressed hairs; cymes 3-5 cm. broad, the 5-7 rays pubescent with subappressed hairs, sparsely
NASH: FLORA OF GUATEMALA 287
red-glandular; bracts linear, ciliate; calyx tube to 1.5 mm. long, more or less pubescent and somewhat red-glandular especially near the base, the lobes triangular, ciliate; corolla white, glabrous, about 3 mm. long; stamens exserted; style glabrous; fruits not seen.
Although the type is cited by Lundell as Matuda 5854, the isotype in the herbarium of Field Museum, mounted with Matuda's original label, bears his number 15854.
Viburnum blandum Morton, Contr. U. S. Nat. Herb. 26: 352. 1933. V. optatum Morton, torn. cit. 358 (type from Huehuetenango, C. & E. Seler 2639). V. optatum var. vagum Morton (type from El Quiche, Heyde & Lux 3042). V. blandum var. vulcanicum Morton, Proc. Biol. Soc. Wash. 49: 154. 1936 (type from Volcan de Atitlan, Suchitepequez, Skutch 2125). V. montanum Lundell, Lloydia 2: 106. 1939 (type from Chiapas, Mexico).
Damp or wet, mixed, pine or Abies forest, 1,300-3,000 m.; Baja Verapaz; Chiquimula; Huehuetenango; Quezaltenango; El Quiche; San Marcos; Solola; Suchitepequez; Zacapa. Mexico (Chiapas, the type from Pinabete); El Salvador.
Shrubs or small trees, commonly 5-9 m. tall, the branches slender, often reddish, glabrous or nearly so; leaves short-petiolate, the blades ovate, oblong-ovate, lanceolate, or lance-oblong, mostly 5-9 cm. long, 2-4.5 cm. wide, sharply acuminate to long-acuminate, cuneate or rounded at the base, the margins entire or with a few remote, glandular teeth, ciliate or not, glabrous above or nearly so, hirsute on costae and veins beneath or glabrate, often barbate beneath along the costae or in lower vein axils, the lateral veins 4-6 pairs; inflorescences pedunculate, the peduncles glabrous, green or reddish, 1.5-5 cm. long; bracts filiform or linear to narrowly elliptical, usually caducous; compound cymes 3-6 (-8) cm. in diameter, the 5-7 primary rays glabrous, often rose or reddish; calyx glabrous, the lobes triangular, acute or obtuse, not or scarcely ciliate; corolla white, glabrous, the limb 5-7 mm. in diameter; stamens slightly exserted; styles usually glabrous, sometimes more or less pubescent; fruits purple-black, subglobose to broadly ovoid.
Specimens previously determined by Morton and by others to be V. blandum and V. optatum appear to exhibit no important differences. The styles of most specimens are glabrous, but a few are more or less pubescent. Morton's separation of these two species was apparently based mainly on the "densely white-villous" styles of V. optatum. Although this is true of the type specimen, two collections determined by him to be V. blandum (Standley 67366 and Steyermark 36853) have styles with some indument (those of the former are pubescent near the base while those of the latter are sparsely pubescent on the upper portions); two collections (Steyer- mark 30535 and Carlson 2384) determined by Morton to be V. optatum have glabrous styles.
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The variety vulcanicum is said to have dentate and somewhat broader leaves than the typical form with entire leaves, but there is great variation in this character, both kinds of leaves often occurring on the same plant. The lower surfaces of leaves of the typical form of the species usually have hirsute veins; in the variety vagum, the veins are glabrous.
Viburnum discolor Benth. PI. Hartweg. 83. 1842. Oreinotus discolor Oerst. Kjoeb. Vidensk. Meddel. 1860: 287. 1861. V. discolor f. integrum Morton, Proc. Biol. Soc. Wash. 49: 153. 1936 (type from Quezaltenango, Skutch 819). V. discolor var. subcordatum Morton, I.e. (type from Chimaltenango, Skutch 755). Membrillo de montana (Jalapa).
Damp or wet, mixed forest, or sometimes in oak, pine-cypress, or Cupressus forest, 1,800-3,500 m.; Chimaltenango; Guatemala; Huehuetenango; Jalapa; Quezaltenango; El Quiche; Sacatepequez; San Marcos; Solola; Totonicapan (type from mountains above Totonicapan, Hartweg 579).
Large shrubs or small trees, sometimes as much as 12 m. tall; branchlets stellate- pubescent; leaves on stellate-pubescent petioles 0.5-2 cm. long, the blades ovate to obovate or sometimes elliptic, mostly 3-8 (-10) cm. long, acute, short-acuminate, or acuminate, rounded to obscurely subcordate or cuneate at the base, the margins commonly sparsely and irregularly glandular-dentate but may be entire, often revolute, deep green above and sparsely or densely stellate-pubescent, commonly white stellate-tomentose beneath, the lateral veins 4-5 pairs; peduncles mostly 1.5-6 cm. long (rarely subsessile); cymes mostly 4-8 cm. in diameter, the 5-7 primary rays stellate-pubescent to stellate-tomentose; bracts caducous, linear; calyx tube about 1.5 mm. long, white stellate-tomentose, the lobes ovate to triangular, ciliate; corolla white, more or less sparsely pubescent outside with simple and/or stellate hairs, the limb about 5 mm. in diameter; stamens a little exserted; style glabrous; fruits purple- black, sparsely and minutely stellate-puberulent or glabrate, subglobose to broadly ovoid, about 8 mm. long.
Viburnum disjunct urn Morton, Contr. U. S. Nat. Herb. 26: 355. 1933.
Damp or wet, mixed or pine forest, sometimes in second growth, 1,100-2,600 m.; Alta Verapaz (type from Coban, Tuerckheim 977); Huehuetenango; El Progreso; El Quiche. Mexico (Chiapas); Honduras.
Shrubs or small trees, sometimes 10 m. tall, the young branches and stems densely tomentose with brown or yellowish, stellate hairs; leaves short-petiolate, the blades mostly lance-ovate, lanceolate, or lance-oblong, 7-20 (-25) cm. long, 4-10 cm. wide, acuminate to long-acuminate, usually rounded at the base, sometimes subcordate, rarely broadly cuneate, the margins entire or remotely and inconspic- uously glandular-dentate, minutely and rather sparsely stellate-puberulent above,
NASH: FLORA OF GUATEMALA 289
usually densely stellate-tomentose below with brown or yellowish hairs, the lateral veins 5-7 pairs; peduncles 2-15 cm. long, densely stellate-pubescent; bracts at base of inflorescence linear, caducous; cymes mostly 7-15 cm. in diameter, the primary rays 6-7, densely stellate-pubescent; calyx tube about 1 mm. long, densely tomentose, the lobes broadly triangular, obtuse, about 0.5 mm. long, stellate-pubescent, ciliate; corolla white, more or less pubescent outside with usually simple hairs (sometimes stellate hairs also present), the limb 5-6 mm. in diameter; stamens slightly exserted; style glabrous; fruits purple-black, broadly ovoid or subglobose, sparsely stellate- puberulent, about 8 mm. long.
Viburnum disjunction var. mendax (Morton) D. Gibson, Phytologia 25: 114. 1973. V. mendax Morton, Proc. Biol. Soc. Wash. 49: 154. 1936.
Brushy slopes or damp forest, sometimes in second growth, 1,500-2,500 m.; Huehuetenango; El Quiche (type from Nebaj, Skutch 1065).
Differs in indument, the leaves usually rather sparsely stellate- pubescent beneath (sometimes glabrate), and the calyx tube appearing green and somewhat viscid with relatively fewer hairs, these both simple and stellate.
Viburnum elatum Benth. PL Hartweg. 59. 1840. V. densum Benth. I.e.
A widespread species in Mexico. Not reported from Guatemala but may be expected, as it has been collected in Chiapas at elevations from 1,200-2,300 m.
Large shrubs or small trees to about 7 m. tall, the branches and stems glabrous, more or less dark-punctate; leaves short-petiolate, the petioles dark-punctate, winged to the base, the blades ovate, ovate-oblong, or lance-oblong, obtusely or acutely acuminate, cuneate to rounded at the base, mostly 2-7 cm. long, 1-3 cm. wide, entire or minutely serrulate, glabrous, conspicuously punctate beneath, the punctae dark red to black; inflorescences sessile, the 3-5 primary rays glabrous and dark-punctate, the compound cymes 3-6 cm. in diameter; calyx glabrous, the lobes triangular, obtuse; corolla white, the limb about 5 mm. in diameter; stamens slightly exserted; style glabrous; fruits purple-black, to 1.5 cm. long, conspicuously flattened, the stone not grooved.
Viburnum euryphyllum Standl. & Steyerm. Field Mus. Bot. 23: 90. 1944.
Known only from the type, Huehuetenango, trail between Tunima and Quisil, Sierra de los Cuchumatanes, 2,500-3,100 m., Steyermark 48425.
A tree of about 9 m., the branchlets stout, densely covered with brownish tomentum; leaves rather thick, on petioles about 1.5 cm. long, the blades rounded-
290 FIELDIANA: BOTANY, VOLUME 24
ovate or suborbicular, 9-11 cm. long, 7-10 cm. wide, acute, broadly rounded or almost truncate at the base, the margins remotely denticulate with usually 10-15 teeth on each side, grayish green above when dry, densely pubescent with minute, stellate hairs about 0.3 mm. in diameter, appearing whitish beneath with very dense, soft, stellate tomentum, the hairs about 0.3 mm. in diameter, averaging 10 or more rays each; peduncles stout, 1-3.5 cm. long; cymes about 10 cm. broad with 6-7 rays, these densely stellate-tomentose, the flowers short-pedicellate or sessile; calyx tube about 2 mm. long, densely white-tomentose, the lobes scarcely 1 mm. long, broadly ovate, obtuse, stellate-puberulent; corolla about 2.5 mm. long, densely or sparsely pubescent with mostly simple hairs, the few stellate ones usually near the base; stamens exserted; style glabrous.
This has been reported from Guatemala as V. ferrugineum (Oerst.) Donn.-Sm., a South American species.
Viburnum hartwegii Benth. PI. Hartweg. 84. 1842. Oreinotus hartwegii Oerst. Kjoeb. Vidensk. Meddel. 1860: 289. 1861. V. guatemalense Gandoger, Bull. Soc. Bot. France 65: 33. 1918 (type from Coban, Alta Verapaz, Tuerckheim s.n.). Malacate (San Marcos); mosiche (Chimaltenango); palito de jalisia (Quezal- tenango); salamo (Guatemala); tzumocte (Alta Verapaz); yech-ba- tza (Huehuetenango).
Damp or wet thickets or forest, sometimes in pine-oak forest, occasionally in second growth, 800-2,800 m.; Alta Verapaz; Chimaltenango; Chiquimula; Escuintla; Guatemala; Huehue- tenango; Jalapa; El Progreso; Quezaltenango; El Quiche; Sacatepe- quez (type said to have been collected in "montibus Santa Maria prope urbem Guatemala", Hartweg 580, probably the Volcan de Agua near Santa Maria de Jesus); San Marcos; Santa Rosa; Solola. Mexico; El Salvador; Honduras; Nicaragua.
Large shrubs or small trees, sometimes to 10 m. tall, the young branches strigose or densely pubescent with a mixture of simple hairs and antrorse, fasciculate, paired or few-rayed, yellowish hairs; leaves on petioles 0.5-2 cm. long, the blades elliptic to oblong-lanceolate, mostly 6-15 cm. long, 2.5-6 cm. wide, obtusely acuminate at the apex and often mucronulate, cuneate to narrowly rounded and often oblique at the base, the margins entire or rarely with 1-2 glandular teeth usually near the base, essentially glabrous or glabrate and lustrous above, glabrous beneath or the indument (of simple and fasciculate hairs) confined to costae and veins, or the vein-axils often densely white-barbate; lateral veins 4-5 pairs; peduncles 2-12 cm. long, commonly pubescent with subappressed hairs, compound cymes commonly 6-14 cm. broad, primary rays 5-7, more or less pubescent; bracts at base of primary rays narrow, foliaceous, usually soon caducous; terminal flowers sessile, the lateral ones pedicellate; calyx tube about 1.5 mm. long, glabrous or somewhat pubescent or sparsely glandular, the hairs when present, minute, solitary or fasciculate, the lobes rounded or obtuse, glabrous or sparely pubescent, ciliate; corolla white, about 3 mm. long, the limb 5-7 mm. in diameter, the lobes glabrous or somewhat pubescent outside; style glabrous; fruits purplish black, ovoid or broadly ellipsoid, about 8 mm. long.
FIG. 4. Viburnum hartwegii. A, branch, xVfe; B, determinate cymule, X 10; C, fruits, X 5; D, seed, X 5; E, flower dissected, X 12Vfe.
291
292 FIELDIANA: BOTANY, VOLUME 24
The presumed differences in the amount of indument on calyx and corolla on V. guatemalense and V. hartwegii do not hold true. Apparently, the pubescent calyx and corolla described for V. hartwegii represent the less common condition encountered.
Viburnum jucundum Morton, Contr. U. S. Nat. Herb. 26: 355. 1933. V. matudae Morton, Proc. Biol. Soc. Wash. 51: 215. 1938. V. chiapense Lundell, Bull. Torrey Bot. Club 66: 603. 1939. V. brunnescens Standl. & Steyerm. Field Mus. Bot. 23: 89. 1944 (type from Huehuetenango, Steyermark 49855).
Damp or wet thickets or forest, sometimes with Pinus and Abies, 2,200-3,900 m.; Huehuetenango; Quezaltenango; San Marcos. Mexico (type from Chiapas, Ghiesbreght 729).
Shrubs or commonly small trees to 9 m. or more, the branchlets stout, usually densely stellate-tomentose; leaves short- petiolate, the blades broadly rhombic to very broadly ovate, 6-19 cm. long, 5-13 cm. wide, short-acuminate, commonly rounded to shallowly cordate at the base but sometimes broadly cuneate, the margins conspicuously and rather coarsely dentate with mostly 10-26 teeth on each side, fairly densely pubescent or pilose above with fascicled and /or stellate hairs, densely stellate-tomentose to stellate-pilose below, the lateral veins 5-8 pairs; peduncles densely stellate-pubescent or stellate-tomentose, mostly 2-7 cm. long; compound cymes umbelliform, 6-16 cm. broad, the primary rays 5-7, usually densely stellate- tomentose; bracts caducous; calyx tube 1.5-2 mm. long, densely stellate-pubescent to stellate-tomentose, the 5 lobes ovate or triangular, 0.6-1 mm. long, usually obtuse, densely stellate-tomentose to nearly glabrous, usually somewhat ciliate near the apex; corolla white, the limb 4-6 mm. broad, essentially glabrous or more or less puberulent outside, the hairs simple and/or stellate; stamens exserted; style glabrous; fruits bluish black or purple, broadly ovoid to subglobose, to about 8 mm. long.
The alleged differences in the indument of leaf surfaces and/or of corollas ascribed to the various "species" in this group do not hold true although there is wide variation in degree of indument. The type collection of V. chiapense Lundell (Matuda 2925), determined by Morton in 1941 as V. matudae, exhibits the same type and degree of stellate pubescence on the lower leaf surface as that of V. jucundum Morton. Both simple and stellate hairs may be found on the outer surface of the corollas of V. jucundum; sometimes the stellate hairs are confined to the lower portion of the corolla with the simple hairs above, while in other specimens the stellate hairs are common over most of the external corolla surface.
Indument of all specimens in this group consists of three kinds of hairs: simple, fasciculate (usually 2-6 antrorse hairs) and true stellate hairs, some sessile, some stipitate. Simple and antrorse hairs in pairs or in groups of 3-6 are nearly always present along the
NASH: FLORA OF GUATEMALA 293
lower portion of the costae, in the vein axils, and sometimes along the veins. They may also be present on the leaf tissue between the veins, but are usually sparse in these areas. True stellae may be absent or nearly so in glabrate forms, or scattered over the leaf surface, or in various degrees of density may obscure the simple and antrorse hairs. Sometimes the simple hairs, which are usually a little longer, remain conspicuous along the costae. In my opinion, to describe the many variations possible in such a situation would be to describe specimens rather than species. However, because color and size of hairs on leaf surfaces and the number of rays in the stellae have been considered diagnostic, sample "patches" of indument were removed from numerous specimens, anchored in mounting medium on slides, and the hairs measured. The indument on leaf tissue of the type of V. brunnescens appears brownish or yellowish, is fairly dense, composed of sometimes overlapping but never layered hairs, the stellae commonly about 0.5 mm. in diameter, averaging 10 or more rays each; rarely an isolated, fasciculate hair was found with slightly longer rays. The lower leaf surfaces of V. jucundum appear yellowish, grey-green, or greenish as the hairs become more scattered, allowing more of the leaf tissue to show through; these hairs range from 0.5 to 0.9 mm. in diameter, each commonly with 10 or more rays but a few with only 6-7 rays.
All of these plants in many ways resemble V. rhombifolium (Oerst.) Hemsl. (from Veracruz, Mexico) and V. tiliaefolium (Oerst.) Hemsl. (Veracruz and Orizaba, Mexico), but the calyx tubes of both of these species are more or less red-glandular and both have pubescent styles. All of ours have pubescent to tomentose calyx tubes and glabrous styles. A third species, V. stellato-tomentosum (Oerst.) Hemsl. (Costa Rica and Panama), is also similar and has glabrous styles, but the calyx tubes are red-glandular and glabrous or only sparsely pubescent. The indument of the style is probably not a dependable character; see the discussion under V. blandum Morton.
Viburnum jucundum var. detractum (Standl. & Steyerm.) D. Gibson, Phytologia 25: 114. 1973. V. detractum Standl. & Steyerm. Field Mus. Bot. 23: 89. 1944. Palo canahuatch (Quezaltenango).
Damp thickets and forest, 2,000-3,800 m.; Chimaltenango; Quezaltenango (type from upper slopes of Volcan Zunil, Steyermark 34590); San Marcos; Solola.
Differs from the typical variety only in indument or lack of it.
294 FIELDIANA: BOTANY, VOLUME 24
The young branches are sparsely and laxly pilose instead of tomentose, with a mixture of stellate, fascicled, and simple hairs, and the leaf blades are essentially glabrous on the tissue between the veins. The upper leaf surfaces bear a few scattered, stellate hairs and /or fascicled hairs, except on costae and veins where the hairs may be dense; the lower leaf surfaces are nearly glabrous except on costae, veins, and in the lower vein axils, where they appear more or less barbate, the hairs mostly simple or paired. A few stellae sometimes occur along the costae and occasionally may be found scattered along the leaf margins. When present they range from 0.4- 0.7 mm. in diameter, with 8-14 rays each.
Viburnum lautum Morton, Contr. U. S. Nat. Herb. 26: 351. 1933.
Damp forest, 2,900-4,000 m.; Huehuetenango; San Marcos. Southern Mexico (the type from Chiapas).
Shrubs to 3 m. or more, the young branches and stems more or less hirsute with spreading, white hairs, these both simple and fasciculate; leaves short-petiolate, the blades broadly ovate to oblong-elliptic, mostly 2-5 cm. long, 1.5-3.5 cm. wide, acute or short-acuminate, rounded or cuneate at the base, the margins entire or sometimes with a few remote, glandular teeth, glabrous except hirsute beneath along the costae and sometimes on the veins, the hairs mostly simple, often glabrate in age, the margins more or less ciliate (not always easily seen due to the often revolute margins), the lateral veins 4-6 pairs; inflorescences on peduncles to 5 cm. long or infrequently subsessile, the peduncles usually hirsute with spreading white hairs about 1 mm. long, sometimes glabrate but then with a few long hairs at the base of the rays; bracts linear-oblanceolate to li near-elliptic, about 1 cm. long, usually ciliate, usually caducous; compound cymes 3-6 cm. in diameter, the primary rays 5-7, hirsute with spreading white hairs about 1 mm. long, or rarely glabrate; calyx tube 1-2 mm. long, usually glabrous and eglandular, the lobes broadly triangular, obtuse, ciliate or not; corolla white, glabrous, the limb about 5 mm. in diameter; stamens slightly exserted; style glabrous; fruits purple-black, obovoid to subglobose, less than 1 cm. long.
Both V. lautum and V. tacanense Lundell appear to have much in common with V. acutifolium Benth. However, according to Morton, the peduncles and rays of V. acutifolium Benth. are sparsely stellate-pubescent, with rays and calyx tubes sparingly red- glandular.
Viburnum mortonianum Standl. & Steyerm. Field Mus. Bot. 22: 294. 1940.
Known only from the type, Chiquimula, Cerro Tixixi, 3-5 miles north of Jocotan, on slope of ravine, 1,500 m., Steyermark 31606.
NASH: FLORA OF GUATEMALA 295
A shrub or small tree, the branchlets stout, densely long-hirsute with a mixture of paired, fasciculate, and stellate, fulvous hairs; leaves on petioles 12-16 mm. long, the blades oblong or lance-oblong to ovate-oblong, 9-13 cm. long, 4.5-7.5 cm. wide, acute to long-acuminate, obtuse to broadly rounded or subtruncate at the base, subentire or remotely and minutely glandular-denticulate, hirsute above with paired or few-rayed hairs 1 mm. long or more, densely stellate-hirsute beneath with stiff, sessile hairs 1-2 mm. in diameter, the lateral veins 5-6 pairs; peduncles 3-4 cm. long; compound cymes 4.5-8.5 cm. broad, densely hirsute, the primary rays 5-7; calyx tube densely hirsute, the lobes linear-lanceolate, hirtellous, long-ciliate; corolla white, sparsely hirsute outside, the limb 5-6 mm. in diameter; stamens exserted; style glabrous; fruits unknown.
Viburnum obtusatum D. Gibson, Phytologia 25: 114. 1973.
Not reported from Guatemala but may be expected: type from Chiapas, Mexico, 2,530-2,840 m., Breedlove & Raven 8118 (flowering) and Breelove 9077 (fruiting).
Shrubs or small trees to about 6 m. tall; leaves short-petiolate, the blades ovate to lanceolate, mostly 2-5 (-6) cm. long, 1.5-2.5 cm. wide, obtusely short-acuminate at the apex, rounded or acute at the base and usually abruptly decurrent on the petiole for one-fourth to one-half its length, essentially glabrous and shining on both surfaces, or with some indument of short, simple and fasciculate hairs, usually confined to the costae, the margins entire, ciliate or not; inflorescences on peduncles 1-3 cm. long, these finely puberulent with a mixture of simple, paired, and fascicled (falsely stellate) hairs; bracts caducous, not seen; cymes twice compound, 1.5-3 cm. broad, the primary rays 5-7, minutely puberulent, some of the rays more of less red- glandular; pedicels frequently red- glandular; calyx tube at anthesis 1-1.5 mm. long, somewhat red-glandular or not, the lobes broadly triangular, obtuse or acute, glabrous, about 1 mm. long, sparsely short-ciliate or not; corolla white, glabrous, the limb about 5 mm. in diameter; stamens slightly exserted; style glabrous or a little pubescent near the base; fruits purple-black, obovoid, less than 1 cm. long, the stone grooved.
Viburnum siltepecanum Lundell, Wrightia 3: 171. 1966.
Not reported from Guatemala but may be expected there. Known only from the type, Matuda 5111 from Chiapas, Mexico, 1,600 m.
Small tree, 4-5 m. tall, the branches angled, the younger branches minutely hirsute with fascicled (falsely stellate) hairs; leaves on glabrous petioles 5-7 mm. long, the blades lanceolate to elliptic, mostly 3-5 cm. long, 1.3-4 cm. wide, acuminate to long-acuminate, acute or almost rounded at base, the margins entire or obscurely and remotely glandular-dentate, glabrous above with the exception of some indument on the lower part of the costae, glabrous below except some indument usually present on the costae and/or in the vein axils; peduncles 1-3.5 cm. long, minutely hirsute with both simple and falsely stellate hairs or glabrate, sparsely red-glandular; cymes mostly 2-5 cm. broad, the 5-7 primary rays glabrous or sparsely and minutely hirsute, sparsely glandular; bracts caducous; calyx tube glabrous, the lobes triangular, glabrous, mostly eciliate; corolla white, glabrous; stamens exserted; style minutely pubescent at base.
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According to Morton's description of V. acutifolium Benth., and his photograph of the type specimen (Hartweg 449 from Oaxaca), V. siltepecanum appears to be very similar and may prove to be synonymous with it.
Viburnum tacanense Lundell, Lloydia 2: 108. 1939.
Damp forest, 3,000-4,000 m.; Quezaltenango; San Marcos. Mexico (the type from Chiapas).
Shrubs or small trees to about 7 m. tall, the young branches and stems pubescent with appressed or subappressed, simple and fasciculate hairs; leaves short-petiolate, the blades mostly obovate, sometimes broadly ovate or nearly elliptic, mostly 2-5 cm. long, 1-2.5 cm. wide, abruptly short-acuminate or almost rounded and apiculate, cuneate to rounded at the base, the margins entire or with a few glandular teeth, ciliate or not, glabrous above except for appressed hairs along the costae, glabrous below except along the costae and sometimes on the veins, sometimes barbate in the vein axils, the lateral veins 4-5 pairs; inflorescences subsessile or on short peduncles to 1.5 cm. long, these pubescent with appressed or subappressed hairs; bracts caducous; cymes 2-4 cm. in diameter, the 5 primary rays pubescent with short, appressed or subappressed hairs, the pedicels frequently somewhat red-glandular; calyx tube 1.5-2.4 mm. long, usually sparsely red-glandular, the lobes broadly triangular, obtuse, sparingly short-ciliate or not; corolla white, glabrous, the limb about 5 mm. in diameter; stamens slightly exserted; style glabrous; fruits purple- black, obovoid to subglobose, less than 1 cm. long.
The subsessile to very short-pedunculate inflorescences de- scribed as characteristic of V. tacanense sometimes also occur on specimens matching in most other respects the description of V. lautum Morton. The latter species, however, has peduncles and rays usually conspicuously long-hirsute with spreading, white hairs, or at least with some long hairs about the nodes.
VALERIANACEAE. Valerian Family By DOROTHY L. NASH
Annual or perennial herbs, rarely somewhat woody, erect or scandent, glabrous or sometimes pubescent, the roots with a distinctive and persistent odor; leaves often chiefly basal, the cauline leaves commonly small, sometimes none, opposite or several crowded on short, lateral, opposite branches, the blades frequently pinnatifid or pinnatisect, the margins entire or dentate; flowers small, sessile or short-pedicellate, commonly disposed in dichotomous or aggregate dichasia, the flowers perfect or sometimes pistillate ones also present or rarely the plants dioecious; calyx tube adnate to the ovary, the limb in anthesis usually small, after anthesis unchanged or variously modified, often with filiform segments or appendages; corolla gamo- petalous, mostly white or pink, funnelform or campanulate to subsalverform, the tube terete or gibbous or even calcarate, usually ampliate above, the limb spreading, mostly 5-lobate, regular, oblique, or bilabiate, the lobes imbricate in bud; stamens 1-
NASH: FLORA OF GUATEMALA 297
4, the filaments free, inserted on the corolla tube, alternate with the lobes, often exserted; anthers versatile or attached near the base, the 2 thecae parallel, distinct, dehiscent by longitudinal slits, sometimes the thecae sulcate and the anthers then 4- lobate; ovary inferior, with 1 perfect locule, 2 empty locules usually present and sometimes larger than the fertile one; style filiform, undivided and with a subtruncate stigma or more commonly 2-3-lobate at the apex; ovule 1 in the fertile locule, anatropous, pendulous from the apex of the locule; fruit achenelike, dry, indehiscent, crowned by the small or enlarged, often pappuslike or plumose calyx limb; seed pendulous, the testa membranaceous; endosperm none or scant; embryo straight, the cotyledons oblong, longer than the superior radicle.
About 11 genera, with more than 300 species chiefly in temperate and tropical regions; in the American tropics found mostly in the higher mountains, except for a few weedy species. Only one genus occurs in Guatemala.
VALERIANA Linnaeus
Reference: Frederick G. Meyer, Valeriana in North America and the West Indies, Ann. Mo. Bot. Gard. 38: 377-503. 1951.
Perennial or annual herbs from tap-roots or rhizomes, erect or scandent, glabrous or pubescent or villous; leaves often predominantly basal, usually petiolate, the margins entire or dentate, or the cauline and basal leaves often pinnatifid or pinnatisect, the cauline ones opposite; inflorescence a compound dichasium (dichotomous throughout, the ultimate dichotomies more or less flat-topped in anthesis) or an aggregate dichasium (more or less pyramidal in anthesis), both types becoming more diffuse in age; unisexual as well as perfect flowers sometimes present, or the plants dioecious; bracts usually present; calyx in flower inconspicuous, the limb segments commonly inrolled, later developing into 5-20 or more dentate, setose, or plumose segments; corolla white or pink, funnelform or campanulate to subsalverform, the tube short or rarely elongated, often gibbous at the base, the 5 lobes spreading; stamens 3, included or exserted, the anthers bithecous but the thecae sometimes sulcate and the anthers then 4-lobate; style included or exserted, the stigma 2-3-lobate; fruit compressed, the posterior face with one rib, the anterior face usually with 3 ribs, these conspicuous or not, one or both faces glabrous or more or less hirtellous, rarely scabrous.
A genus of perhaps 200 species, in all continents except Australia, with 12 species in Guatemala.
Leaf blades undivided, oblong-linear to spathulate, attenuate to the base; anthers
appearing 4-lobate, the thecae sulcate V. prionophylla.
Leaf blades divided or not, but when undivided, never oblong-linear to spathulate
and not attenuate to the base; anthers bilobate, the thecae not sulcate. Plants scandent.
Leaves all or mostly trifoliolate V. scandens.
Leaves simple.
Leaf blades cordate at the base; inflorescence appearing more or less pyramidal or paniculate V. scandens var. candolleana.
298 FIELDIANA: BOTANY, VOLUME 24
Leaf blades obtuse, rounded, or truncate at the base; inflorescence appearing
corymbose, flat-topped or nearly so V. clematitis.
Plants erect. Leaves mostly simple (rarely some leaves trifoliolate in V. palmatiloba), the
margins entire or dentate.
Leaf blades palmately 5-7-lobate, simple or sometimes trifoliolate, the 2 lower leaflets when present much smaller and narrower than the principal blade .
V. palmatiloba. Leaf blades simple, not lobate, the margins somewhat angulate, dentate, or
undulate-dentate.
Leaves predominantly basal, long-petiolate, the petioles sometimes to 30 cm. long, the blades mostly 6-18 cm. long, cordate at the base, the margins somewhat angulate or undulate-dentate, the smaller, cauline leaves 2-4
pairs V. cucurbitifolia.
Leaves cauline, 3-8 pairs, sessile or short-petiolate, the petioles less than 3 cm. long, the blades mostly 1-4 (-5) cm. long, rounded or subacute at base, the margins almost regularly dentate or sinuate-dentate.
V. urticaefolia. Leaves mostly pinnatifid or pinnatisect.
Inflorescence a compound dichasium, dichotomous throughout, appearing
corymbiform in anthesis (later more diffuse). Cauline leaves usually 1 pair; calyx limb segments 11-12, or obsolete.
V. pulchella. Cauline leaves usually 2-3 pairs; calyx limb segments 8, or obsolete.
V. deltoidea. Inflorescence an aggregate dichasium, appearing more or less pyramidal in
anthesis (later becoming laxly peniculiform). Stamens exserted.
Cauline leaves 3-8 pairs, mostly 1-6 cm. long including the petiole, the blades commonly bipinnatisect (the lobes incised-pinnate) or about 5- foliolate (the lobes then irregularly dentate or crenate); terminal branches of the inflorescence scorpioid; calyx limb segments 6 or 7.
V. robertianifolia.
Cauline leaves 2-4 pairs, mostly 4-13 cm. long including the petiole, the
blades commonly pinnate to pinnatifid (or some leaves rarely
undivided); terminal branches of the inflorescence not scorpioid; calyx
limb segments 10-12.
Plants with perfect flowers or sometimes with perfect and pistillate
flowers V. densiflora.
Plants dioecious V. densiflora var. affinis.
Stamens included.
Lateral segments of leaves more or less decurrent on the winged rachis;
achenes more or less winged on the adaxial margins V. palmeri.
Lateral segments of leaves distinct; achenes not winged, the adaxial margins more or less revolute.
Achenes pilosulous or glabrous V. sorbifolia.
Achenes more or less scabrous V. sorbifolia var. mexicana.
Valeriana clematitis HBK. Nov. Gen. & Sp. 3: 327. 1819; Meyer, Ann. Mo. Bot. Gard. 38: 438, f. 25. 1951. V. laurifolia HBK.
NASH: FLORA OF GUATEMALA
299
FIG. 5. Valeriana clematitis. A, habit, x Vr, B, staminate flower dissected, x 12; C, segments of inflorescences and a pistillate flower, all x 12.
torn. cit. 328. V. subincisa Benth. PL Hartweg. 39. 1839. V. pauonii Poepp. & Endl. Nov. Gen. & Sp. 3: 16, t. 215. 1845. V. hispida Turcz. Bull. Soc. Nat. Mosc. 25(2): 172. 1852. V. pavonii var. yungasensis Briq. Ann. Conserv. Jard. Bot. Geneve 17: 337. 1913. V. ghies- breghtii Briq. torn. cit. 345. V. laxissima Standl. & L. Wms. Ceiba 1: 252. 1951.
300 FIELDIANA: BOTANY, VOLUME 24
Open or dense, damp or wet thickets or forest, frequently in Abies-Cupressus forest, 2,500-3,800 m.; Chimaltenango; Huehue- tenango; El Progreso; Quezaltenango; El Quiche; San Marcos; Solola; Totonicapan. Mexico; Costa Rica; Panama; Colombia; Peru; Venezuela.
Perennial, herbaceous or suffrutescent vines, the stems sometimes several m. long, pubescent or glabrate; leaves opposite or several often crowded on short, lateral, opposite branches, at least the lower ones commonly long-petiolate, the blades simple, lanceolate, lance-elliptic, or ovate, mostly 2-8 cm. long, acuminate, obtuse, rounded, or truncate at the base, the margins usually entire or nearly so, rarely some leaves dentate or denticulate, usually finely pubescent above, glabrous or sparsely to densely pilosulous below; inflorescence an aggregate or compound dichasium, usually dense at first and appearing corymbose but becoming diffuse; perfect and pistillate flowers both present, fragrant; calyx limb with 10 or 12 segments; corolla white, pinkish, or white with lavendar lobes, funnelform, the perfect flowers 2-5.5 mm. long, the pistillate ones 1-3 mm. long; stamens and style exserted; achenes oblong-linear, frequently more or less falcate, glabrous or minutely puberulent.
The leaves of South American specimens and of some Costa Rican material are thicker than those of Mexican and Guatemalan specimens but I can find no other differences.
Valeriana cucurbitifolia Standl. Field Mus. Bot. 22: 58. 1940 (type from Siltepec, Chiapas, Mexico). V. cacalioides Standl. torn, cit. 125 (type from Mt. Ovando, Chiapas).
On bluffs, 1,800-3,000 m.; Huehuetenango (Cumbre Papal, between summit and La Libertad, Steyermark 50966). Mexico (Chiapas).
Erect perennials from thick roots, the robust stems sometimes more than 1 m. tall, glabrous or pilosulous, the nodes always pilosulous; leaves predominantly basal, on long petioles mostly 12-30 (-40) cm. long, the blades simple, reniform to rounded- ovate, mostly 6-18 cm. long, 5-15 cm. wide, rounded or obtuse at the apex, cordate at the base, the margins irregularly sinuate-dentate to undulate-dentate or angulate, usually glabrous above or nearly so, or the indument confined to costae and veins, usually sparsely pilose or pilosulous beneath, at least on costae and veins, cauline leaves 2-4 pairs, similar to the basal but much smaller, the uppermost pair sessile or nearly so and much reduced, more conspicuously dentate than the basal leaves; inflorescence usually large, long-pedunculate, the compound (or rarely aggregate) dichasium appearing corymbiform, the branches usually hirtellous; flowers perfect; calyx limb segments 10-14; corolla white, funnelform, about 4 mm. long, the tube gibbous at the base; stamens and style exserted; achenes oblong-ovate, often falcate, glabrous on one side, more or less pubescent on the other, the abaxial ribs usually conspicuous.
Valeriana deltoidea F. G. Meyer, Ann. Mo. Bot. Gard. 38: 459, /. 36. 1951.
NASH: FLORA OF GUATEMALA 301
Not reported from Guatemala but as it has been collected in Chiapas (Breedlove 10429 and 14789, determined by Meyer) as well as in western Mexico, it may be expected in Guatemala.
Erect perennials to about 50 cm. tall, the stems glabrous or puberulent; cauline leaves usually 2-3 pairs, the lower ones long-petiolate, those uppermost short- petiolate or sessile, all leaves quite variable in form, pinnatisect or pinnate with 2-3 pairs of distinct leaflets, palmately trifoliolate, palmately lobate (the terminal leaflet or lobe usually elliptic to obovate, rarely linear, considerably longer than the lateral segments or lobes), or entire and oblong-oval to spathulate, mostly 2-5 cm. long; inflorescence a compound dichasium, dichotomous throughout, appearing corymbiform at anthesis, later becoming diffuse; bracts narrowly oblong to linear; flowers perfect; calyx limb segments 8 or obsolete; corolla white or pink, funnelform, 4-5 mm. long, the tube gibbous at the base; stamens and style exserted; achenes more or less bellows-shaped, glabrous, often spotted, the keel usually prominent.
Valeriana densiflora Benth. PL Hartweg. 39. 1839. V. pilosiuscula Mart. & Gal. Bull. Acad. Brux. 11(1): 122. 1844. V. retrorsa Fern. Proc. Amer. Acad. 36: 502. 1901.
Not reported from Guatemala but may be expected there as it has been collected in Chiapas, Breedlove 5991 (Cerro San Cristobal, San Cristobal Las Casas), Breedlove 6731 (Piedracitas), Breedlove 11099 (Municipio Tenejapa), and Breedlove 14544 (near Municipio Venustiana Carranza).
Perennials from simple or forked taproots, the stems to 90 cm. tall, leafy or subscapose, more or less pilosulous or glabrate; basal leaves usually pinnate to pinna tif id, rarely undivided, mostly 3-15 cm. long including the petiole, the terminal lobe or leaflet linear, oblong, elliptic, or suborbicular, irregularly dentate or entire, glabrous or somewhat hirtellous on costae and veins, the lateral ones usually distinct, sometimes decurrent on the rachis, 1-6 pairs, considerably smaller than the terminal one; cauline leaves 1-4 pairs; inflorescence an aggregate dichasium, more or less pyramidal at anthesis, later diffuse; bracts 5-15 mm. long, glabrous; flowers perfect or sometimes pistillate ones also present, these smaller; calyx limb segments 10-12; corolla white or pinkish, mostly 3-6 mm. long; stamens and style exserted; achenes linear to oblong-ovate or ovate, 1.4-3 mm. long, hirtellous or glabrous, the abaxial ribs conspicuous or not.
V. densiflora var. affinis (Mart. & Gal.) F. G. Meyer, Ann. Mo. Bot. Gard. 38: 457. 1951. V. affinis Mart. & Gal. Bull. Acad. Brux. 11(1): 123. 1844.
Stony ledges, Sierra de los Cuchumatanes, 2,500 m.; Huehue- tenango. Southern Mexico.
Differs from V. densiflora principally in that the plants are dioecious. Meyer described the leaves of V. densiflora as "pre- dominantly cauline" and those of the var. affinis as "predominantly
302 FIELDIANA: BOTANY, VOLUME 24
basal" and this is true of most Mexican material. However, this character does not hold true in all cases as a number of Guatemalan and Mexican specimens annotated by Meyer as var. affinis have predominantly cauline leaves. The alleged differences in taproots and achenes were seldom discernible in herbarium material available to me.
Valeriana palmatiloba F. G. Meyer, Ann. Mo. Bot. Gard. 38: 448, /. 31. 1951.
Not reported from Guatemala but may be expected there. Known only from the type collection, an immature specimen, Purpus 419 from Chiapas, Mexico.
Erect perennial to about 30 cm. tall, the stem glabrous or more or less pilosulous with spreading hairs, the nodes pilose; basal leaves mostly long-petiolate, simple or 3- parted, the blade when simple 5-7-lobate, or when divided the terminal leaflet usually 5-lobate, acute, hastate to cordate at base, 2-7.5 cm. long, 1.5-3.5 cm. wide, the margins sometimes very sparsely dentate, more or less hirtellous along costae and veins, the 2 lateral leaflets mostly 1-1.7 cm. long; cauline leaves 2-3 pairs, similar to the basal ones, petiolate, except the uppermost pair much reduced and sessile or subsessile; inflorescence an aggregate or compound dichasium about 1.5 cm. broad in anthesis, more or less hirtellous; bracts 4-6 mm. long, glabrous or sometimes ciliate; flowers perfect; calyx limb immature; corolla funnelform, white, the tube gibbous, about twice as long as the lobes; stamens and style exserted; achenes glabrous on one side, short-hirtellous on the other.
Valeriana palmeri Gray, Proc. Amer. Acad. 22: 417. 1887. V. langlassei Briq. Ann. Conserv. Jard. Bot. Geneve 17: 341. 1914. V. fistulosa Briq. torn. cit. 343.
Damp or wet thickets and forest, often on rocky slopes in pine- oak forest, 1,100-2,100 m.; Chiquimula; Guatemala; Huehue- tenango; Jalapa; Santa Rosa. Mexico; Honduras; El Salvador; Nicaragua; Costa Rica.
Erect annuals from subnapiform taproots, the stems sometimes as much as 2 m. tall, slender or stout; cauline leaves 3-7 pairs, petiolate, mostly 3-20 cm. long including the petiole, the blades pinnate to pinnatifid (rarely undivided), the margins usually serrate-dentate, rarely almost entire, rachis more or less winged, the terminal lobes ovate to elliptic, acuminate or acute, more or less decurrent on the rachis, lateral lobes 1-5 pairs, linear-oblong and smaller than the terminal lobe, somewhat decurrent; inflorescence an aggregate dichasium, mostly 10-40 cm. long, the terminal branches scorpioid, glabrous; bracts minute, glabrous; flowers perfect or some pistillate ones also present; calyx limb segments 8-11; corolla white, funnelform to subcampanulate, 1-2 mm. long; stamens and style included; achenes mostly 1-3 mm. long, broadly elliptic, winged on the adaxial margins, glabrous or pilosulous.
Valeriana prionophylla Standl. Field Mus. Bot. 18: 1384. 1938. V. skutchii Standl. op. cit. 22: 59. 1940 (type from
NASH: FLORA OF GUATEMALA 303
Huehuetenango, Skutch 1240). V. pumilio Standl. & L. Wms. Ceiba 1: 253. 1951. Pericon de monte (Huehuetenango).
In open, damp, pine forest, in damp or wet alpine meadows, or in rather dry crevices of rocky cliffs, sometimes on limestone, 2,100- 4,200 m.; Chimaltenango; Guatemala; Huehuetenango; Quezal- tenango; Sacatepequez; San Marcos; Solola; Totonicapan. Mexico (Chiapas); Costa Rica.
Erect perennials from long, often forked, taproots, the stems 10-80 cm. tall, sparsely pilose or almost glabrous, the nodes pilosulous; leaves predominantly basal, usually numerous and appearing somewhat cespitose, the blades undivided, oblong- linear to spathulate, mostly 3-30 cm. long, 0.5-3 cm. wide, obtuse, attenuate to the subpetiolar base, the margins serrate, serrate-dentate, undulate-dentate, crenate, or rarely entire, usually ciliate, glabrous to pilosulous, cauline leaves 2-3 pairs, mostly 2- 20 cm. long, usually sessile and clasping, sometimes short-petiolate; inflorescence long-pedunculate, the numerous flowers disposed in an aggregate dichasium, dense or diffuse; bracts linear; calyx limb with 9-11 segments; corolla rotate, 1.5-3 mm. long, white, pink, or pale violet, glabrous; stamens exserted, the anthers appearing 4- lobate, the thecae sulcate; style exserted; achenes 2-3 mm. long, smooth or transversely rugulose, glabrous or pilosulous, the adaxial ribs usually conspicuous.
Valeriana pulchella Mart. & Gal. Bull. Acad. Brux. 11(1): 123. 1844. V. woodsonii Standl. Ann. Mo. Bot. Gard. 27: 346. 1940.
Damp alpine meadows, 3,300-3,500 m.; Huehuetenango; Quezal- tenango; San Marcos. Mountains of southern Mexico; Costa Rica; Panama.
Erect perennials from thick roots, the stems glabrous or nearly so, to about 30 cm. tall; basal leaves long-petiolate, glabrous or nearly so, pinnatisect with 1-5 (-7) pairs of lateral leaflets very variable in shape, the terminal leaflet larger and broader than the others, mostly 1.5-3 cm. long, subreniform to elliptic or obovate, rounded to acute at the apex, the margins sinuate-dentate to undulate; cauline leaves usually only one pair, much reduced; inflorescence a compound dichasium, dichotomous throughout, densely flowered, compact and appearing corymbiform at anthesis; bracts linear to spathulate; flowers perfect or rarely some pistillate flowers also present; calyx limb segments 11-12 or obsolete; corolla funnelform, white or pinkish, mostly 4-6 mm. long, the tube gibbous at the base; stamens and style exserted; achenes oblong to ovate, glabrous or hirtellous, the keel prominent or not.
Valeriana robertianifolia Briq. Ann. Conserv. Jard. Bot. Geneve 17: 342. 1914. V. delicata Standl. & Steyerm. Field Mus. Bot. 23: 255. 1947 (type from Huehuetenango, Steyermark 50936).
Damp banks, ravines, in thickets or forest, often in pine or oak- pine-cypress forest, 1,500-3,100 m.; Chimaltenango; Huehuetenango; Jalapa; Quezaltenango; San Marcos; Solola; Totonicapan. Mexico; Honduras; Costa Rica; Venezuela.
304 FIELDIANA: BOTANY, VOLUME 24
Erect or ascending herbs mostly 15-40 cm. tall, the stems sparsely or moderately pilose to hirtellous; leaves predominantly cauline, small, the 3-8 pairs mostly 1-6 cm. long including the petiole, the blades commonly bipinnatisect with incised-pinnate lobes, or about 5-foliolate, the lobes then irregularly dentate or crenate, more or less pilose or pilosulous, the terminal leaflet or lobe larger than lateral ones, dentate or variously incised; inflorescence an aggregate dichasium mostly 6-38 cm. long, 3-12 cm. wide, the terminal branches scorpioid; flowers sessile, both perfect and pistillate ones present; calyx limb segments 6 or 7; corolla white or pinkish, not more than 1 mm. long, campanulate-funnelform, the tube short; stamens and style exserted; achenes ovate to elliptic, smooth, glabrous on one side, densely hirtellous on the other.
Valeriana scandens L. Sp. PI. ed. 2: 47. 1762. V. volubilis Sesse & Mocino ex DC. Prodr. 4: 634. 1830, nomen nudum. V. phaseoli Braun, Ind. Sem. Hort. Berol. 13 (App. 2). 1851. V. scandens var. genuina Muell. in Mart. Fl. Bras. 6(4): 344. 1885. V. scandens var. angustiloba Muell. I.e.
Damp or wet thickets or forest, sometimes in pine forest, near sea level to 1,500 m.; Alta Verapaz; Chiquimula; Escuintla; Izabal; Peten. Florida; Mexico; British Honduras; Honduras; El Salvador; Nicaragua; Costa Rica. West Indies; South America.
Perennial, herbaceous vines, the stems glabrous or somewhat pilosulous; leaves opposite, long-petiolate, simple or trifoliolate, when simple the blades broadly ovate to triangular-ovate, acute to acuminate, cordate at the base, the margins undulate- dentate or entire, when trifoliolate, the leaflets sessile or short-petiolate, the terminal ones ovate to oval, acute or acuminate, rounded at the base, undulate or entire, the lateral ones more or less lance-ovate and oblique; inflorescence an aggregate dichasium with both perfect and pistillate flowers, pedunculate, commonly 10-40 cm. long; bracts 1.5-2.5 cm. long; calyx limb segments 9-15; corolla white, 1-2.5 mm. long, campanulate-funnelform; stamens and style exserted; achenes more or less ovoid to bellows-shaped, glabrous or pubescent.
Valeriana scandens var. candolleana (Card.) Muell., Fl. Bras. 6(4): 344. 1885. V. alpina Veil. Fl. Flum. 28. 1825. V. candolleana Gardn. in Hook. Lond. Journ. Bot 4: 112. 1845. V. mikaniae Lindl. Journ. Hort. Soc. 3: 316. 1848. V. scandens var. dentata Muell. Mart. Fl. Bras. 6(4): 344. 1885. Camchiri-sumaljop (Guatemala, fide Aguilar); genciana (Jalapa); chansi and shautzi (Alta Verapaz, Quecchi).
Damp or wet thickets or forest, 600-3,800 m.; Alta Verapaz; Chimaltenango; Guatemala; Huehuetenango; Jalapa; El Progreso; El Quiche; Quezaltenango; Retalhuleu; Sacatepequez; San Marcos; Santa Rosa; Solola; Suchitepequez; Zacapa. Mexico; Honduras; El Salvador; Nicaragua; Costa Rica; Panama; West Indies.
NASH: FLORA OF GUATEMALA 305
Leaves simple, the blades broadly ovate to triangular-ovate, mostly 3-7 cm. long, acute to acuminate, cordate at the base, the margins undulate-dentate or entire.
Valeriana sorbifolia HBK. Nov. Gen. & Sp. 3: 332. 1819.
Damp or wet forests or thickets, open or shaded banks, rocky or grassy slopes, 1,200-2,700 m.; Alta Verapaz; Chimaltenango; Escuintla; Guatemala; Huehuetenango; Jalapa; El Quiche; Sacate- pequez; San Marcos; Zacapa. Mexico to Panama; northern South America.
Erect or ascending (rarely subscandent) annuals to about 1 m. tall, the stems usually simple, sometimes branched, pilosulous or glabrous; leaves cauline, petiolate, usually 4-7 pairs, mostly 3-15 cm. long including the petiole, the blades usually pinnate, the terminal segment 2-6 cm. long, lanceolate to obovate or elliptic, obtuse to acuminate, the margins usually serrate to subcrenate, the lateral segments commonly 3-6 pairs (sometimes only 1-2 pairs, especially on younger plants), distinct, smaller than the terminal one; inflorescence an aggregate dichasium, commonly 10-50 cm. long, the terminal branches scorpioid, usually glabrous, occasionally tufted on the nodes; bracts minute, glabrous; flowers both perfect and pistillate; calyx limb segments 6-11; corolla white, funnelform to subcampanulate, 1-3 mm. long; stamens and style included; achenes about 2 mm. long, smooth, pilosulous or glabrous.
Valeriana sorbifolia var. mexicana (DC.) F. G. Meyer, Ann. Mo. Bot. Gard. 38: 475. 1951. V. mexicana DC. Prodr. 4: 640. 1830.
Damp or wet forest, on shaded banks and slopes, 1,300-2,200 m.; Chimaltenango; Huehuetenango; Jalapa; Sacatepequez. Mexico.
Differs from V. sorbifolia in its scabrous achenes.
Valeriana urticaefolia HBK. Nov. Gen. & Sp. 3: 330, t. 275. 1819. V. scorpioides DC. Prodr. 4: 635. 1830. V. erysimoides Poepp. & Endl. Nov. Gen. & Sp. 3: 16. 1844. V. rhomboidea Green, Pittonia 1: 154. 1888. V. sallei Briq. Ann. Conserv. Jard. Bot. Geneve 17: 339. 1914.
Damp, shaded banks, wet thickets, or in rather open forest, often in pine or pine-oak forest, 900-2,400 m.; Alta Verapaz; Chimaltenango; Chiquimula; Guatemala; Huehuetenango; Jalapa; Santa Rosa; Sacatepequez; Solola; Zacapa. Central and southern Mexico; Honduras; Nicaragua; Costa Rica; Panama; Andes of western South America.
Erect plants from subnapiform taproots, the stems simple or branched, to about 40 cm. tall, pubescent or pilosulous, at least near the base, with reflexed hairs; leaves predominantly cauline, the 3-8 pairs sessile or short-petiolate, simple, the blades mostly rounded-ovate, ovate, or flabelliform, sometimes triangular, mostly 1-4 cm. long, rounded or acute at the apex, rounded or subacute at the base, the margins almost regularly dentate or sinuate-dentate, hirsutulous above, usually glabrous or
306 FIELDIANA: BOTANY, VOLUME 24
nearly so beneath or the indument confined to costae and veins; inflorescence an aggregate or compound dichasium, small or large, repeatedly dichotomous or paniculate with slender branches, the terminal ones scorpioid; flowers perfect; calyx limb segments 10-13; corolla usually white, sometimes pinkish, narrowly funnelform, 2-4 (-5) mm. long, the tube gibbous at the base; stamens and style exserted; achenes ovate, glabrous or usually pubescent on one side, the midribs conspicuous.
DIPSACACEAE. Teasel Family
By DOROTHY L. NASH
The plants of this family are all natives of the Old World, but a few have become naturalized in temperate North America, and several are cultivated for ornament. The common teasel, Dipsacus sylvestris Huds., was observed growing in the Jardin Botanico, where it was not known to have been planted, and may have been introduced accidentally. It is a coarse, branched, herbaceous plant with opposite, simple leaves, and flowers in long-pedunculate, dense heads subtended by an involucre of curved, linear bracts. Its very rough flower heads were much used formerly for raising the nap on woolen cloth. One member of the family is grown commonly in Guatemalan gardens for ornament, at almost all elevations. It is Scabiosa atropurpurea L., native of southern Europe, known in Guatemala as "ambarina," "amarines," "flor de viuda," and "viuditas." In El Salvador it is called "mirame-linda." It is an erect, branched annual about 50 cm. tall, the basal leaves lyrate- pinnatifid and coarsely dentate, the cauline leaves opposite, pinnately parted. The rather showy flowers, crowded in large long- stalked heads, vary from very dark purple to rose or white. The fruit is a small achene, similar to that of some Compositae. In Guatemala the plant is a favorite garden flower at middle elevations, and the flowers are often sold in the markets.
CUCURBITACEAE. Gourd Family
By JENNIE V. A. DIETERLE UNIVERSITY OF MICHIGAN
References: Alfred Cogniaux, Cucurbitaceae, in DC. Monogr. Phan. 3: 325-1008. 1881; Cucurbitaceae, Pflanzenr. IV. 275, I, II. 1916 and 1924. John Hutchinson, Cucurbitaceae, in Genera Fl. PI. II: 376-419. 1967. C. Jeffrey, Notes on Cucurbitaceae, including a
DIETERLE: FLORA OF GUATEMALA 307
proposed new classification of the family, Kew Bull. 15: 337-372, 1962; Corrections in Cucurbitaceae, op. cit. 16: 483. 1962; A note on pollen morphology in Cucurbitaceae, op. cit. 17: 473-477. 1964. Thomas W. Whitaker and Glen N. Davis, Cucurbits. London. 1962.
Mostly vines, prostrate or climbing, rarely without tendrils, annual or perennial, monoecious or dioecious, glabrous or variously pubescent, often scabrous; stems herbaceous or woody, sometimes arising from a thickened rootstock or a tuber; sap watery; leaves alternate, usually petiolate, simple and entire, or angular, or variously lobed, sometimes palmately or pedately compound, generally cordate at the base, generally membranaceous, sometimes bearing distinctive glands or villae, venation usually palmate-pedate; stipules absent; petiole sometimes with a stipuliform bract in its axil; tendrils lateral with the petioles (not opposite as in Vitaceae, not in the axils as in Passifloraceae), usually one at each node, simple or branched, the branching of two kinds: 1) proximal, with spiralling only above the point of division, or 2) distal, with spiralling above and below the point of division; inflorescences borne in leaf axils (in monoecious species the staminate and pistillate flowers often arising simultaneously or successively in shared axils); flowers unisexual (very rarely bisexual), small or large, regular, mostly pentamerous (usually with a reduction to 3 in the pistil), arranged in fascicles, racemes, or panicles, or solitary, the pistillate more commonly solitary than the staminate, white, yellow, green, rarely red, or very rarely lilac; floral structure consists of a central portion, very shallow to elongated-tubular in shape, interpreted by some as coherent and adnate portions of calyx and corolla, or of the calyx alone, with sometimes a basal disc, by others as an extension of the receptacle, this central portion (referred to in literature as the calyx tube, or the receptacle tube, or simply the receptacle) supporting the sepals (also called calyx lobes, limb, or teeth) and the corolla (or corolla lobes or petals) on its rim, the stamens on its base or walls, or the pistil with an inferior ovary at its center (for simplicity, the terms receptacle, sepals and petals have been used in the present treatment); sepals (3-) 5 or obsolete, free, open in the bud or valvate; petals (3-) 5, free or united, or partly united, alternate with the sepals; stamens basically 5 (sometimes appearing to be 3 or 2 or 1 through cohesion), inserted on the receptacle or on a basal disc (never on the petals), generally monothecous (the thecae also called sacs, locules, or cells, in the literature), often variously combined to appear to be reduced in number, frequently combined in a 2 - 2 - 1 pattern (two double and one single); filaments long or short or almost lacking, free or united into a column; anthers free, coherent, or confluent, the thecae straight, arcuate, hooked, replicate, or variously contorted, or united in a horizontal ring; connectives narrow or broad, often joined in a globose head, sometimes prolonged as apical appendages; rudimentary ovary sometimes present in staminate flowers; pollen markedly varied in size and morphology within the family, but mostly uniform within genera or species; pistillate flowers with perianth usually like that of the staminate flower, though sometimes larger or smaller; rudimentary stamens sometimes present; ovary usually wholly inferior, placentation parietal, apical, or basal; ovules anatropous, 1-many, horizontal, ascending or pendulous; style 1 (rarely 3, spreading); stigmas linear, globose, spoon-shaped, or bilobed; fruits small or large, dry or fleshy, indehiscent or variously dehiscent, spiny or unarmed, variously shaped, sometimes angulate, gibbous, or winged; seeds large or small, smooth or rough or sculptured, marginate or immarginate, generally compressed, rarely alate; testa usually rigid; endosperm absent.
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Most Cucurbitaceae exhibit, in addition to the above charac- ters, more or less stiff hairs on disclike or bulbous bases, which account for the commonly found scabridity on leaves, also glandlike structures terminating nerves at leaf margins, these structures perhaps useful for riddance of excess water. Great morphological variation may be found in a population, or even on a single vine. This is especially evident in leaf outline, i.e., in the degree of lobing and in the depth of the basal sinus.
Hybridization in the family is apparently extremely rare. Intergeneric or interspecific crosses are seldom achieved, and rarely produce seed beyond the first generation.
A high degree of endemism is exhibited in patterns of geographical distribution. Ranges for most species tend to be restricted. Almost no genera or species, except for a few valued as cultivated plants, and except for a few weedy introduced species, are common to both the Old World and the New.
About 130 genera, best represented in warm or tropical regions in both hemispheres, with 29 in Guatemala. The family is of considerable economic importance, yielding the world's melons, squashes, pumpkins, cucumbers, and gourds. The volume "Cucur- bits" by Whitaker and Davis should be consulted for the more economically important Cucurbitaceae.
Most classifications for the family have been based on fruit and floral characters, especially those of stamen structure. Valuable additional characters are to be found in seeds, pollen, and roots.
Descriptions and keys in the present treatment were written for use with Guatemalan material only, not for greater ranges. Because adequate flowering and fruiting material are seldom collected together, it seemed desirable to prepare two generic keys. The first is based principally on floral characters; the second on fruit and seed characters, with a few necessary exceptions. Many genera are poorly known, or have long been in need of revision. Some of the names used here may prove to be misapplied. Published descriptions and herbarium material at hand were often inadequate. I have not been able to examine types, and have had to rely perhaps too heavily on the literature and on conclusions of preceding in- vestigators. Portions of a manuscript for Guatemalan Cucurbitaceae left by Dr. Paul C. Standley have been quoted extensively.
Two species often mistaken for Cucurbitaceae, Tourrettia lappacea (L'Her.) Willd. (Bignoniaceae), and Gronovia scandens L. (Loasaceae), are discussed at the end of the family.
DIETERLE: FLORA OF GUATEMALA 309
KEY I.
1. Stamens united or partly united, either all the filaments or all the anthers connate. 2. Filaments forming a central column. 3. Pistillate flowers solitary or rarely geminate; ovules numerous, except solitary
in Sechium and Ahzolia. 4. Staminate flowers solitary; sepals markedly elongated, ligulate, 15-20 mm.
long Tecunumania.
4. Staminate flowers disposed in racemes, panicles, or in umbelliform clusters;
sepals small, not ligulate, 0.5-6 mm. long. 5. Filaments connate less than full length; anthers free; ovules and seeds
solitary, pendulous. 6. Plants cultivated; fruits fleshy, edible Sechium.
6. Plants feral; fruits hard-fleshed, not used as food Ahzolia.
5. Filaments connate full length; anthers on an androecial head; seeds more
than one, not pendulous.
7. Fruits gibbous, bursting explosively.
8. Anther locules arranged in a horizontal ring Cyclanthera.
8. Anther locules vertically plicate or arched.
9. Seeds few, orbicular, more than 1 cm. in diameter Hanburia.
9. Seeds numerous, turtle-like in outline, less than 1 cm. long.
Rytidostylis.
1. Fruits symmetrically ovoid or ellipsoid except in Echinopepon torquatus, dehiscent by an apical operculum, not bursting explosively .
Echinopepon.
3. Pistillate flowers clustered on a common peduncle; ovules and seeds always
solitary, pendulous.
10. Perianth of Staminate flowers 4-merous, filaments united to about mid- length Microsechium.
10. Perianth of stamina te flowers 5-merous, filaments united full length. 11. Receptacle conspicuously 10-foveolate, fruits alate, 5-6 cm. long.
Pterosicyos. 11. Receptacle without conspicuous foveolae.
12. Staminate flowers in panicles, fruits fleshy, 3-5 cm. long Parasicyos.
12. Staminate flowers in racemes (except S. kunthii), fruits dry, less than 1 cm. long Sicyos.
2. Filaments 3, separate, anthers united.
13. Flowers night-blooming, always white Polyclathra.
13. Flowers day-blooming or matutinal, some shade of yellow, or greenish. 14. Corolla tubular in part.
15. Filaments elongate, slender, inserted on middle of receptacle tube.
Schizocarpum.
15. Filaments short, thickened, inserted at the bottom of receptacle tube.
Cucurbita. 14. Corolla segments separate to base, or nearly so.
16. Staminate flowers large, petals 15-20 mm. long; fruits fleshy, peponiform .
Cionosicyos. 16. Staminate flowers small, petals 5-7 mm. long; fruit a fibrous berry.
Cayaponia.
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1. Stamens free, 5, or united to appear as 2 or 3 free stamens. 17. Stamens 5, wholly free.
18. Thecae multiflexuous; petal midribs not appendaged; tendrils branching proximally; style 1 Luffa.
18. Thecae simply curved; petal midribs appendaged; tendrils branching distally;
styles 3 Feuillea.
17. Stamens 2 or 3.
19. Staminate flowers solitary. 20. Corolla tubular below.
21. Sepals broadly triangular-ovate, reflexed Sicana.
21. Sepals linear, spreading or erect Cucurbita.
20. Corolla 5-parted to the base, or nearly so.
22. Flowers white, night-blooming; petioles biglandular at apex Lagenaria.
22. Flowers yellow or greenish yellow, day-blooming petioles without glands
at apex. 23. Peduncles of staminate flowers filiform (ours with a collarlike bract
midlength or lower); tendrils simple Momordica.
23. Peduncles of staminate flowers short; bract in axil; tendrils 2-3-parted.
Citrullus.
19. Staminate flowers disposed in spikes, racemes, fascicles, or panicles. 24. Staminate inflorescences racemose, spicate, or subumbellate at the apex of a common peduncle.
25. Stems and tendrils woody, stem trunklike at base Doyerea.
25. Stems and tendrils herbaceous, stem never trunklike at base.
26. Thecae flexuous Luffa.
26. Thecae straight or hooked.
27. Fruits ellipsoid or long-cylindrical, 0.8-6 cm. long; thecae straight.
28. Plants monoecious Melothria.
28. Plants dioecious.
29. Sepals much shorter than petals Anguria.
29. Sepals much longer than petals Gurania.
27. Fruits globose, 8-10 cm. in diameter; thecae hooked at apex.
Posadaea. 24. Staminate inflorescences fasciculate or paniculate.
30. Plants dioecious; staminate flowers in panicles; pistillate flowers in
panicles; tendrils distally bifurcate; styles 3; seed solitary Sicydium.
30. Plants monoecious; staminate flowers in fascicles; pistillate flowers usually solitary; tendrils simple; style 1; seeds numerous Cucumis.
KEY II.
1. Seeds solitary, pendulous from apex of locule.
2. Fruits samaroid Pterosicyos.
2. Fruits various but never samaroid.
3. Tendrils distally furcate; fruits disposed in panicles Sicydium.
3. Tendrils simple or proximally branched. 4. Fruits mostly solitary (rarely geminate).
5. Cultivated plants; fruits used as food Sechium.
5. Feral plants; fruits not used as food Ahzolia.
DIETERLE: FLORA OF GUATEMALA 311
4. Fruits in clusters.
6. Fruits small, not more than 1 cm. long, hard and dry Sicyos.
6. Fruits larger, 1-4.5 cm. long, fleshy.
7. Pericarp conspicuously spotted, without spines; staminate inflorescences
paniculate; perianth 5-merous Parasicyos.
7. Pericarp concolorous or obscurely spotted, with few spines; staminate
inflorescence racemose; perianth 4-merous Microsechium.
1. Seeds few to numerous, not pendulous from apex of locule. 8. Fruits sparsely to densely echinate. 9. Fruits dehiscing explosively.
10. Seeds few, orbicular, more than 1 cm. in diameter Hanburia.
10. Seeds numerous, turtle-shaped, less than 1 cm. long.
11. Receptacle patelliform Cyclanthera.
11. Receptacle elongated, tubular Rytidostylis.
9. Fruits indehiscent, or if dehiscent, not explosive.
12. Fruits rostrate and apically operculate Echinopepon.
12. Fruits not rostrate, indehiscent Cucumis.
8. Fruits not echinate.
13. Seeds 4.5 cm. broad or larger Feuillea.
13. Seeds less than 2 cm. broad.
14. Dehiscence (in ours) by pericarp shattering irregularly from the base up.
Schizocarpum. 14. Dehiscence not as above, or fruits indehiscent.
15. Seeds with conspicuous, fleshy, red covering Momordica.
15. Seeds without conspicuous colored covering. 16. Seeds erect or ascending.
17. Tendrils simple Doyerea.
17. Tendrils 2-5-parted.
18. Pericarp thin, fruits 1-2 cm. long Cayaponia.
18. Pericarp thick, fruits 3-5 cm. long Cionosicyos.
16. Seeds horizontal. 19. Plants dioecious. 20. Tendrils simple. 21. Sepals much shorter than corolla; petals broad, spreading.
Anguria. 21. Sepals much longer than corolla; petals short, thick, erect.
Guarania.
20. Tendrils 2-4-parted Tecunumania.
19. Plants monoecious. 22. Staminate flowers in racemes; pistillate flowers solitary.
23. Fruits dry, fibrous, operculate at apex Luffa.
23. Fruits fleshy, indehiscent.
24. Peduncles slender; fruits ellipsoid, less than 6 cm. long; seeds
3-6 mm. long Melothria.
24. Peduncles grossly thickened; fruits globose, 8-10 cm. in
diameter; seeds not seen Posadea.
22. Staminate and pistillate flowers solitary. 25. Flowers nocturnal, white.
26. Fruits fleshy; petioles biglandular at apex Lagenaria.
26. Fruits dry, petioles eglandular Polyclathra.
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25. Flowers diurnal, or matutinal, yellow or greenish yellow.
27. Pericarp dark red to purple when mature Sicana.
27. Pericarp green, yellow, or orange, never red or purple.
28. Seeds narrowly ovate-oblong Cucumis.
28. Seeds broadly ovate.
29. Fruits firm-fleshed or fibrous, not markedly juicy, seeds
(in ours) marginate Cucurbita.
29. Fruits tender, very juicy, seeds not marginate Citrullus.
AHZOLIA Standley & Steyermark
Scandent herbs, probably monoecious, almost glabrous; leaves long-petiolate, membranaceous, nearly orbicular in outline, deeply cordate at the base, angulate or shallowly 5-lobate, scabrous above, smooth below, the margins entire or remotely and minutely denticulate; tendrils commonly 4-5-parted, with a sturdy peduncle; staminate flowers: inflorescence an elongated, interrupted raceme or narrow panicle with short, few to several- flowered branches, pedicels capillary, perianth 5-parted, receptacle shallowly pateriform, 10-foveolate, corolla spreading, 5-lobed nearly to the base, the lobes broadly ovate-triangular, often granular-tomentulose outside, filaments connate to the middle into a slender column, divaricately separating above the middle, anthers 5, monothecous, thecae replicate; pistillate flowers unknown, but apparently solitary, pedunculate, sharing axils with staminate flowers; fruit moderately large, hard-fleshed, 1-seeded, indehiscent, obovoid, smooth or said to be sometimes shallowly sulcate.
The genus consists of a single species. The name commemorates Ahzol, one of the chieftains of the Guatemalan army of Tecun Uman, who died in battle at the Rio Xequijel near Olintepeque, where the Indian hosts were overwhelmed by the army of Pedro de Alvarado.
Ahzolia seems little different from the closely related genus Sechium. Materials at hand suggest that it may be distinguished from the latter principally through having hard-fleshed, unpala- table fruits. Herbarium specimens of Ahzolia exhibit longer, more slender pedicels, more openly spreading corollas, smaller sepals, and more delicate stamens. The two genera are markedly alike in habit of growth, vegetative characters, floral morphology and pollen structure. Both are monotypic. Further study may reveal that Ahzolia composita and Sechium edule are congeneric, perhaps even conspecific. Sechium fruits vary considerably in size, color, spininess, and in quality of flesh. Perhaps Ahzolia is only an unpalatable form of Sechium which is able to reproduce in a wild state because its fruits are not gathered for food.
Ahzolia composita (Donn.-Sm.) Standl. & Steyerm. Field Mus. Bot. 23: 92. 1944. Microsechium compositum Donn.-Sm. Bot. Gaz. 35: 2. 1903. Huisquil de raton.
FIG. 6. Ahzolia composita. A, leaf, X Vfe; B, staminate inflorescence, X V2; C, staminate flower, upper side, X 3l/2; D, staminate flower, lower side, X 3'/2; E, two views of a shriveled fruit (Standley s.n., Field Museum herbarium sheet no. 991399), X '/2, thought to be of this species. See also Figure 35.
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314 FIELDIANA: BOTANY, VOLUME 24
Dense, wet, mixed forest, often in deep ravines; 200-1,500 m.; Santa Rosa (type from Malpais, Heyde & Lux 6146, PI. Guat. quas ed. Donn.-Sm.); Escuintla; Suchitepequez; Quezaltenango. Mexico (Chiapas).
Large, herbaceous, scandent vines, the stems rather stout, sulcate, smooth, almost glabrous; petioles glabrous, sulcate, 8-20 cm. long; leaf blades up to 22 cm. long and about as broad or broader, scaberulous above, smooth below, deeply and usually narrowly cordate at the base, acute or short-acuminate, angulate, or shallowly 5-7-lobate, the lateral lobes often obtuse, margins entire or remotely and minutely denticulate, principal leaf veins at apex of petiole 5; tendrils unequally 4-5 (-6)-parted, glabrous, on peduncles 2.5-8 cm. long; staminate inflorescence: peduncle glabrous, sulcate, 30-40 (-65) cm. long, laxly many- flowered, the flowers in simple racemes or in narrow panicles, the pedicels slender, 7-15 mm. long, somewhat longer after anthesis, glabrous or glabrate, flowers green or pale green, 9-15 mm. in diameter, receptacle shallowly saucer-shaped, 5 mm. broad, with a cushionlike, 10-radiate center, ringed by 10 strongly defined, circular or semi-circular nectary pits, the pits 0.5-1 mm. wide, sepals green, subulate, reflexed in anthesis, 1-2.5 mm. long, corolla spreading, the 5 lobes broadly ovate, entire, 5-6 mm. long, 7-nerved, often tomentulose outside; filaments glabrous, slender, about 5 mm. long, connate to the middle, the anthers 1 mm. long, monothecous, the thecae doubly sigmoid flexuous; pollen prolate spheroid, finely reticulate, 9-sulcate, short-spiny, about 50x55 ju; pistillate flowers unknown; attached fruits not seen.
Standley left the following notes: "Fruit obovoid, obtuse at each end, in the dry state 6-7 cm. long and 4 cm. broad, when fresh considerably larger, deep green, when dry showing 10 conspicuous winglike ridges, somewhat scabrous on the angles. . . This plant is abundant in many wooded ravines of the Pacific slope, often forming dense mantles of leaves over medium-sized trees. The large fruits, similar to those of the chayote or huisquil, are produced in great abundance, and after the beginning of the dry season fall to the ground in large numbers and are conspicuous there. The fruits are hard and heavy, with solid white flesh, and they remain unchanged for a long time after falling, withering and drying only after many weeks. Mature fruits laid aside to dry for deposit in the herbarium did not lose their moisture until after several months. The fresh fruits are almost wholly smooth, but on the dry ones it may be noticed that the edges of the crests are somewhat scabrous. It may be that the young ovary is somewhat aculeolate. . . Apparently the fruits of Ahzolia are not eaten in Guatemala."
ANGURIA Jacquin
Vigorous vines (? from perennial roots), often climbing several m. into trees and over tall shrubs, glabrous or nearly so, dioecious; leaves petiolate, simple and entire or lobed, or palmately 3-(5-)foliolate; tendrils robust, unbranched; staminate
DIETERLE: FLORA OF GUATEMALA 315
inflorescence spicate or narrowly racemose at the apex of an elongated peduncle; staminate flowers: receptacle tube cylindric or flask-shaped, sepals 5, short, petals 5, free, orbicular to obovate, spreading red or orange-red, membranceous, stamens 2, free, bithecous, described by some authors as one dithecous, one trithecous, dorsifixed, sessile or subsessile in the recetacle tube, connective usually produced into a short appendage which may be glabrous, papillose, or puberulent, anthers linear or oblong, straight, thecae linear, rudimentary pistil none; pistillate flowers: usually solitary in the leaf axils, larger than the staminate flowers, but with a similar perianth, ovary long-cylindric; fruit fusiform-cylindric, short-rostrate, green, striped with lighter green or white, shallowly angled or sulcate, or terete; seeds numerous, horizontal, flattened, asymmetrically ovate, not marginate but sometimes subalate at apex, testa shiny, minutely pitted and roughened, grey. Further details of floral structure, according to the literature: ovary unilocular with 2 placentas; style filiform, bifid, stigmas 2; rudimentary stamens in pistillate flower 2.
Species 30-40, all in tropical America. The name Anguria is an old one taken up by Jacquin and Linnaeus and applied to this genus of New World Cucurbitaceae.
The surfaces of the petals, especially the outer surfaces, are often described as "furfuraceous", which commonly means scaly; however, the petals have not scales but short, multicellular hairs bearing numerous small concretions.
Leaves of Anguria are exceptionally heteromorphic. A single vine may display extreme degrees of lobing or division (e.g., Gentle 5469), or considerable variation in the shape of lobes or leaflets. Hence the outline of the blade, as shown on herbarium specimens, should be used with reservation in distinguishing species. The type of adornment on the appendage of the stamens may be a similarly variable character, and also of questionable value. Setting aside these two characters, specimens from Guatemala and Chiapas appear to be fairly readily separable into two groups, each group presenting a complex of consistently correlated characters, and each group representing, therefore, possibly only one species. Further assessment, when more material is available, may expose this appraisal as oversimplified. The two groups may be distinguished as follows:
Receptacle tube flask-shaped, not speckled; calyx teeth subulate, arching; staminate flowers sessile, a few villae interspersed on the peduncle; older scars from fallen flowers disc-shaped; plants completely glabrous except on or near vegetative buds, and between staminate flowers; leaves usually 2-3-lobed A. diversifolia.
Receptacle tube cylindric, speckled with small, pigmented areas; calyx teeth broadly triangular, obtuse, pigmented at apex; staminate flowers short-pedicellate; older scars from fallen flowers shriveled; portions of stems, petioles, petiolules, or tendrils minutely puberulent; leaves usually 3-petiolulate A. warscewiczii.
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Anguria diversifolia Cogn. in Donn.-Sm. Bot. Gaz. 16: 10. 1891.
Wet forest and thickets, often along streams, in ravines or on high ridges, near sea level to 1,360 m.; Alta Verapaz; Izabal; Peten; Retalhuleu; San Marcos. Mexico; British Honduras; Honduras.
Extensive vines, completely glabrous except for dense hispidity on young vegetative buds in leaf axils, and some villae on the peduncles at the bases of staminate flowers; stems striate, sparingly branched, internodes mostly 7-16 cm. long; petioles striate, 1-3 cm. long; leaves entire or lobed, sometimes 3-partite, the margins usually entire, the principal nerves rather prominent below, surface smooth above and below; staminate flowers: numerous, sessile or nearly so, on upper portion of sturdy peduncles often much longer than the leaves, older scars from fallen flowers flat, disc-shaped, receptacle tube elongate, flask-shaped, sepals subulate to triangular or ovate-triangular, arched-spreading, 1-2 mm. long, petals ovate to obovate, red or red-orange, spreading, appendage of the connective short, glabrous; pistillate flowers and fruits: see description for the genus.
Anguria warscewiczii Hook, f., Curt. Bot. Mag. 88 (ser. 3), t. 5304. 1862.
Wet forest and thickets, sea level to 600 m.; Izabal; Peten. British Honduras; Honduras.
Spreading vines, nearly glabrous throughout, but minutely puberulent on portions of stems, petioles, petiolules, and tendrils; stems striate, sparingly branched, the internodes mostly 8-16 cm. long; leaves membranaceous, smooth, usually trifoliolate, the lateral folioles a little shorter and auriculate, sometimes bifid, the petioles 2-5 (-6) cm. long, the petiolules 0.6-1 cm. long, the margins entire, undulate, or distantly and shallowly toothed, sometimes coarsely serrate, the surfaces smooth above and below, the principal veins rather prominent below; staminate flowers: numerous, short-pedicellate or subsessile on sturdy peduncles much longer than the leaves, pedicels short and thick, older scars from fallen flowers short-stalked, shriveled, receptacle tube long-cylindric, speckled with small, pigmented areas, sepals triangular, obtuse, pigmented at the tips, scarcely 1 mm. long, petals ovate-oblong or suborbicular, spreading, more or less pubescent on both surfaces; pistillate flowers and fruits: see description for the genus.
CAYAPONIA Manso
References: A. Cogniaux in DC. Monogr. Phan. 3: 738-743. 1881. C. Jeffrey, Kew Bull. 25: 201-234. 1971.
Herbaceous or suffrutescent vines, generally scandent, from perennial roots, monoecious; leaves petiolate, palmately 3-7-lobate, or entire, or rarely 3-5-foliolate, sometimes decurrent on the petiole, the blade often bearing disc-shaped glands, these flat- margined, oval or round; tendrils 2-5-parted or rarely simple; flowers usually not more than 1 cm. across, yellowish green or whitish, solitary or the inflorescence fasciculate or racemose; staminate flowers: receptacle usually campanulate, the sepals 5, corolla campanulate or spreading, deeply 5-parted, the lobes ovate or oblong,
FIG. 7. Anguria diversifolia A, habit, X '/2; B, and C, immature staminate flowers, two dissected to uncover stamens, X 4; D, diagrammatic sketch of older flower with partly expanded petals, X 4.
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318 FIELDIANA: BOTANY, VOLUME 24
5-7 mm. long, stamens 3, inserted on the base of receptacle, the filaments free, remote, anthers generally coherent, one monothecous, the others dithecous, the thecae longitudinally replicate, the connective not produced beyond the thecae; pistillate flowers: perianth similar to that of staminate flower but smaller, ovary globose, ovoid, or oblong, the locules said to be 3, sometimes fewer by abortion, ovules 1-few in each locule, ascending from the base, style erect, inserted on a basal trilobate or annular disc, stigmas 3, dilated, reflexed, papillose or smooth, staminodia 3, minute; fruit small, 1-2 cm. long (in ours), ellipsoid or globose, fibrous or corky, indehiscent, unarmed, 1-several-seeded; seeds erect, ovate or oblong, subcompressed, not or only faintly marginate, the testa usually rigid and smooth.
One of the largest genera in the Cucurbitaceae; numerous species in tropical South America, a few in Mexico and Central America. All the fertile Guatemalan specimens examined by me can be referred to either of two widespread species: C. attenuata (Hook.
6 Arn.) Cogn., or C. racemosa (Mill.) Cogn. Two other species with comparatively larger, peponiform fruits, transferred by Jeffrey from Cayaponia to the genus Cionosicyos [as Cionosicys], are described here under Cionosicyos. Four sterile sheets at hand, from Huehuetenango and San Marcos, Standley 68961, Steyermark 37157, 48812, and 51731, all with trifoliate, long-petiolulate leaves, if properly referable to Cayaponia, resemble C. petiolulata Cogn., but may represent an undescribed species.
Unfortunately, leaves and flowers of the two species treated here, C. attenuata and C. racemosa, are so similar that the plants can be positively distinguished only when fruits are available.
Fruits globose; leaves commonly deeply trilobate to or below the middle; receptacle of pistillate flowers about 1 mm. in diameter C. attenuata.
Fruits ovoid to ellipsoid; leaves commonly shallowly lobate but sometimes some leaves may be deeply lobate; receptacle of pistillate flowers about 2 mm. in diameter C. racemosa.
Cayaponia attenuata (Hook. & Arn.) Cogn. in DC. Monogr. Phan. 3: 769. 1881. Anguria dubia Hook. & Arn. Bot. Beechey Voy. 292. 1840. Bryonia attenuata Hook. & Arn. op. cit. 424. 1840. C. dubia (Hook. & Arn.) Rose, Contr.U. S. Nat. Herb. 5: 121. 1897.
Moist or dry thickets, hedges, second growth at low altitudes, 900 m. or lower; Chiquimula; Escuintla; Jutiapa; Retalhuleu; Santa Rosa; Suchitepequez; Zacapa. Mexico; British Honduras to El Salvador and Panama; West Indies.
Vigorous, much-branched vines; stems tough, deeply sulcate, glabrous or subglabrous; leaves on slender petioles mostly 1-5 (-10) cm. long, the blades mostly 3-
7 (-15) cm. long, commonly deeply 3-lobed to or below the middle, usually the upper leaves not lobed and much smaller, lobes obtuse or acute, the median one larger and usually constricted at the base, veins glabrous on lower surface, densely short-hispid
FIG. 8. Cayaponia attenuata. A, habit (portion of a lower stem with lobed leaves), x Vfc; B, habit (portion of an upper stem with unlobed leaves), X Vfc; C, detail of a branchlet with pistillate flowers and a foliar gland (one greatly enlarged at left), X 2; D, trilobate disc at base of style in opened pistillate flower, X 5; E, pistillate flower, X 3; F, staminate flower, X 3; G, stamens and basal disc in opened staminate flower, X 3. See also Figure 36.
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320 FIELDIANA: BOTANY, VOLUME 24
on upper surface and often scabrous, margins remotely denticulate or subentire, base of the blade usually attenuate, then narrowly decurrent on the petiole and bearing several disc-shaped glands; tendrils simple or 2 (rarely more)-branched; flowers mostly on branchlets with suppressed or somewhat reduced leaves and tendrils, single or frequently paired in the axils, or in short axillary racemes, staminate and pistillate flowers on the same vines but seldom equally abundant together; staminate flowers: pedicels slender, about 5 mm. long, receptacle broadly campanulate, 3-4 mm. broad, sepals short-triangular to linear, erect, remote, corolla deeply parted, subrotate, about 8 (-10) mm. across, petals separate nearly to the base, oblong or ovate, the upper margins flared laterally, minutely and densely spiculate or papillose outside, appressed-villous inside, the flared portion membranaceous, whitish or yellowish, the median portion firmer and greenish, stamens inserted at base of receptacle, filaments slender, villous at base, anthers 2 to 2'/2-plicate, basal disk 3-lobed; pollen globose, spiny, 8-porate, about 70-75 fi in diameter; pistillate flowers: much smaller than the staminate, perianth similar but with petals relatively narrower and not flared along the upper margins and with the sepals relatively narrower and longer, pedicels about 5 mm. long, glabrous or nearly so, ovary globose, nearly glabrous, ovules said to be about 3, receptacle short-campanulate, 1-1.5 mm. across, abruptly and naprowly constricted at the base, nearly glabrous, basal disc 3-lobed, free from the receptacle, sepals minute, dentiform, erect, remote, corolla about 6 mm. across, petals oblong, obtuse, erect or subrotate, minutely and densely spiculate or papillose outside, appressed-villous inside, staminodia very small, linear, glabrous, style slender, linear, stigmas smooth, not papillose; fruit globose, commonly about 1 cm. in diameter, green, thin-walled, commonly 3-seeded, very bitter; seeds ovate, subcom pressed, notched at the hilum, about 7X4x2 mm., smooth, brown or brindled, the edge often banded with a narrow, tawny stripe.
A frequent vine in disturbed growth at low altitudes. The crushed, dry stems and fruits are said to be used in place of soap for washing clothes.
Cayaponia racemosa (Mill.) Cogn. in DC. Monogr. Phan. 3: 768. 1881. Bryonia racemosa Mill. Gard. Diet., ed. 8. 1768. Melocotoncillo de raton (Guatemala, fide Aguilar).
Damp thickets or forest, near sea level to 1,500 m.; Alta Verapaz; Chiquimula; Escuintla; Guatemala; Izabal; Jalapa; Ju- tiapa; Peten; Quezaltenango; Retalhuleu; Santa Rosa. Mexico; British Honduras and Honduras to Panama; West Indies; Colombia and Venezuela.
Usually large, branching vines; stems tough, sulcate, essentially glabrous; leaves on slender petioles mostly 2-6 cm. long, the blades mostly 4-13 cm. long, 3-17 cm. wide, commonly shallowly trilobate, sometimes some of the lower leaves deeply lobed, rarely 5-lobed, or the upper leaves not at all lobate and smaller, lobes obtuse or acute, the median one larger, veins usually glabrous on lower surface, the upper surface short-hispid and often scabrous, margins remotely denticulate or subentire, base of blade usually attenuate or cuneate and then narrowly decurrent on the petiole, with several disc-shaped glands; tendrils simple or bifid (rarely more than two-parted); flowers mostly on branchlets with suppressed or reduced leaves and
DIETERLE: FLORA OF GUATEMALA 321
tendrils, single or paired in axils, or in short axillary racemes, white or greenish white, staminate and pistillate flowers on same vines but seldom equally abundant together; staminate flowers: pedicels slender, about 5 mm. long, receptacle broadly campanulate, about 5 mm. broad, sepals short-triangular or almost linear, erect, remote, corolla deeply parted, subrotate, about 1 cm. across, petals ovate, minutely spiculate outside, villous inside, stamens inserted near base of receptacle, filaments slender, villous at base, anthers 2 to 21/2-plicate, basal disc trilobate; pistillate flowers: much smaller than the staminate, perianth similar but with narrower petals, pedicels 5-8 mm. long, receptacle about 2 mm. across, abruptly constricted at base, sepals minute, erect, ovary ovoid, essentially glabrous, ovules said to be 3, basal disc trilobate, corolla about 8 mm. across, petals oblong, obtuse, erect or subrotate, minutely and densely spiculate or papillose outside, appressed-villous inside, staminodia minute, style slender, stigmas smooth, not papillose; fruit ovoid to ellipsoid, 1.5-2 cm. long, about 8 mm. in diameter; seeds narrowly ovate to oblong, notched at the hilum, about 11 X 5 X 2.5 (X 3) mm., smooth, greenish brown or brindled.
CIONOSICYOS Grisebach
Reference: C. Jeffrey, Further Notes on Cucurbitaceae: II, Kew Bull. 25: 200-201. 1971.
Scandant vines, herbaceous or suffrutescent, probably perennial, monoecious; leaves petiolate, the blades commonly palmately 3-5-lobate, sometimes unlobed, often decurrent on the petiole and bearing at the base one or more disc-shaped, flat- margined glands; tendrils bifid or sometimes simple; flowers of medium size, at least the staminate ones more than 1 cm. across (often 2-4 cm.), white or greenish, usually solitary in the axils or disposed in short racemes or on branchlets with suppressed or reduced leaves and tendrils; staminate flowers: receptacle more or less campanulate but when dried often appearing funnelform, sepals 5, corolla campanulate, 5-parted almost to the base, petals suberect, ovate or oblong, 15-20 mm. long, stamens 3, filaments free, villous at base, anthers generally coherent, one monothecous, the others dithecous, thecae tightly replicate; satisfactory pistillate flowers not seen; ovules said to be numerous, borne on 3 large, intrusive, parietal placentae, the ovules ascending in the lower part, more or less horizontal in the upper; fruits medium size, globose, pepolike, smooth, unarmed, the pericarp thick; seeds ovate or oblong-ovate, subcompressed, not margined, testa rigid and smooth.
Grisebach's name for the genus appears in his Flora of the British West Indies Islands (p. 288. 1860) as Cionosicys (not Cionosicyos). Hooker, f. (in Benth. & Hook, f., Gen. PI. 1: 826. 1867) and Cogniaux (in DC. Monogr. Phan. 3: 516. 1881) apparently believed this spelling to be unintentional and wrote the name as Cionosicyos. Jeffrey (1971) reverted to the spelling of the original publication when he transferred two species from Cayaponia. Since the Greek word for cucumber is olkuos, and I am not convinced that Grisebach really wished the omission of the final o, I have chosen to join Hooker, f., and Cogniaux in using the revised form Cionosicyos.
322 FIELDIANA: BOTANY, VOLUME 24
Three species, all American, with two in Guatemala.
Sepals lance-ovate, constricted at the base, 7-13 mm. long C. excisus.
Sepals dentiform or triangular, 2-3 mm. long C. macranthus.
Cionosicyos excisus (Griseb.) C. Jeffrey (as Cionsicys), Kew Bull. 25: 201. 1971. Sicana excisa Griseb. Cat. Fl. Cub.: 112. 1866. Cayaponia excisa Cogn. in DC. Monogr. Phan. 3: 747. 1881. C. alata Cogn. torn. cit. 746.
Damp to dry thickets and forest, sea level to 300 m.; Peten. Yucatan Peninsula of Mexico; British Honduras; Cuba.
Large or small herbaceous or suffrutescent vines, climbing over shrubs and small trees, the stems essentially glabrous; leaves on slender petioles 1-4 cm. long, the blades broadly angulate-ovate or cordate-ovate in outline, 3-15 cm. long, shallowly or deeply 3-5(-7)-lobate or sometimes merely rounded-angulate, the lobes acute or obtuse, the central one often constricted or narrowed at the base, the upper surface glabrous or nearly so, the lower sparsely pubescent or sometimes hispidulous on costa and veins, or essentially glabrous, often punctate and more or less scabrous, margins minutely denticulate or entire; staminate flowers: solitary, the pedicels 0.5-3 cm. long, receptacle broadly campanulate, acute at the base, glabrous or more or less puberulent or sparsely hispid, 7-12 mm. long, sepals broadly lance-ovate, acute, constricted at the base, mostly 7-13 mm. long, foliaceous, 3-5-nerved, nearly glabrous or more or less scabridulous or punctate, usually minutely ciliolate, corolla broadly campanulate, petals oblong, almost 2 cm. long, white, greenish white, cream, or greenish pale yellow, papillose outside, villous within, especially near the base, acute, short-acuminate, or truncate and somewhat apiculate, filaments villous, especially at the base, the anther head about 9 mm. long; pistillate flowers: outwardly similar to the staminate ones but considerably smaller, ovary hairy; fruit globose, 3-5 cm. in diameter, green, said to become pale tomato-red when mature; immature seeds 5-7 mm. long; mature seeds not seen.
Cionosicyos macranthus (Pittier) C. Jeffrey (as Cionosicys), Kew Bull. 25: 200. 1971. Cayaponia macrantha Pittier, Contr. U. S. Nat. Herb. 13: 118. 1910. C. microdonta Blake, op. cit 24: 27. 1922 (type from Izabal, Los Amates, Blake 7730).
Damp or wet forest, often pendant from trees, in open thickets or along streams, 200-500 m.; Alta Verapaz; Izabal; Peten. Mexico; British Honduras; Honduras; Costa Rica.
Large herbaceous or suffrutescent vines, the stems sulcate, essentially glabrous, said to be as much as 9 m. long and 2.5 cm. thick below; leaves on petioles 1-3 (-4) cm. long, the blades firm, broadly angulate-ovate or cordate-ovate in outline, commonly deeply 3-5-lobate, sometimes unlobed, mostly 7-12 cm. long, the central lobe longer, obtuse, acute, or rarely abruptly acuminate, often constricted at the base, the lateral lobes rounded or auriculate at the base, blades often cuneate at the base and then more or less decurrent on the petiole, essentially glabrous on both surfaces or more or less hispidulous beneath, especially on costae and veins, both surfaces often somewhat scabrous or scabridulous, margins subentire or obscurely
FIG. 9. Cionosicyos macranthus. A, habit (portion of a vine with 3-lobed leaves and a young staminate flower), x Vz; B, portion of a vine with unlobed leaves and nearly mature fruits, x Vz C, seed, X 1; D, another leaf form and a staminate bud, x Vz; E, pistillate flower, partially dissected, X %; F, diagrammatic sketch of receptacle of pistillate flower showing stigmas, style, and staminodia (usually 3 in number, not 5), x 3/4; G, stigma, x 2Vz; H, staminate flower opened to show stamens, X Wz; I, another leaf form, X Vz.
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324 FIELDIANA: BOTANY, VOLUME 24
denticulate, basal sinus obscure or broad and shallow; tendrils bifid or rarely simple; flowers solitary, on pedicels 1.5-4 cm. long; staminate flowers: receptacle tubular- campanulate, 10-12 (-14) mm. long, minutely puberulent and sparsely or densely hispid outside, pilose within, sepals dentiform or triangular, acute or acuminate, remote, 2-3 mm. long, corolla campanulate, petals oblong-ovate, about 1.5 cm. long, white or cream-colored, papillose outside, long-villous within, anthers about 11 mm. long; pistillate flowers: similar to staminate but smaller, petals about 12 mm. long; fruit green striped with cream or pale green with darker green stripes, subglobose or globose, 3-5 cm. in diameter; seeds pale, about 1 cm. long.
CITRULLUS Forskal. Watermelon.
Annual or perennial vines, monoecious; leaves petiolate, rounded to triangular- ovate in outline, deeply lobed, the lobes lobulate; solitary axillary bracts present; tendrils proximally 2-4-parted or simple; flowers moderately large, solitary, yellow, short-pedunculate; staminate flowers: receptacle broadly campanulate, short, the sepals 5, narrow, remote, corolla spreading or broadly campanulate, deeply 5-parted, the lobes ovate to ovate-oblong, stamens 3, inserted on the base of the receptacle, filaments short, free, anthers free or slightly coherent, one monothecous, the other two dithecous, thecae flexuous, the connective broad, not appendaged, rudimentary pistil present; pistillate flowers: perianth like that of the staminate flowers, ovary globose or ovoid to ellipsoid, triplacentiferous, ovules numerous, horizontal, style short, stigmas 3, sub-bilobate, fleshy, staminodia 3, short, slender; fruit globose or ellipsoid, large, fleshy, not hollow, indehiscent; seeds numerous, ovate in outline, compressed, smooth or slightly rough, not marginate but with a pair of rounded protuberances on each side of the hilum.
Several species, one in Guatemala, ruderal or cultivated.
Citrullus lanatus (Thunb.) Matsumura & Nakai, Cat. Sem. Hort. Bot. Univ. Tokyo 38. 1920. Cucurbita citrullus L. Sp. PI. 1010. 1753. Momordica lanata Thunb. Prodr. Fl. Cap. 13. 1794; op. cit 36. 1807. Citrullus vulgaris Schrader ex Ecklon & Zeyher, PI. Afr. Austr. 279. 1834. Sandia.
Commonly cultivated in Guatemala and other Central Amer- ican countries mostly at 300 m. elevation or less.
Strong- growing, annual vines, usually prostrate, softly hairy to densely lanate on younger portions, becoming scabrous in age; stems herbaceous, long-hairy at first, then glabrous; leaves on long-hairy petioles 4-12 cm. long, the blades membrana- ceous, ovate-triangular in outline, deeply 3-5-lobed, the lobes lobulate, the central lobe much the largest, margins shallowly sinuate-toothed; tendrils usually bifid; axillary bracts obovate-spathulate, hooded, glabrous, to 14 mm. long; staminate flowers: peduncles 1-2 (-3) cm. long, hairy, receptacle broadly campanulate, sepals linear- triangular, 3-4 mm. long, corolla 5-parted to the base or nearly so, the petals broadly ovate, 10-15 mm. long, obtuse and mucronate, dull yellow with green nerves, filaments sturdy, densely hairy at the base on the ventral side, anthers slightly exserted; pistillate flowers: peduncles 5-9 mm. long, hairy, ovary globose to ovoid, villous, perianth like that of the staminate flowers, rudimentary stamens linear, style
FIG. 10. Citrulhis lanatus. A, habit, X '/a; B, seedling plants, X '/•>; C, staminate flower, X 2'/2j D, seeds, X Vfc; E, small mature fruit (from a photograph). See also Figure 37.
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326 FIELDIANA: BOTANY, VOLUME 24
glabrous; fruit glabrous, green or marked with longitudinal, broken stripes of light and dark green, inner flesh yellowish to red; seeds 7-10 mm. long, 4-6 mm. wide, pale to dark brown, or mottled.
Standley, in manuscript, has stated, "Watermelon plants have been seen growing along railroad embankments and occasionally in waste places where seeds have been discarded. It is doubtful if the melons could reproduce themselves for any fruit produced would be eaten by people or by animals."
CUCUMIS Linnaeus Reference: A. Cogniaux, Pflanzenreich IV, 275, II: 116. 1924.
Annual, monoecious vines of moderate length, usually scabridly or setosely pubescent, mostly prostrate; leaves long-petiolate, angulate, or 3-7-lobate or unlobed; tendrils simple (in ours); flowers yellow, rather small; staminate flowers: axillary, fasciculate, rarely solitary, pedicels slender, receptacle campanulate, usually narrowly so, sepals 5, subulate, corolla deeply 5-parted, spreading, stamens 3, free, inserted on the tube of the receptacle, the filaments short, the anthers oblong (one of them monothecous, the others dithecous), the connective produced at the apex into an appendage; pistillate flowers: usually solitary, perianth like that of the staminate flower, staminodes 3, setiform or liguliform, ovary globose, ovoid, or subcylindric, tubercled or softly spiny, glabrous or hairy, usually 3-placentate, ovules horizontal, numerous, style short, columnar, surrounded by a disk at the base, stigmas 3, globose, obovate or flattened and somewhat bilobate, sometimes connivent; fruits globose to long-cylindric, fleshy, many-seeded, indehiscent; seeds rather small, numerous, white, smooth, compressed, narrowly ovate-oblong, subacute at each end, immarginate.
An Old World genus of about 30 species, principally represented in Africa. In addition to the cultivated cucumbers and melons produced in Guatemala, two other Cucumis entities, not cultivated, occur there. One of these, C. anguria L., an easily distinguished species, thought by Standley to be native in America, is regarded by most authors as of African origin. The other, commonly determined as a variety of C. melo (var. dudaim Naud. or var. chito Naud.) resembles the cultivated C. melo in having densely hairy ovaries, unarmed fruits, and more or less rounded leaf blades. Widely distributed in southern United States, Mexico, Central America, and perhaps elsewhere, its status as a variety of C. melo rather than as a separate species needs study.
Fruit peduncles more than 4 cm. long (usually 6-15 cm.); leaves deeply lobed.
C. anguria.
Fruit peduncles less than 4 cm. long (often less than 3 cm.); leaves entire, angulate, or shallowly lobed.
Ovary densely hairy; fruits unarmed C. melo.
Ovary hispid or tuberculate; fruits sparingly armed C. sativus.
DIETERLE: FLORA OF GUATEMALA
327
FIG. 11. Cucumis anguria. A, habit (node with a mature leaf and a fruiting branchlet), X Vz; B, fruits, two sectioned to show seeds, X Vz; C, two flowers (pistillate at left, staminate at right), X 2l/2; D, staminate flower opened to show stamens, X 4; E, pistillate flower opened to show stigmas, style, and cup-shaped disc at base of style, x 4.
Cucumis anguria L. Sp. PL 1011. 1753. Sandia de raton.
Mostly on sandy or rocky plains, or rocky slopes, at low altitudes, rarely above 700 m., probably an early introduction to our region; Chiquimula; Escuintla; Peten; Retalhuleu; Santa Rosa; Zacapa. Mexico; British Honduras; Honduras; West Indies; Colombia.
328 FIELDIANA: BOTANY, VOLUME 24
Rough, annual, herbaceous vines; stems 1-2 m. long, hispid or spiculate, prostrate, branched; leaves on slender, coarsely hispid petioles mostly 3-12 cm. long, the blades villous-hispid, broadly ovate in outline, 3-8 cm. long, deeply 3-5-lobed, sometimes compoundly lobed, the lobes obtuse, basal sinus relatively broad and shallow, margins finely undulate-denticulate; tendrils unbranched, slender, rather short; staminate flowers: small, about 1 cm. broad or less, pale yellow, borne in few- flowered, axillary fasciculate or subfasciculate clusters, the flowers of these reaching full anthesis one at a time, pedicels filiform, 1-2 cm. long, hispid, receptacle tube narrowly campanulate to funnelform, 5-7 mm. long, hispid, sepals subulate, 3-4 mm. long, hispid, corolla lobes ovate, acute; pistillate flowers: very long-pedunculate, solitary, usually produced at first nodes of slender branchlets off principal stems, peduncles at first 0.5-2 cm. long, soon elongating to as much as 15 cm., ovary globose, ovoid, or subcylindric, hispid or aculeate, perianth similar to that of staminate flowers; fruit subglobose to oblong-ellipsoid, to about 5 cm. long, sparsely and shortly echinate, fleshy with a juicy, many-seeded center and a firm, thin rind, pale green becoming pale yellow, edible when young, odor like that of cucumbers, peduncles more than 4 cm. long, commonly 6-15 cm. long, usually of sturdier girth than the supporting branchlets; seeds dull white, 4-5 mm. long.
Young tender fruits are said to be used like cucumbers, raw or prepared as pickles.
Cucumis melo L. Sp. PI. 1011. 1753. Melon; muskmelon; cantaloupe. (Fig. 38).
Believed to be native of southern Asia and of tropical Africa; widely cultivated in temperate and tropical regions.
Medium-sized, annual, herbaceous vines, usually prostrate, seldom more than 2 m. long, hairy when young, coarsely hispid or scabrous in age; leaves suborbicular to ovate in outline, sometimes shallowly lobed, the lobes rounded, margins shallowly undulate-denticulate to nearly entire, basal sinus rectangular to broadly arched; tendrils of moderate length, unbranched; pistillate flowers usually produced on short branchlets from the principal stems, ovary ovoid to subcylindric, densely pubescent; fruit fleshy, globose or nearly so, terete or shallowly furrowed lengthwise, the surface smooth or more or less roughly netted, nonaculeate; seeds 7-8 mm. long.
The quality of the cantaloupe produced in the Central American region has improved greatly in recent years due to agronomic research and the production of high quality fruits suitable to our climate. The melons seem to do best in the river valleys where irrigation is possible.
The species varies greatly, e.g., in the dimensions and texture of the vines, and in the size and quality of the fruits. Slender, small- fruited, very scabrous variants, encountered mostly on roadsides or ruderal areas, are often determined as C. melo var. dudaim Naud., also as var. chito Naud., and as var. agrestis Naud., following a segregation of categories attempted by Naudin (Ann. Sci. Nat. IV:
DIETERLE: FLORA OF GUATEMALA 329
34, 73. 1859), one not unanimously regarded as phylogenetically acceptable. The taxonomic status of these plants needs clarification.
Cucumis sativus L. Sp. PL 1012. 1753. Pepino; pepino de ensalada; pepino de Espana; cohombro. Cucumber.
A cultivated species, sometimes ruderal as an escape from cultivation, about which Standley wrote: "Native of the East Indies but grown for its fruit generally in tropical and temperature regions; planted commonly in Guatemala, chiefly at middle but also at low elevations."
Medium-sized, herbaceous vines, trailing or scandent, sparingly branched, more or less scabrous and spiculate; tendrils rather long; leaves petiolate, simple, membranaceous, angulate or shallowly angulate-lobed, the lobes triangular, acute; staminate flowers: fasciculate, pedicels slender, receptacle narrowly campanulate, densely villous, sepals subulate, spreading, corolla up to 3 cm. broad, petals broadly ovate, patent; pistillate flowers: solitary, peduncle short, robust; fruit usually oblong, cylindric or obscurely trigonous, sparsely tuberculate, spiculate on the tubercles, yellow and smoother when mature.
CUCURBITA Linnaeus
References: L. H. Bailey, The domesticated Cucurbitas, Gentes Herb. 2: 63-115, /. 29-64. 1929; Species of Cucurbita, op. cit. 6: 267- 322, f. 140-165. 1943; E. F. Castetter and A. T. Erwin, A systematic study of squashes and pumpkins, Bull. Iowa Agric. Exp. Sta., 244: 107-135. 1927; Paul D. Hurd, Jr., E. Gorton Linsley, and Thomas W. Whitaker, Squash and gourd bees (Peponapis, Xenoglossa] and the origin of the cultivated Cucurbita, Evolution 25: 218-234. 1971; Cucurbits, Thomas W. Whitaker and Glen N. Davis, London 1962.
Annual or perennial vines, herbaceous, monoecious, prostrate or scandent, hispid or setulose-pilose; leaves petiolate, the blades more or less lobate with a deep sinus at the base; tendrils simple or multiparted; flowers large, solitary, showy, matutinal or diurnal, the staminate usually long pedunculate; staminate flowers: receptacle campanulate, sepals 5, linear, corolla some shade of yellow or orange, campanulate, 5-lobate to the middle or more deeply, stamens 3, inserted at the base of the receptacle, filaments short and fleshy, closely positioned but usually free, anthers linear, connivent to form a cylindrical column, one of them monothecous, the others dithecous, the thecae elongate, vertically and tightly sigmoid-flexuous, the connective narrow, not produced at the apex; pollen large, globose, spiny, multiporate, non- colpate; pistillate flowers: short-pedunculate, receptacle short, perianth like that of staminate flowers, staminodia 3, short, triangular, ovary ellipsoid, ovoid, or globose, 3-5-placentate, style short, thick, the stigmas usually 3, bilobate, ovules numerous, horizontal; fruit a pepo, fleshy or fibrous; seeds numerous, ovate, strongly compressed, smooth, marginate or immarginate.
330 FIELDIANA: BOTANY, VOLUME 24
Perhaps about 20 valid species, including five cultivated ones, probably all of them American in origin. The genus is doubtless the most economically important in the family, but its taxonomy is far from well understood. The species to be found in Central America, and ways to distinguish them, are as yet unclear and are likely to remain so until more field work is undertaken. Ripe fruits and seeds, roots, buds and flowers, fully developed leaves, habit and habitat notes, are among items needed for delineating the species. Herbarium specimens are often hardly more than a flower and a leaf. Only two species are treated here: one a feral lowland plant with small bitter fruits (C. lundelliana), the other a highland perennial, feral or cultivated, with edible fruits (C. ficifolia). Descriptions given here for these two taxa have been amplified with excerpts from the literature. When better materials are available, it will doubtless become evident that more species are represented in Guatemala. No attempt has been made to include the cultivated Cucurbitas. A general discussion of these may be found in the bulletin by Castetter and Erwin (I.e.) and in "Cucurbits" by Whitaker and Davis (I.e.).
Cucurbita ficifolia Bouche, Verh. Ver. Gartenb. Berlin 12: 205. 1837. C. melanosperma Gaspar. Act. Acad. Napol. 1847; Ann. Sci. Nat. IV. 9: 208, t. 2, f. 4-7. 1855. Chilacayote; cidracoyote (Jalapa); ccoocc, elaoc, ooc (El Quiche).
Cultivated for its edible fruit, grown abundantly in Guatemala at middle and high elevations, mostly at 1,500 m. or more, said to be thoroughly naturalized in some localities, producing fruit, and reproducing itself.
Plants perennial but producing fruit the first year from seed and often cultivated as an annual, sometimes growing to the tops of tall trees, but when cultivated usually prostrate, the stems stout, sparsely or densely setulose-pilose throughout; leaves on petioles 5-20 cm. long, the blades subreniform to circular-ovate in outline, often as much as 25 cm. broad, shallowly or deeply lobate, the lobes usually rounded, the basal sinus deep and open, or the basal lobes subauriculate; tendrils multifid on sturdy peduncles; staminate flowers: long-pedunculate, receptacle tube 5-7 mm. long, sepals linear, acuminate, corollas yellow to light orange; pistillate flowers: short- pedunculate, the corollas as much as 12 cm. broad; fruit large and heavy, ovoid to globose, 15-35 cm. long, green with white blotches and stripes, the shell hard and durable, the flesh firm and white, the peduncle of medium length, 5-6 cm. long, somewhat expanded at attachment to fruit; seeds round-ovate, narrow-margined, flat, 1.5-2 cm. long, said to be black when fully mature.
These plants are grown extensively in the mountains of Guatemala. In the highlands during the dry season, one may see
DIETERLE: FLORA OF GUATEMALA 331
immense numbers of the heavy fruits piled about the houses. Often large numbers are laid along the eaves or on the roofs of the low houses to ripen, or perhaps also to keep them away from domestic animals. The fruits are highly esteemed for making desserts of various kinds, the flesh usually cooked in thick syrup. They are also grown as food for cattle, horses, and sheep during the long dry season when there is scant pasture.
According to Standley, "At some places in barrancos on the lower slopes of the Volcan de Fuego, toward Antigua, chilacayote vines have become thoroughly naturalized and look like native plants, climbing to the tops of high trees, and forming dense tangles of stems and foliage."
Cucurbita lundelliana Bailey, Gentes Herb. 6: 297. 1943. Ayote de caballo (Retalhuleu); ixbulut (Peten); bitter pumpkin, wild pumpkin (British Honduras).
Dry thickets, plains or low hills, sea level to 300 m.; Escuintla; Peten; Retalhuleu. British Honduras.
Herbaceous annual to perennial vines, prostrate or scandent over low shrubs, the stems slender and elongated, sometimes rooting at the nodes, older parts sulcate, thinly hirsute or glabrate; tendrils mostly simple; leaves on slender, softly pilose petioles usually longer than the blades,, mostly 2-12 cm. long, blades broadly ovate to subreniform in outline, usually as broad as long or broader, mostly 4-12 cm. wide, broadly and openly cordate at the base, sometimes white -spot ted, usually rather deeply 5-lobate, the lobes obtuse but mucronulate, at least the central one constricted at the base, margins finely denticulate, upper surface thinly short-pilose with appressed hairs or hispidulous and scabrous, lower surface more densely pilose, appearing grayish; staminate flowers: peduncles slender, sometimes twice as long as the subtending petiole, receptacle tube 5-9 mm. long, sepals narrowly linear, 5-15 mm. long, the apex acute or obtuse, sometimes spathulate, corolla yellow to orange- yellow, green-veined, mostly 5-7 cm. long, petals obtuse or acute, filaments short, anther head about 15 mm. long; pistillate flowers: peduncles short, 0.5-2 (-3) cm. long, perianth similar to that of the staminate flowers but larger; fruit peduncles stout, sulcate, fruits globose or ovoid-globose, glabrous, to about 7 cm. long and 6 cm. broad, deep green or yellowish, longitudinally striped with lighter color, the shell thin but hard, the flesh white, stringy, bitter; seeds cuneate-obovate, 7-10 mm. long, pale olive-white, with a thickened, wrinkled margin.
According to Standley, the plant treated here is a wild one, of which no use is made locally. The fruits are too bitter to be eaten. During the dry season the ripe fruits may be seen in large numbers, hanging on low shrubs, particularly on the Pacific plains.
FIG. 12. Cucurbita hindelliana. A, habit (portion of vine with a staminate flower), x '/2; B, pistillate flower, top view, x Vr, C, fruit and seed, X Vz.
332
DIETERLE: FLORA OF GUATEMALA 333
CYCLANTHERA Schrader1
Climbing vines, annual or perhaps sometimes perennial, monoecious, the stems more or less branched and angulate-sulcate, glabrous to pubescent; leaves on angulate-sulcate petioles, the blades lanceolate to orbicular, almost entire or 3-9- lobate or pedately 3-5-foliolate, with or without glands at the base; tendrils unequally parted, usually bifid, glabrous to pubescent; flowers small, pedicellate; staminate inflorescences: racemose to narrowly paniculate, receptacle saucer-shaped or cup-shaped, the sepals 5, or sometimes obsolete or obscure, corolla rotate, deeply 5- parted, the triangular segments acute, stamens united into a central column, the anthers connate into a head, the thecae united into a horizontal ring; pistillate flowers: usually solitary, in the same axil with the male inflorescence, perianth similar to that of the male flower but usually larger, ovary obliquely ovoid, rostrate, said to be 2-multiloculate, ovules few-many, style short, stigma subglobose; fruit obliquely ovoid, somewhat fleshy, echinate or rarely smooth, explosively dehiscent or sometimes the cultivars indehiscent, few-many-seeded; seeds broadly ovoid to rectangular, sometimes tear-shaped, sometimes turtle-shaped, rather compressed, the margin often appendaged.
The genus contains about 30 species distributed from the southwestern United States through Mexico, Central America, and much of South America.
Cyclanthera pedata (L.) Schrad. is perhaps native of or selected for cultivation from Ecuador or Peru. It is a minor food plant and cultivated, or possibly escaped, in Guatemala and other Central American countries, and in the Old World.
Leaves with 3-5 primary leaflets, 3-5-petiolulate.
Primary leaflets 5 C. pedata.
Primary leaflets 3.
Androecial head stalked, petiole long-villous at apex C. langaei.
Androecial head sessile, petiole not long-villous at apex.
Lateral leaflets deeply to completely bisected (rarely trisected); usually
occurring above 2,000 m C. integrifoliola.
Lateral leaflets deeply or completely 3-7-times dissected; usually occurring
below 2,000 m C. multifoliola.
Leaves nearly entire or merely lobed.
Staminate inflorescence longer than petiole, or usually as long as or longer than the leaf (or more than 3.5 cm. long).
Peduncles of fruit less than 2.5 cm. long C. ribiflora.
Peduncles of fruit longer than 2.5 cm C. steyermarkii.
Staminate inflorescence shorter than or equalling petiole (or less than 3.5 cm. long) C. explodens.
'Dr. C. E. Jones, Jr. h?i prepared a thesis, as yet unpublished, of the genus and annotated specimens in many heibaria. Where possible, his annotations have been followed. Description r.nd synonym/ from the thesis have been abstracted and used largely without revision except as necessary to conform with the "Flora of Guatemala" format.
334 FIELDIANA: BOTANY, VOLUME 24
Cyclanthera explodens Naud. Ann. Sci. Nat. ser. IV, 12: 160. 1859.
In thickets, 800-2,000 m.; Chiquimula. Mexico; El Salvador; Costa Rica; Andean South America.
Vines to several m. long, mostly annual; stems glabrous, branched; leaves on petioles to 4 cm. long, the blades ovate to orbicular, subentire to deeply 3-5-lobate, denticulate, more or less scabrous above, glabrous to puberulent beneath, apex acute, base shallowly to deeply cordate, or with an open sinus, 2-10 cm. long and 2-11 cm. broad; tendrils unequally bifid; staminate flowers: inflorescence short, delicate, receptacle 1-1.5 mm. broad, sepals none or obscure, corolla white, minute papillate, 1.5-2.5 mm. broad, the segments 5; pistillate flowers: peduncles glabrous or puberulent, the stigma 1-1.5 mm. broad, style short; fruits greenish, gibbous, 2-4 cm. long, short-beaked, the base rounded to cuneate; seeds grayish brown, turtle-shaped, 7-9 mm. long.
Rare in Guatemala, often cultivated for the edible fruits.
Cyclanthera integrifoliola Cogn. Mem. Cour. Acad. Belg. 28: 65. 1877. C. donnellsmithii Cogn. Bot. Gaz. 20: 290. 1885 (type from El Quiche, Heyde & Lux 3345).
Thickets and old fields, mostly above 2,000 m.; El Quiche; Huehuetenango. Mexico.
Annual vines, the stems somewhat branched, glabrous; leaves on petioles to 5.5 cm. long, these sparsely pilose or glabrate, the blades broadly ovate to orbicular, trifoliolate, sparsely serrulate, glabrous except more or less pubescent on the veins beneath, 2.5-14 cm. long and about as broad, the central leaflet lanceolate or elliptic to oblong, lateral leaflets similar, bi-trisected; staminate flowers: inflorescence racemose to narrowly paniculate, receptacle 1-2 mm. broad, sepals none, corolla more or less papillate, especially within, 2-5 mm. broad, androecial head sessile; pistillate flowers: 1.5-2 mm. broad, stigma sessile; fruits greenish white, gibbous, explosively dehiscent, to 4 cm. long and 2.5 cm. in diameter, apex acuminate, the base rounded to cuneate, covered with setae to 5 mm. long; seeds 8-10 mm. long, turtle -shaped, grayish brown.
Cyclanthera langaei Cogn. Mem. Cour. Acad. Belg. 28: 67.
1877.
Thickets, old fields, and disturbed forests, growing over trees and shrubs, 1,400-2,400 (-3,200) m.; Alta Verapaz; Huehuetenango; Retalhuleu; Sacatepequez; San Marcos; Solola. Mexico; El Sal- vador; Nicaragua; Costa Rica.
Vines, annual or perhaps perennial, often of considerable length; leaves on glabrous petioles, these to 7.5 cm. long and usually bearing 2 tufts of villae at the apex, the blades ovate to orbicular, trifoliolate, 5-21 cm. long and 5-11 cm. broad, glabrous or nearly so, central leaflet lanceolate to elliptic-oblong, lateral leaflets trilobulate; staminate flowers: inflorescences racemose to narrowly paniculate, glabrous, much exceeding the leaves, receptacle 3-4 mm. broad, sepals obscure to 1
FIG. 13. Cyclanthera langaei. A, habit (portion of a flowering stem), x Vfe; B, leaf, X 1; C, pistillate flower, X 1; D, pistillate flower, X 3'/2; E, diagram of androecium, X 5; F, staminate flower, X 3'/2 (note detached stamen above, X 5); G, staminate flower, underside, X l!/2; H, seeds and an immature fruit (typical gibbous shape not yet fully developed), X 11A.
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336 FIELDIANA: BOTANY, VOLUME 24
mm. long and filiform, corolla lobes 2.5-5 mm. long, filament column 0.5-1 mm. long; pistillate flowers: solitary, peduncles to 5 cm. long; fruits pale green, gibbous, to 3.5 cm. long and 2.5 cm. in diameter, beak 7-15 mm. long, echinate with glabrous spines to 7 mm. long; seeds grayish brown, turtle-shaped, 7-8 mm. long.
The fruits are sometimes used as food by the country people, often filled with ground meat.
Cyclanthera multifoliola Cogn. Mem. Cour. Acad. Belg. 28: 66. 1877. C. filifera Cogn. torn. cit. 72 (type from Solola, Bernoulli 1201).
Thickets and forest edges, 1,000-1,500 m.; Jutiapa; Santa Rosa; Solola. Mexico; Costa Rica; Ecuador; Peru; Brazil.
Annual vines with branched, glabrous stems or with nodes more or less pubescent; leaves ovate to orbicular, trifoliolate, 3.5-15 cm. long and 4.5-20 cm. broad, glabrous or the veins beneath pubescent, the central leaflet elliptic to lanceolate, lateral leaflets deeply 3-7 times dissected; staminate inflorescences: many-flowered, much longer than the leaves, receptacle 0.8-2.5 mm. broad, sepals none or obscure, corolla somewhat papillate, greenish white, 2-3.5 mm. broad, androecial head sessile; pistillate flowers with broad, sessile stigma; fruits greenish white, gibbous, to 4 cm. long and 2 cm. in diameter, acuminate, beak 2-8 mm. long, the base rounded or cuneate, densely setose, the setae to 7 mm. long; seeds brown, turtle-shaped, 8-13 mm. long.
Cyclanthera pedata (L.) Schrad. Linnaea 8, Litt.-Bericht: 23. 1833. Momordica pedata L. Sp. PL 2: 1009. 1753. (Fig. 39).
In Guatemala perhaps only in cultivation or as an escape, about 1,500 m.; Baja Verapaz; Guatemala. Mexico (?); Honduras; Costa Rica; Andean South America.
Annual vines to several m. long, the stems branched, glabrous; leaves on glabrous petioles 1-8 cm. long, blades broadly ovate to orbicular, pedately lobate, margins serrulate to deeply dentate, essentially glabrous, 2.5-20 cm. long and about as broad, lateral leaflets pedately lobed; staminate flowers: in panicles with few-several (rarely many) flowers confined to upper half or less of peduncle, receptacle to 3.5 mm. wide, sepals obscure or to 2 mm. long, corolla obscurely papillate, whitish, 3-6 mm. across, filament column to 0.75 mm. long, androecial head to 1.5 mm. wide; pistillate flowers: subsessile, stigmas sessile, to 2.5 mm. broad, with a depressed center; fruiting peduncles 0.5-2 cm. long; fruits greenish white, striate, lacrimiform and somewhat gibbous, not explosively dehiscent, to 15 cm. long and 6 cm. in diameter, setae none or few, confined to abaxial part of fruit; seeds black, more or less turtle-shaped, 12-16 mm. long.
Formerly common in Guatemalan markets but in recent years less so; the "shell" was stuffed with ground or chopped meat and cooked to produce a quite acceptable dish. The fruits are less setose than in other species.
DIETERLE: FLORA OF GUATEMALA 337
Cyclanthera ribiflora (Schlecht.) Cogn. Mem. Cour. Acad. Belg. 28: 63. 1877. Elaterium ribiflorum Schlecht. Linnaea 7: 388. 1832.
Thickets or old fields, about 1,500 m.; Huehuetenango; Sacatepequez. Mexico.
Annual vines with glabrous, branched stems, nodes sometimes pubescent; leaves on petioles to 4 cm. long, blades ovate to orbicular, to 8 cm. long and 10 cm. broad, shallowly to deeply 3-5-lobate, margins denticulate, glabrous or often puberulent along the veins, central lobe ovate to triangular, the lateral ones more or less angulate; staminate inflorescences narrowly paniculate, few-many-flowered nearly to base of peduncle, to about 12 cm. long, receptacle 1-1.5 mm. broad, sepals none or obscure, corolla whitish, papillate, 2-3.5 mm. broad, androecial head sessile, to 0.75 mm. broad; peduncles of pistillate flowers glabrous, to 4 mm. long, stigma 1-2 mm. broad, nearly sessile; fruiting peduncles 5-15 mm. long, the fruit greenish, gibbous, explosively dehiscent, to 2.5 cm. long and 1.7 cm. in diameter, setae entirely covering fruit, these glabrous, to 6 mm. long; seeds dark brown, turtle-shaped, 4.5-7 mm. long.
Cyclanthera steyermarkii Standl. Field Mus. Bot. 23: 93. 1944.
In forest or thickets, about 2,300 m.; Chimaltenango; Quezal- tenango (type from Volcan de Santa Maria, Steyermark 34047)', Solola; Zacapa. Mexico.
Annual vines with branched, glabrous stems, the nodes sometimes puberulent; leaves on petioles 0.5-8 cm. long, the blades broadly 3-5-angulate-lobate, margins denticulate or dentate, glabrous or puberulent on veins, the central lobe triangular, .acuminate; staminate inflorescences racemes or reduced panicles 2-9 cm. long, few- flowered, receptacle about 1.5 mm. broad, sepals none or obscure, corolla white, papillate, 2-2.5 mm. wide, androecial head sessile, about 0.8 mm. broad; peduncles of pistillate flowers 0.7-2.5 cm. long, nearly sessile, stigmas 1.5 mm. broad; fruiting peduncles 2.5-4 cm. long, the fruits greenish, gibbous, mostly 1.5-2 cm. long and about 1.5 cm. in diameter, often crowned by the persistent sepals, the body covered with setae 1-2.5 mm. long; seeds light brown, turtle-shaped, about 7 mm. long.
DOYEREA Grosourdy
Reference: J. Hutchinson, The Genera of Flowering Plants, II: 386-387. 1967.
Perennial, woody, scandent, probably dioecious vines, the base trunklike, short and thick, the stems becoming obscurely fractiflex in age and conspicuously lenticular; leaves petiolate, the blades rounded-cordate, palmate or rather bluntly or obscurely trilobate, broadly cordate at the base, 3-nerved from the base; tendrils simple, woody; flowers disposed in short, axillary racemes on thick, fleshy peduncles, those of the pistillate flowers shortened to a tight cluster; staminate flowers: 6-20 (-40) in each raceme, sepals 5, valvate, acute, corolla yellowish green, 5-lobate, the lobes shortly acuminate, stamens 3, the filaments inserted near the rim of the receptacle, two anthers dithecous, one monothecous; pistillate flowers: fewer, short-
338 FIELDIANA: BOTANY, VOLUME 24
pedicellate or subsessile, bracteolate, perianth similar to that of the staminate flowers, staminodes 3, style thick, simple, stigmas said to be 4, ovary bilocular with 4- 6 ovules on each parietal placenta; fruit ellipsoid to oblong, fleshy, shortly rostrate, 1- 2 cm. long, first green with broken white stripes, later red, mottled or striped with greenish white.
Only one species is known.
Doyerea emetocathartica Grosourdy, Fl. Med. Antill. 2: 338. 1864. Corallocarpus emetocatharticus Cogn. Bull. Soc. Bot. Belg. 30, pt. 1: 279. 1891. C. millspaughii Cogn. Field Mus. Bot. 1: 322, t. 20. 1896. C. guatemalensis Standl. & Steyerm. op. cit. 23: 93. 1944 (type collected near Zacapa, Standley 74623).
Damp or dry thickets, rocky slopes, brushy hillsides, sea level to 500 m.; Chiquimula; Retalhuleu; Zacapa. Mexico; British Honduras; Honduras; Nicaragua. West Indies; Venezuela; Colombia.
Woody climbers with stems first fleshy, becoming woody, said to attain more than 6 m. in length, puberulent when young, glabrous or glabrate in age; leaves on petioles 1.5-4 cm. long, the blades rounded-cordate, somewhat angular, or more commonly more or less trilobate, mostly 4-9 cm. long, the apex sometimes abruptly acuminate when young but usually obtuse and minutely apiculate in age, commonly minutely scabridulous above, sometimes short-pilose, densely and softly pilosulous beneath, the margins dentate to remotely denticulate or almost entire, the basal sinus deep, rectangular or more broadly arched; flowers subsessile in short racemose clusters; staminate flowers: peduncle 3-4 mm. long, densely hairy, the short pedicels glabrous, receptacle less than 2 mm. long, sepals about 1 mm. long, corolla lobes greenish yellow, stamens 3, inserted near the rim of the receptacle, thecae nearly 2 mm. long, curved, the connective not appendaged; pistillate flowers: perianth like that of the staminate flowers, the ovary said to be bilocular; fruits ellipsoidal or oblong, 1.3-2 cm. long, short- rostrate, rather thin- walled, appearing rugose when dry, first green with broken whitish lines, becoming red, mottled or striped with greenish white when mature; seeds pear-shaped, 3-4 mm. long, with a pale, winglike margin near the apex.
ECHINOPEPON Naudin
References: A.Cogniaux, Echinocystis, sect. Echinopepon (Cu- curbitaceae) in DC. Monogr. Phan. 3: 798-815. 1881; K. M. Stocking, Some considerations of the genera Echinocystis and Echinopepon in the United States and northern Mexico, Madrono 13: 84-100. 1955.
Annual, herbaceous, monoecious vines, the stems usually slender, scandent or trailing, glabrous or variously pubescent, striate. drying angular-sulcate; leaves petiolate, the blades thin, usually angulate-cordate, commonly more or less 3-5(-7)- lobate, the basal sinus various, margins almost entire to denticulate; tendrils bifid or trifid: staminate flowers: pedicellate, disposed in racemes or panicles, perianth 5-
FIG. 14. Doyerea emetocathartica. A, habit (thickened stem at ground level, from a photograph); B, habit (section of a fruiting vine), X Vz; C, staminate inflorescence (from a photograph); D, fruit (from a photograph); E, opened fruit, seeds, X IVr, A, C, and D from photographs by Erich Halbinger.
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340 FIELDIANA: BOTANY, VOLUME 24
merous, receptacle cup-shaped to patelliform, sepals small or minute, often subulate, corolla campanulate or spreading, white or greenish white or cream, stamens 5. filaments fused into a column, anther connectives almost or entirely fused to form globose, hemispherical, or cuplike heads, thecae straight, curved, or replicate; pistillate flowers: mostly solitary, sessile or pedunculate, perianth similar to that of staminate flowers but usually larger, ovary bilocular with 2-5 ascending (rarely horizontal) ovules in each locule; fruits ovoid or ellipsoid, operculate, glabrous or hairy, conspicuously echinate, rostrate; seeds quadrangular or angular-ovate, compressed, rugose or verrucose.
Preliminary studies of Echinopepon pollen, using scanning electron microscopy, indicate that very different types of exine morphology are to be found among the species included, each type possibly associated with a particular shape of anther thecae; non- colpate grains with tightly replicate thecae, for example, multicol- pate grains with U-shaped thecae, etc.
About a dozen species, all American, with two in Guatemala. We have treated three species here, since E. torquatus (DC.) Rose has been collected in nearby Chiapas, Mexico and may be expected in Guatemala.
Petioles glabrous save for a tuft of hairs at the apex; staminate flowers very small. 1.5-2.5 mm. across; thecae U-shaped; fruits conspicuously asymmetrical at base
and apex; spines glabrous E. torquatus.
Petioles pubescent to long-villous; staminate flowers 6-10 mm. across; thecae replicate, hooked, arched, or straight, but never U-shaped; fruits vertically symmetrical; spines hairy. Staminate inflorescences racemose; receptacle cup-shaped; thecae tightly replicate;
mature fruits (excluding spines) 1.5-2.5 cm. wide E. horridus.
Staminate inflorescences paniculate; receptacle saucer-shaped; thecae hooked, arched, or straight; mature fruits (excluding spines) 0.8-1.5 cm. wide.
E. paniculatus.
Echinopepon horridus Naud. Ann. Sci. Nat. Bot. ser. 5, 6: 19. 1866. Echinocystis lanata Cogn. Diagn. Cucurb. 2: 92. 1877; Mem. Acad. Sci. Belg. 8, 28: 92. 1878. Micrampelis lanata O. Ktze. Rev. Gen. 1: 257. 1891. Echinopepon lanatus Rose, Contr. U. S. Nat. Herb. 5: 117. 1897, in syn. Echinopepon jaliscanus Rose, loc. cit. Quimiche (Chimaltenango); quix (Guatemala, fide Aguilar); quisquil de culebra (Zacapa).
Damp or wet thickets, sometimes along roadsides, 900-1,900 m.; Alta Verapaz; Chimaltenango; Chiquimula; Guatemala; Jutiapa; El Quiche; Zacapa. Mexico; British Honduras and Honduras south to Costa Rica; possibly South America.
FlG. 15. Echinopepon horridus. A, habit, X Vfc; B, pistillate flower, X 4; C, spine from ovary, X 8; D, nearly mature fruit, X 1; E, seed, X 1 above, X 4 below; F, staminate flower, X 3'/>; G, underside of staminate flower, X 21A. See also Figure 40.
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342 FIELDIANA: BOTANY, VOLUME 24
Small or large vines, the stems sparsely or densely villous with multiseptate hairs; leaves on long- villous petioles mostly 2-10 cm. long, the blades thin, commonly broader than long, angulate or shallowly 3-5(-7)-lobate, rather deeply cordate at the base with a broad, open sinus, acute or obtuse, mostly 7-14 cm. long, the margins almost entire or rather remotely denticulate, glabrate or sparsely pilose above, sometimes scabridulous, usually softly and sparsely pilose beneath; tendrils trifid or bifid; staminate flowers: in racemose inflorescences, peduncles 3-8 cm. long, pubescent, pedicels mostly 5-12 mm. long, the 15-25 flowers commonly 8-14 mm. across, receptacle cup-shaped, conspicuously green-veined, sepals 1-3 mm. long, corolla white, more or less glandular within, the petals ovate-triangular, acute or subobtuse, filament column glabrous, thecae tightly replicate; pistillate flowers: solitary or geminate, sessile or subsessile, ovary ovoid, densely lanate, style 2-3 mm. long, stigma globose or subglobose; fruit oblong-ovoid, commonly 3-4 (-5) cm. long (excluding beak), 1.5-2.5 cm. broad (excluding spines), densely echinate, rostrate, the beak 0.5-2 cm. long, the spines unequal, 4-17 mm. long, as much as 3 mm. broad at the base, pubescent; seeds oblong-elliptical, 7-8 mm. long, 4-6 mm. wide, about 2.5 mm. thick, verrucose.
Echinopepon paniculatus (Cogn.) Dieterle, comb. nov. Ech- inocystis paniculata Cogn. Diagn. Cucurb. 2: 90. 1877.
Damp thickets, savannas, brushy or rocky slopes, forest clearings, cornfields, near sea level to 1,700 m.; Chiquimula; Huehuetenango; Jalapa; Peten. Mexico; British Honduras.
Annual, herbaceous, branching, scandent vines, the stems puberulent or pubescent and more or less villous, in age glabrate or sometimes sparsely villous, deeply sulcate, the nodes often long-villous; leaves on puberulent and more or less long-villous petioles 2-8 cm. long, the blades broadly triangular-ovate, shallowly 3-5- lobate or merely angulate, the apex acuminate, the base subauriculate, the basal sinus deeply and broadly arched, bordered in part by the lowermost lateral veins, the margins subentire to shallowly denticulate, both surfaces more or less short-pilose or the lower surface glabrate except for costa and veins; tendrils bifid or trifid, sulcate, pubescent; staminate flowers: in paniculate inflorescences as much as 20 cm. long and 5-9 cm. broad, peduncles and pedicels more or less pubescent or viscid-pubescent, the slender pedicels mostly 6-15 mm. long, receptacle and corolla white or greenish cream, glandular, the receptacle patelliform, with a conspicuous spot of dark pigment at the center, sepals minute, subulate, corolla patent, 9-11 mm. across, the lobes triangular to ovate-triangular, punctate-glandular to stipitate-glandular, staminal column short, anthers 5, thecae straight, hooked, or arched, the connectives joined and appearing hemispherical or cuplike; pistillate flowers: solitary, subsessile or short-pedunculate, the perianth like that of the staminate flowers but larger, commonly 10-12 mm. across, ovary oblong, densely tomentose, style short, stigma subglobose; fruiting peduncles 1-2 (-3) cm. long, the fruit ellipsoidal, tapering toward the base, the body densely echinate, about 3-4 cm. long, 0.8-1.5 cm. wide, the beak 1-2 cm. long, the spines unequal, hairy, 3-16 mm. long; seeds angulate-obovate, (4-) 5-8 mm. long, verrucose.
There is some question as to whether E. paniculata may include two specific or subspecific entities or whether it is simply a
DIETERLE: FLORA OF GUATEMALA 343
highly variable species. Included are plants with straight thecae and plants with hooked thecae. Those with straight thecae tend to have conspicuously stipitate glands on the corolla lobes while those with hooked thecae tend to have punctate glands on the lobes. I feel that a decision should not be made until additional material can be examined and further studies made.
Echinopepon torquatus (DC.) Rose, Contr. U. S. Nat. Herb. 5: 118. 1897. Elaterium torquatum DC. Prodr. 3: 310. 1828. Echinocystis torquata Cogn. Mem. Cour. Acad. Belg. 8, 28: 99. 1878. (Fig. 41).
Not reported from Guatemala but included here, as several collections have been made in nearby Chiapas, Mexico (1,500- 2,500 m.).
Annual vines, the stems glabrous; leaves on petioles mostly 3-14 cm. long, these glabrous save for a tuft of long, multiseptate hairs at the apex, the blades broadly angular-ovate in outline, cordate at the base, shallowly or fairly deeply 5-lobate, mostly 4-8 cm. long, acute or acuminate, more or less scabrous or scabridulous on both surfaces or essentially glabrous, margins dentate, denticulate, or subentire, basal sinus broadly rounded or rarely almost V-shaped; tendrils trifid; staminate flowers: in racemes 3-10 cm. long, the pedicels 5-9 mm. long, the perianth very small, 1.5-2.5 mm. across, receptacle cup-shaped, sepals minute, corolla white, petals triangular, filament column glabrous, thecae U-shaped; pistillate flowers: solitary or geminate, subsessile or short-pedunculate, perianth similar to that of the staminate flowers, style short, stigma globose; fruiting peduncles 0.3-2 cm. long, the fruits oblong, about 2 cm. long, 0.5 cm. wide, more or less curved below the beak, usually conspicuously so, the body echinate with unequal, broad-based, glabrous spines, the longest to about 9 mm.; mature seeds not seen, said to be ovate, almost smooth.
FEUILLEA Linnaeus
References: A. Cogniaux, Pflanzenreich IV. 275, I: 4-9. 1916. H.E. Baillon, The Natural History of Plants VIII: 375-379. 1886. C. Jeffrey, Kew Bulletin 16: 200. 1962.
Large vines, herbaceous or suffrutescent, dioecious; leaves petiolate, the blades mostly membranaceous or subcoriaceous; tendrils distally bifid, coiling both above and below bifurcation; staminate inflorescences paniculate, the flowers small, greenish or yellow; staminate flowers: receptacle short, campanulate or cupular, sepals 5, imbricate in bud but not completely enclosing the petals, spreading in anthesis, petals 5, unguiculate, imbricate in bud, spreading in anthesis, the limb ovate or oblong, each often with a median, ventral, uncinate appendage, stamens 5, free, monothecous, inserted in the center of the receptacle, thecae short, curved, anthers longitudinally dehiscent, connective thick, not produced at the apex, pistillodium none; details of pistillate flower structure, as reported in the literature: perianth like that of staminate flower, ovary oblong, imperfectly trilocular, free at apex, ovules 6 in each locule or fewer by abortion, pendulous from the margins of the carpels in 2 vertical series, styles 3, distinct, the stigmas reniform, bilobate, staminodia 5 or none;
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FIELDIANA: BOTANY, VOLUME 24
FIG. 16. Feuillea cordifolia. A, portion of stem with a leaf and an immature staminate inflorescence, X Vz\ B, seed, two views, X %; C, staminate flower, X \Vz; D, staminate flower, underside, X l'/2.
fruit large, gourdlike, apically operculate, girdled above the middle by the scar of the limb of the receptacle and marked at the apex by a raised triradiate line; seeds many, large, orbicular, compressed, the testa thick.
About 10 species in tropical America. Only one cultivated species is treated here.
Feuillea cordifolia L. Sp. PL: 1013. 1753. Chichimora. Sometimes planted in Guatemalan gardens.
Stems stout, high-climbing, glabrous; leaves subcoriaceous, nearly glabrous, usually cordate in outline, 8-16 cm. long, margins entire but with 4-8 conspicuous marginal glands terminating main lateral veins; tendrils very long, robust, bifid at apex, glabrous or nearly so; staminate inflorescences paniculate, branchlets of staminate panicles usually subtended by a cordiform bracteole, the panicles 20-60 cm. long, densely short-villous, staminate flowers: numerous but reaching anthesis only a few at a time, the pedicels 2-6 mm. long, slender, puberulent, receptacle shallowly cupular, glabrous within, puberulent without, sepals ovate, 2 mm. long, margins ciliate, petals yellowish, ovate-orbicular, about 4 mm. long, each bearing a median ventral uncinate appendage, stamens 5, free, monothecous, radiate from center of receptacle, filaments linear-clavate, recurved; fruit globose, green, 10-12 cm. in diameter, seeds orbicular, 4.5 cm. broad or larger, 1.2 cm. or more thick, brown, complanate, margined, smooth.
DIETERLE: FLORA OF GUATEMALA 345
According to Standley, this species may be native in the West Indies and tropical South America but it is cultivated in some or all of the Central American countries. The seeds are to be found in small numbers in most Central American markets, and particularly in those of Guatemala.
GURANIA Cogniaux
Scandent dioecious herbs; leaves long-petiolate, membranaceous to subcoriac- eous; tendrils undivided; staminate flowers: in long-pedunculate, spicate, racemose, or subcapitate inflorescences, receptacle-tube subglobose, ventricose, or cylindric, sepals elongate, narrow, red, petals shorter than the sepals, separate to the base, lingulate, yellow, stamens 2, free, dithecous (or, reported by some authors as one dithecous and one tritheous), sessile or nearly so, dorsifixed near the middle of the receptacle tube, rudimentary pistil none; pistillate flowers: solitary, or the inflorescence fasciculate, or capitate, perianth like that of the staminate flower, ovary slenderly cylindric (said in literature to have 2 placentas), style columnar, bifid, stigmas 2, rudimentary stamens 2, sometimes 1 or none; fruit long-cylindric; seeds ovate, horizontal, compressed, numerous.
Many species have been described from tropical South America, several from Central America. Collections from Guatemala avail- able to me provide scant floral material for dissection. The similarity in their general aspect, however, combined with the paucity of fertile material, makes it difficult to dispel the suspicion that no more than one species is represented. More field work is required.
Gurania makoyana (Lem.) Cogn. Diagn. Cucurb. 17. 1876. Anguria makoyana Lemaire, Fl. Serres 3, t. 222. 1847 (originally described from plants grown in Belgium from seeds said to have been sent from Guatemala). G. donnell-smithii Cogn. in Donn.-Sm. Bot. Gaz. 16: 10. 1891 (type from Alta Verapaz, J. D. Smith 1511).
Wet forest, thickets, swamps, sea level to 900 (1,500) m.; Alta Verapaz; Chiquimula; Izabal; Peten. British Honduras; Honduras.
Stout vines, probably perennial, sometimes climbing high in trees, the stems short-villous; tendrils undivided, sturdy, short-villous on outer surface of coils; leaves on stout, hirsute-tomentose petioles 5-12 cm. long, the blades membranaceous to subcoriaceous, broadly suborbicular in outline, 15-35 cm. long, 18-40 cm. wide, commonly trilobate to about the middle or deeper, the lobes broadly rounded, somewhat constricted below, abruptly acuminate at tips, basal sinus subrectangular, 2-4 cm. broad and deep, partly bordered at the top by the 2 main lateral veins of the blade, paler green below, more or less villous-pilose on both surfaces, especially on the principal veins, margins minutely spinulose-denticulate; staminate flowers: in densely capitate-spicate inflorescences, the flowers sessile or subsessile, the stout common peduncle (12-) 18-32 (-34) cm. long, densely to moderately villous, receptacle
FIG. 17. Gurania makoyana. A, section of vine with a staminate inflorescence, X Vfe; B and C, leaf forms, x 1/10; D, pistillate flower dissected to expose style and staminodia, x 2; E, staminate flower, x 4; F, bud of staminate flower, x 2W, G, fruit (with opening to show seeds within), X %; H, two views of seed, X IVz.
346
DIETERLE: FLORA OF GUATEMALA 347
subglobose to globose-ellipsoid, often densely lanate, 8-10 mm. long, sepals erect, linear, acuminate, densely or moderately tomentose or pubescent, 10-24 mm. long, almost 1 mm. broad at base, petals erect, connivent, oblong-ellipsoid, obtuse, papillose, 7-8 mm. long, anthers oblong, thecae replicate at base, the connective narrow, the appendage about 1 mm. long, flat, more or less ovate, glabrous or papillose; pistillate flowers: borne in few-several-flowered clusters on nodes of branches with repressed leaves and tendrils, pedicels rather slender, short-puberulent, perianth like that of the staminate flower, sepals 8-14 mm. long, ovary narrowly cylindrical, thinly pilose, about 2 cm. long, style fleshy, 0.5-1 cm. long, about 0.3 cm. thick, deeply bifid, the segments connivent, placentae parietal; fruit cylindric, somewhat narrowed at each end, green, many-seeded; seeds whitish, 8x5x2 mm., oval in outline, compressed, with a slight margin.
HANBURIA Seemann
Reference: B. Seemann, Bonplandia 6: 293. 1858; 10: 189, t. 12. 1862.
Coarse, perennial, monoecious vines, almost glabrous; leaves petiolate, the blades broadly rounded in outline, simple or lobed; staminate flowers; in racemose inflorescences, corollas white, deeply 5-lobed, stamens united, the filaments in a central column, the anthers in a connate head depressed at the apex, thecae vertically replicate and linearly dehiscent, pollen grains large, globose, 5-6-colpate, smooth, finely pitted; pistillate flowers: solitary, perianth similar to that of staminate flower, ovary obliquely ovoid or subglobose, more or less echinate, ovules erect or ascending, style columnar, stigma large, capitate; fruit large, to about 10 cm. long, obliquely ovoid, acute, carnose, rupturing explosively to expose a pendulous placental arm which hurls the seeds; seeds few, large, about 2 cm. in diameter (in ours), orbicular, compressed.
Two species are known, in Mexico and Central America, one of them in Guatemala.
Hanburia parviflora Donn.-Sm. Bot. Gaz. 13: 299. 1886; 15: 27. 1890.
Wet forest, 250-1,100 m.; Alta Verapaz. Mexico (Chiapas).
Extensive vines in clearings and second growth, glabrous, the stems many m. long, sulcate; leaf blades broadly orbicular or ovate in outline, 8-16 cm. long, deeply trilobate, the lobes oblong-lanceolate, acuminate, the margins entire or distantly and shallowly dentate, the base subtruncate and then obtusely cuneate to the petiole and bearing small, scattered, disciform glands on the underside; petioles 1-3 cm. long, much shorter than the leaf blades; tendrils bifid, on peduncles about one and one- half times as long as the adjoining petioles; staminate flowers: in short racemes about one-half to twice as long as the petioles, the racemes few-flowered, nodding, the pedicels slender, spreading, receptacle elongate-tubular, 9-15 mm. long, expanded at base and apex, sepals small, dentiform, remote, corolla lobes ovate-triangular, reflexed, about half as long as the receptacle tube, filament column glabrous, androecial head obpyriform, exserted outside the corolla; pistillate flowers: solitary
FlG. 18. Hanburia parviflora. A, habit X Vz; B, base of leaf blade, underside, X 1'4 (one gland, much enlarged, at left); C, section of lower stem, x 2; D, staminate flowers in bud, x IVfe; E, staminate flower opened to expose filament column and androecial head, X 5; F, nearly mature fruit, X Vz; G, fruit dissected to show placenta and seed, X Vfc; H, two views of seed, X '/2.
348
DIETERLE: FLORA OF GUATEMALA 349
on peduncles about 10 cm. long, perianth similar to that of staminate flowers; fruits about 8-10 cm. long, sparsely echinate on the proximal portion, the echinae soft, fleshy, seeds few (2-4), nearly orbicular, about 23 X 18 X 3 mm., dull brown, smooth, the margin irregular.
LAGENARIA Seringe
Herbaceous, annual vines, densely and softly pubescent, monoecious; leaves simple, long-petiolate, the blades rounded-cordate, the petiole with a pair of apical, lateral glands; tendrils proximally bifid; flowers solitary, rather large, white, night- blooming; staminate flowers: receptacle funnelform to campanulate, sepals 5, relatively small, remote, petals 5, free, entire, stamens 3, two dithecous, the third monothecous, the thecae triplicate and usually also much contorted, inserted low on the receptacle tube, the filaments free, the anthers free or slightly coherent, anther connectives not produced beyond the thecae, pollen globose, tricolporate, rudiment- ary pistil none or represented by a gland; pistillate flowers: perianth similar to that of the staminate flowers but somewhat smaller and with a very short receptacle, ovules numerous, horizontal, style short, thick, the stigma trilobate, thick, fleshy, staminodes 3, small or minute; fruit large, terete, indehiscent, hard-shelled, fleshy, the pulp spongy; seeds numerous, obovate-oblong. slightly flattened, smooth, more or less distinctly marginate, truncate and often bicornute at one end, pale grey or striped.
The genus consists of a single species.
Lagenaria siceraria (Molina) Standl. Field Mus. Bot. 3: 435. 1930. Cucurbita lagenaria L. Sp. PI. 1010. 1753. C. siceraria Molina, Sagg. Chil. 133. 1782. C. leucantha Duch. in Lam. Encycl. 2: 150. 1786. L. vulgaris Ser. Mem. Soc. Geneve 3, pt. 1: 25, t. 2. 1825. L. leucantha Rusby, Mem. Torrey Bot. Club 6: 43. 1896. Tecomate (fruit), tol, tecomate de grillo, chu, bux, lee (Peten); chuj, suj (Quecchi, Alta Verapaz); suj (Pocomam); suy (Cacchiquel); cala- baza, tecumat (Pipil of Salama).
Probably native in tropical Asia and Africa, but introduced into America long before the advent of ancient man in the Americas. Commonly cultivated and said to be frequently naturalized in tropical America; grown in Guatemala at low and middle elevations, 250-1,900 m., and sometimes at higher altitudes, naturalized in many places in thickets or waste ground; Alta Verapaz; Baja Verapaz; Chimaltenango; Chiquimula; Guatemala; Jalapa; Jutiapa; Peten; Retalhuleu; Sacatepequez; Santa Rosa; Zacapa; probably in all departments.
Vigorous vines, often large with coarse stems, prostrate or scandent, softly pubescent, with grey-green foliage and handsome white flowers that open at evening; leaves on petioles 3-9 (-16) cm. long, the blades broadly ovate or reniform to cordate- triangular, mostly 6-15 cm. long, 7-22 cm. wide, often angulate or slightly trilobate, with a broad, open sinus, apiculate, the margins sinuate-dentate; staminate flowers:
FIG. 19. Lagenaria siceraria. A, habit, X Vz; B, staminate flower, x 1 (inset of anthers, greatly enlarged); C, leaf bases with petiolar glands, X 2 (greatly enlarged gland in inset); D, pistillate flowers, x Vfc; E, stigmas, X 8; F and G, two fruits, x 1/20; H, two views of seed, x 1.
350
DIETERLE: FLORA OF GUATEMALA 351
penduncle longer than the associated petiole, receptacle 1-1.5 cm. long, pubescent, the sepals triangular to linear, remote, much shorter than the receptacle, 3-4 mm. long, corolla spreading, the petals broadly obovate, 3-4 cm. long, more or less emarginate and apiculate at the apex, strongly green-veined on the dorsal surface, pubescent on the central portion of the corolla, the margins often ruffled, anthers included, the filaments glabrous; pistillate flowers: peduncle equalling or shorter than the subtending petiole, ovary ovoid to long-cylindric, densely villous, receptacle less than 1 cm. long, sepals remote, petals 2-2.5 cm. long, staminodia 3; fruit large, smooth, terete, very variable in size and shape, at first light green, when ripe whitish or yellowish, glabrous; seeds obovate-oblong to almost rectangular, about 15 mm. long, slightly flattened, more or less bicornute at one end, and each face with two raised, submarginal stripes.
Most of the plants of Guatemala have bottle-shaped fruits, but those with a long, slender neck and an enlarged, globose apex, from which are made the familiar gourd dippers of some parts of the United States, are also frequent. The bottle-shaped fruits, with their hard, durable shells, are of great importance in Guatemala, being used like bottles, and commonly as canteens for carrying water.
LUFFA Adanson
Herbaceous vines, climbing or trailing, monoecious; leaves long-petiolate, simple, the blades 5-7-lobate; tendrils proximally 2-5-parted; staminate flowers: in racemes, the pedicels bracteate, receptacle short, campanulate, sepals 5, valvate in bud, petals 5, free, stamens 5, all monothecous, the stamens separate or variously paired to appear as 4 or 3, the filaments free, inserted low on the receptacle, anthers free or sometimes partly coherent, thecae flexuous; pistillate flowers: solitary or single in axils with the staminate flowers, staminodia 5, ovary trilocular, style one, stigma 3- parted, ovules numerous, horizontal; fruits medium-sized to large, dry, fibrous, dehiscent by an apical operculum; seeds 9-12 mm. long (in ours), broadly oblong- ovate in outline, complanate, smooth but bearing a pair of opposing ridges near the hilar edge of each face.
Perhaps six to eight species, in the tropics of both hemispheres, only two in Central America.
Fruits smooth, terete; petals 3-4 cm. long, bright yellow; lateral leaf-lobes acute.
L. cylindrica.
Fruits tuberculate-spiny, ribbed; petals about 1 cm. long, pale yellow; lateral leaf- lobes obtuse L. operculata.
Luffa cylindrica (L.) Roem., Fam. Reg. Veg. 2: 63. 1846. Momordica cylindrica L. Sp. PL 1009. 1753. Pashte, estropajo.
Wet thickets or open forest, usually less than 900 m.; Chiquimula; Izabal; Jutiapa; Peten; Retalhuleu; Santa Rosa; Zacapa. Mexico (Jalisco, San Luis Potosi, Puebla, Chiapas); British
FIG. 20. Luffa cylindrica. A, habit (a leaf and a young staminate inflorescence), X !/2; B, pistillate flower, opened to expose stigmas and staminodia, x \Vr, C, immature pistillate flower, X % (greatly enlarged gland from sepal at left); D, immature fruit, x '/2; E, two seeds, x Vz, the lower one with alate margin lost (detail of an undamaged margin above); F, center of staminate flower, x 2; G, staminate flowers, X !/2.
352
DIETERLE: FLORA OF GUATEMALA 353
Honduras; Brazil; Colombia; Bolivia. Often cultivated in the American tropics and in many regions thoroughly naturalized.
Rampant, herbaceous, annual vines, often several m. long and profusely branched, finely hairy to nearly glabrous; stems sturdy, ribbed; leaves on striate petioles 3-9 cm. long, the blades broadly ovate in outline, mostly 5-10 cm. long, more or less angulate or lobed, dark green above, lighter below, scabrous or scabridulous on both surfaces, especially the upper, the central lobe longer and broader, often long- acuminate, margins coarsely denticulate to nearly entire, basal sinus broadly arched to rectangular, closely but not actually bordered by the lowermost, main, lateral veins, principal veins 5-7, prominent, arising nearly together at the apex of the petiole; tendrils 3-5-parted, on a sturdy peduncle; flowers rather large and showy, bright yellow, the pistillate flowers co-axial with the staminate but reaching anthesis first, a glanduliferous, triangular bract usually present in leaf axils bearing inflorescences; staminate flowers: borne on upper third or less of sturdy racemes 10- 30 cm. long, 10-30-flowered, the pedicels about 3 mm. long, sturdy, tomentose in youth, persistent and recurved after anthesis, each with a bract at or near its base, these glabrous, triangular, rather fleshy, and each bearing 1-3 disciform glands on its abaxial surface, receptacle shallowly and broadly campanulate, sepals triangular to oblong-triangular, sparsely to densely appressed short-pubescent, valvate in bud, median nerves prominent, petals 5, free, broadly obovate in outline, 3-4 cm. long, thin, more or less villous along the numerous principal veins, stamens 5 or fewer, free, partly exserted, filaments distinct, inserted near the center of the receptacle, hairy at the base, thecae 5, longitudinally triplicate and also much contorted, borne marginally and dorsally on broad, lobed connectives, pollen tricolporate, reticulate, relatively large (more than 100 jii in diameter); pistillate flowers: perianth like that of the staminate flowers but the sepals somewhat longer and narrower, and beset with dark, disciform glands on the outer surface, peduncle sturdy, about 1 cm. long (2-10 cm. long in fruit), ovary narrowly cylindric, tomentose and also marked with numerous dark, linear, glandular areas, sepals and style remaining on its apex long after anthesis, ovules numerous, horizontal, style sturdy, glabrous, stigmas 3, the lobes broad, recurved; fruits large, up to 40 cm. or more long, cylindric, rounded at base and apex, not rostrate, longitudinally striped with 10 narrow lines, dry and fibrous when mature, opening by an apical operculum; seeds numerous, broadly oblong-ovate in outline, strongly compressed, about 12 mm. long, 4-5 mm. broad, nearly white to dark brown, dull, smooth, the margins narrowly alate, and each face ornamented with a pair of almond-shaped ridges near the hilar end.
According to Standley, the interior of the fruit is ". . . composed of a dense, somewhat elastic mass of reticulated fibres, somewhat resembling a marine sponge, and may be used in much the same manner. The name Spanish towel is applied to the fruits or 'sponges' in British Honduras. These sponges are much used in Guatemala for scrubbing and cleaning, and for many other purposes. Strange-appearing dolls are often made from them, and the sponges are a common article in the markets." Again according to Standley, "During the dry season the dry fruits are very conspicuous in the Pacific thickets, as they dangle from naked vines and are moved by the wind. The sap of the fruit has been found to
354 FIELDIANA: BOTANY, VOLUME 24
have purgative properties, and the oily seeds are reported to have emetic-cathartic properties . . ."
Luffa operculata (L.) Cogn. in Mart. Fl. Bras. 6, pt. 4: 12, t. 1. 1878. Momordica operculata L. Syst. ed. 10. 1278. 1759. Pashte. (Fig. 42).
Lowlands, near sea level; Escuintla; Izabal. Mexico (Sinaloa, Sonora, Nayarit, Guerrero, Chiapas); Colombia; Peru.
Herbaceous vines, often forming dense masses of scandent or prostrate growth; stems rather slender, striate, freely branching; tendrils mostly bifid on peduncles 1-6 cm. long; leaves on slender petioles mostly 2-6 cm. long, the blades cordiform- reniform in outline, 3-10 (-15) cm. long, scabrous on both surfaces, usually 3-5-lobate, the lobes constricted at the base, the lateral lobes smaller and obtuse, the margins entire to obscurely denticulate, basal sinus broadly arched to rectangular; flowers pale yellow, rather small, the staminate ones in slender racemes, the pistillate solitary or single in the axils with staminate flowers; staminate flowers: racemes 10-20 cm. long, exceeding the leaves, the 5-30 flowers borne on the distal third or less, only 1 or 2 in anthesis at a time, pedicels slender, 2-3 mm. long, bracteate at or near the base, the bracts glanduliferous. receptacle short, broadly campanulate, sepals triangular to ovate-acuminate, about 3 mm. long, 1-1.5 mm. wide at base, valvate in bud, median nerve prominent, petals 5, free, about 1 cm. long, ovate to wedge-shaped, obtuse and apiculate, 3-nerved, stamens 3 (one monothecous, two dithecous), wholly free or the anthers partly connate, inserted low on the receptacle, thecae gently curved and marginal on broad lobes of connective tissue, filaments linear, about 2 mm. long, 0.5 mm. thick, hairy at the base, pollen 3-colporate, reticulate, about 50 u in diameter; pistillate flowers: sessile or short-stalked, ovary fusiform, long-rostrate, tomentose and also tuberculate-spiny, ovules numerous, horizontal, receptacle very short, campanulate, petals and sepals similar to those of the staminate flowers, rudimentary stamens sometimes present, style linear, glabrous, sometimes 3-parted in the upper quarter, stigmas broadly lobed; fruit about the size of a hen's egg, or smaller, ellipsoid to broadly fusiform, short-stalked, rostrate, dry, operculate, obscurely ribbed, sparsely beset with short, thick spines, thin-walled, coarsely fibrous within, many-seeded; seeds oblong to elliptic in outline, immarginate, about 9 mm. long, nearly white to dark brown, dull, smooth, compressed, the hilar end of each face ornamented with a pair of ridges, hilum bordered by a short, thin collar.
This species is easily confused with Momordica charantia L., which it closely resembles in habit and overall appearance. It may be readily distinguished, however, by its fibrous, operculate fruits, or by its racemes of staminate flowers.
MELOTHRIA Linnaeus
References'. A. Cogniaux in Pflanzenreich IV, 275, 1: 75. 1916. C. Jeffrey, Notes on Cucurbitaceae, including a proposed new classification, Kew Bull. 15: 337 372. 1961 [1962].
Climbing or prostrate vines, monoecious, sometimes several m. long, rooting at the nodes when prostrate, often forming annual, herbaceous stems from a perennial
DIETERLE: FLORA OF GUATEMALA 355
rootstock; stems usually slender; leaves entire to more or less distinctly lobed, usually thin in texture, usually more or less scabrid or hispidulous; tendrils simple; flowers small; staminate flowers: pedicellate, in few to many-flowered, racemose, or subumbellate inflorescences at the apex of a common peduncle; receptacle campanulate, sepals short, dentiform, corolla yellow or white, deeply 5-parted, the segments entire, stamens usually 3, two dithecous, one monothecous, free (or sometimes the anthers lightly coherent), oblong or suborbicular, subsessile at about the middle of the receptacle tube, thecae straight or slightly curved, basal nectary (or rudimentary pistil) globose, often trilobate; pistillate flowers: solitary for the most part, rarely 2 or 3 in one axil, pedunculate, separate or in the same axils as the staminate, perianth similar to that of the staminate flowers, ovary globose, ovoid, or fusiform, contricted below the receptacle, triplacentiferous, ovules numerous, horizontal, style short, surrounded at the base by an annular or cup-shaped nectary, stigmas 3, bilobate, or 1 and trilobate, staminodes 3 or none; fruit small, globose or ellipsoid, fleshy, indehiscent, many-seeded; seeds numerous, ivory-colored, compressed, ovate, smooth, immarginate, the surface appearing satiny because of the long, shiny, tightly appressed hairs.
A large genus. Jeffrey proposed that it be limited to New World species by re-establishing the African genera Zehneria, Solena, and Mukia (Kew Bull. 15: 343. 1961): "Melothria is then left as an entirely New World genus of plants with long-stalked fruits and male racemes, compressed seeds, and 3 stamens, 2 of which are bithecous and the other monothecous."
My own experience is that Melothria can be found wherever there is sufficient moisture, warmth, and protection. It seems quite possible that a few species have attained very widespread distribution and may have been described several times. Vegetative characters are exceedingly variable; many species have been established on the basis of features that do not prove to be sufficiently constant or not easily verifiable because of lack of flower material. Melothria may be one of the genera of Cucurbi- taceae most in need of revision. Many of the described species may be merely growth forms, perhaps not even that. In the field, preparation of good specimens is difficult because the flowers are small and drop off easily. R. Martinez Crovetto (Darwinia 1: 496- 518. 1949) commends Cogniaux's monograph of the genus in DeCandolle, Monogr. Phan. 3: 572-626. 1881, and in Engler's Pflanzenreich IV, 275, I: 75-104, 1916, but I found it of little help with the material at hand.
Until more collections with good flowers and/ or fruits can be made, it seems inadvisable to attempt definition of the several species described as having leaves that are either entire or not deeply lobate. I have therefore chosen the oldest name, Melothria
356 FIELDIANA: BOTANY, VOLUME 24
pendula L., for this complex, and have treated all of our segregate forms as M. pendula L. sens lat.
Leaves deeply 3-5(-7)-lobate; fruits 4-6 cm. long M. trilobata.
Leaves entire, angulate, or shallowly lobate; fruits commonly 0.8-2 cm. long.
M. pendula sens. lat.
Melothria pendula L. sens. lat. Sp. PI. 35. 1753. Bryonia guadalupensis Spreng. Syst. Veg. 3: 580. 1881. M. fluminensis Gardner in Hook. Lond. Journ. Bot. 1: 173. 1842. M. quadalupensis Cogn. in DC. Monogr. Phan. 3: 580. 1881. M. donnell-smithii Cogn. in Donn.-Sm. Bot. Gaz. 16: 9. 1891 (type from Masagua, Escuintla, J. D. Smith 2203). M. donnell-smithii var. hirtella Cogn. loc. cit. (type from Escuintla, J. D. Smith 2206). M. donnell-smithii var. rotundifolia Cogn. loc. cit. (type from San Luis, Escuintla, J. D. Smith 2208). Mexito (Chimaltenango); sandia de raton (Chimal- tenango, Chiquimula, Peten).
Wet or dry thickets, forest, or brushy slopes, sometimes on grassy, open banks or old fields, frequently along roadsides and in hedges, sea level to 1,860 m.; Alta Verapaz; Chimaltenango; Chiquimula; Escuintla; Guatemala; Huehuetenango; Izabal; Ja- lapa; Peten; Quezaltenango; Retalhuleu; San Marcos; Santa Rosa; Suchitepequez; Zacapa. Southern United States; Mexico; British Honduras to Panama; South America.
Scandent or prostrate plants, the slender stems short or elongate, much branched, often rooting at the nodes, essentially glabrous or glabrate; leaves on slender, more or less pubescent petioles 2-6 cm. long, the blades thin, variable in outline, ovate-cordate, suborbicular, or triangular and more or less subsagittate, entire, angulate, or shallowly lobate, the apex acute, acuminate, or obtuse, mostly 3-7 (-10) cm. long, 3.5-7 cm. wide, the margins undulate or rather remotely denticulate, the basal sinus usually rounded, rarely acute, commonly 1-2 cm. deep, the upper surface usually more or less scabrous, paler beneath and sparsely or densely villous- hirsute or scaberulous or glabrate; tendrils simple; staminate flowers: small, disposed in 2-7-flowered racemes, peduncles filiform, pedicels 1-3 mm. long, receptacle narrowly campanula te to urceolate, sparsely pilose, hirtellous, or glabrate, 1.5-2.5 mm. long, sepals 0.2-0.5 (-1) mm. long, corolla yellow, the lobes ovate to obovate- oblong, obtuse or acute, or more or less emarginate, anthers suborbicular to oblong, the connective usually broad; pistillate flowers: on filiform peduncles 3-8 cm. long, ovary oblong to linear-oblong, style short, stigma obscurely lobed; fruits triplacen- tate, subglobose or ellipsoid to ovoid, mostly 0.8-2 cm. long, 0.6-1.2 cm. thick, dark, mottled green, purplish black when mature; seeds obovate, ivory colored, 3-4 mm. long, 2-3 mm. wide.
Material reported from Guatemala as M. scabra Naud. is referable to this variable species.
FIG. 21. Melothria pendula. A, habit, X Vfe; B, immature pistillate flower with receptacle laid open, X 10; C, immature staminate flower, opened, 10; D, immature pistillate flower, X 5; E, immature staminate flower, X 5; F, fruit, X IVa, opened to expose seeds; G, two views of seed, X 2'/2. See also Figure 43.
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Melothria trilobata Cogn. in Mart. Fl. Bras. 6, pt. 4: 26. 1878. Guisquil cimmarron (Huehuetenango); granadia (Izabal).
Damp thickets, often along streams, at altitudes from near sea level to about 1,000 m.; Heuhuetenango; Izabal; Jutiapa; Retal- huleu; Suchitepequez. Southern Mexico; British Honduras; Honduras; Panama.
Small, rather sturdy, herbaceous vines, scandent or creeping; stems pilose or glabrate, ribbed; leaves moderately large, on petioles 3-4 cm. long, these usually markedly villous, the blades 5-10 cm. long, rather firm in texture, deep green above, paler beneath, very scabrous, broadly orbicular to ovate in outline, deeply 3-5(-7)- lobed, the terminal lobe oblong-ovate, contracted at the base, the lateral ones variously lobulate, the margins entire or remotely denticulate, the basal sinus deep, rectangular to broadly arched; tendrils strong, glabrous; staminate inflorescences few-flowered, delicate, shorter than the petioles, flowers not seen; pistillate inflorescences not seen; fruit ellipsoid, 4-6 cm. long, 2.5-3 cm. thick, green or striped with light and dark green, becoming yellow in age; seeds numerous, ovate, 5-6 mm. long.
MICROSECHIUM Naudin
Scandent herbs from thickened, perennial roots, monoecious; leaves long- petiolate, cordate, mostly 3-5-lobate; tendrils proximally branched, 2-6-parted; flowers rather small, greenish white or greenish yellow, green-veined, the staminate and pistillate usually sharing common axils; staminate flowers: in racemes, receptacle short-funnelform to broadly campanulate, without pouchlike nectaries, sepals 4, petals 4, triangular-ovate, stamens 4, the filaments connate into a column, this 4- parted above the middle, the branches spreading, anthers free, monothecous, the thecae loosely replicate, flexuous; pistillate flowers: in subumbellate inflorescences on short peduncles with small leaflike bracts sometimes present in leaf axils at base of peduncle, perianth like that of staminate flowers but with only 3 sepals and 3 petals, ovary ovoid, unilocular, ovule one, pendant from the apex of the cell, style trifid at the apex, stigmas broad, reflexed; fruit clustered on short peduncles, rather small, to 1.5 cm. long (in ours), ovoid or globose, fleshy, few-aculeate, indehiscent, one-seeded.
A poorly known, variously circumscribed genus. I have chosen to adopt in part Cogniaux's conception of it as set forth in Monographiae Phanerogamarum, and to include only plants having 4-merous, staminate flowers with united filaments and free anthers.
Micro sechium helleri (Peyr.) Cogn. in DC. Monogr. Phan. 3: 910. 1881. Sicyos helleri Peyr. Linnaea 30: 56. 1859-60. M. ruderale Naud. in Ann. Sci. Nat. ser. 5, v. 6: 25. 1866. M. guatemalense Hemsl. Diagn. PL Mex. 16. 1878 (type from ridge above Calderas, Volcan de Fuego, Salvin s.n.}. Guisquil de raton.
Moist thickets or forest, often in brushy ravines, 1,000-2,800 m.; Chimaltenango; Chiquimula; Guatemala; Huehuetenango; Quezal-
FIG. 22. Microsechium helleri. A, habit, x Vfc; B, staminate flower, x 3V4; C, pistillate flowers, X 4; D, immature fruit, X 2.
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tenango; San Marcos; Sacatepequez; Suchitepequez. Southern Mexico.
Moderately large, herbaceous vines, sometimes climbing several m. high, puberulent or glabrous, often partly villous; main root thickened and woody; stems slender to rather stout; leaves on slender petioles 2-7 cm. long, the blades thin and soft to subrugose, broadly ovate or circular in outline, mostly 5-12 cm. long, shallowly to deeply 3-5-lobate with a deep basal sinus, green and more or less scabrous or short- hispid above, lighter green and more or less densely short- hirsute below, the margins entire to denticulate; sessile, reduced leaflets often found in leaf axils at base of inflorescences and branchlets; tendrils usually 3-parted; staminate flowers: racemes 7-20 cm. long, puberulent or villous or both, long-pedunculate, the flowers borne on the upper one-half or one-third or less, pedicels persistent, slender, 5-20 mm. long, puberulent, receptacle broadly and shallowly campanulate, without conspicuous foveolae within, sepals 4, subulate, about 3 mm. long, green, petals 4, triangular or ovate-triangular, separate to the base, light green or yellow-green, often conspicuously green-veined, stamens 4, monothecous, filaments united into a column, 4 -branched above the middle, thecae loosely replicate on the edges or undersides of the branch tips; pistillate flowers: disposed in umbelliform clusters on peduncles about 1 cm. long, perianth similar to that of staminate flowers but smaller and usually trimerous, ovary ovoid, unilocular, ovule one, pendulous from apex of locule, style slender, stigmas 3, circular to spoon-shaped, reflexed; fruits ovoid to subglobose, densely short-hairy or glabrous, 1-1.5 cm. long when dry, clustered 2-several on a common peduncle 1-3 cm. long, fleshy, one-seeded, sparsely aculeate.
A very variable species displaying broad divergence in depth of leaf-lobing, and in the amount and character of pubescence. Perhaps more than one species is involved. More study is needed.
MOMORDICA Linnaeus. Balsam-apple
Slender, prostrate or scandent herbs, monoecious, probably annual; leaves petiolate, the blades 3-5-lobate; tendrils simple; flowers rather small, yellow, solitary, the peduncle often bearing a prominent sessile bract; staminate flowers: receptacle shallowly cupular or short-funnelform, sepals 5, entire, rounded to lanceolate, corolla spreading to broadly campanulate, petals 5, free, one or more with an incurved scale inside at the base, stamens 3, inserted near the rim in the receptacle, the filaments free, short, the anthers coherent at least at first, one of them monothecous, the others dithecous, the thecae vertically sigmoid-flexuous, the connective not produced at the apex; pistillate flowers: perianth similar to that of the staminate flowers, ovary elongate-fusiform, strongly constricted at base, and apex, ovules numerous, horizontal, style slender, glabrous, stigmas 3, V-shaped; fruit small or medium-sized, fusiform to ellipsoid, fleshy, few to many-seeded, indehiscent or bursting by 3 valves to expose the seeds hanging from the segments of the wall; seeds medium-sized (about 12 mm. long, in ours), enveloped by a red, pulpy covering, testa smooth or variously sculptured.
About 40-45 species in the Old World, mostly African. One species, M. charantia L., thought to have been introduced from Africa, is firmly established in tropical America, where it occurs
DIETERLE: FLORA OF GUATEMALA 361
widely as a weed. M. charantia and one other species, M. balsamina L. are occasionally cultivated in America.
Momordica charantia L. Sp, PL 1009. 1753. Sorosi.
Moist or wet thickets, near sea level to about 500 m., but rare in most parts of the Guatemalan lowlands; Alta Verapaz; Escuintla; Izabal; Retalhuleu; perhaps in all the Pacific Coast departments. Southern United States; Mexico; British Honduras to El Salvador and Panama; West Indies; South America; Old World tropics; perhaps introduced in America.
Large or small herbaceous vines with usually much-branched stems, the branches glabrous, puberulent, or tomentose; leaves on long, slender petioles, the blades membranaceous, orbicular in outline, deep green, 3-8 cm. long and wide, deeply 5-7- lobate, the lobes mostly obovate, narrowed at the base, acute or obtuse, mucronate; tendrils whiplike, that is, each a slender, coiling thread on a stiff, straight stalk; staminate flowers: peduncles very slender, filiform, 5-7 cm. long, puberulent or glabrous, usually bearing near the middle a sessile, clasping, rounded-cordate to reniform green bract, 4-7 mm. broad, sepals ovate-lanceolate, acuminate, 3-4 mm. long, pubescent, appressed to the lower surface of the petals, petals obovate, obtuse or apiculate, 10-15 mm. long, strongly veined, pale yellow to stronger, brighter yellow, two with a recurved scale inside at the base, anthers coherent in center of flower; pistillate flowers: pedicel slender to rather sturdy, 2-4 cm. long, often bearing a small, green, clasping bract near the base, ovary elongate-fusiform, beaked, ribbed, tuberculate, tomentose, the sepals linear, acute, about 2 mm. long, 0.5 mm. broad, pubescent, petals similar to those of the staminate flowers but rather smaller; fruit 3- 6 cm. long, pendulous on a stalk 3-8 cm. long, bright orange when ripe, broadly ellipsoid to fusiform, tuberculate, beaked and usually bearing remnants of the sepals and pistil at its apex, indehiscent or eventually bursting by 3 valves and exposing the seeds enveloped in scarlet pulp; seeds oblong in outline, slightly flattened, to 12 mm. long, 5-6 mm. wide, the testa sculptured.
Standley wrote: "In some parts of the Central American lowlands this is a very common, weedy vine, often forming dense masses of foliage over tall, dead trees, but we have not found it common anywhere in Guatemala. It sometimes is planted about houses for ornament. The brightly colored fruits are handsome and often produced in abundance. When fully ripe, they split open, exposing the bright red pulp, which sometimes is eaten by children. In Yucatan the fruit is applied as a poultice to cure itch, sores, and burns."
PARASICYOS Dieterle
Reference: J. Dieterle, Parasicyos maculatus, a new genus and species of Cucurbitaceae from Guatemala, Phytologia 32: 289-290. 1975.
FIG. 23. Momordica charantia. A, habit, x Vz; B, small fruit, x 1, inset of sessile bract on peduncle, X 2; C, bursted fruit (normal position pendant) with seeds clinging to wall, X 1A, two views of seed with fleshy coat removed, above, X 1; D, staminate flower, X 4; E, stamens (filaments free, anthers coherent), X 9; F, androecial head from above, X 9. See also Figure 44.
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Herbaceous, monoecious, glabrous vines, the stems sparsely branched; tendrils commonly 3-parted; leaves petiolate, the blades usually trilobate; staminate flowers: pedicellate, disposed in panicles, receptacle tube campanulate, not foveolate, corolla 5-parted to the base, sepals 5, dentiform, stamen filaments connate in a column, the anthers sessile at its apex but more or less free along their upper length; pistillate flowers unknown; fruits 2-several on a common peduncle, fleshy, one-seeded, indehiscent; seeds solitary, pendant from the apex of the fruit locule, elliptic in outline, compressed.
Only one species is known.
Parasicyos maculatus Dieterle, Phytologia 32: 289. 1975.
Thickets along streams, cutover and second growth forest, cornfields, wet roadbanks, disturbed sites in pine forest remnants, coffee and banana plantations, 300-2,300 m.; Alta Verapaz (type from hills north of San Pedro Carcha, Williams, Molina, Williams, & de Molina 40205); Baja Verapaz; El Progreso.
Herbaceous, monoecious vines, the slender stems glabrous, sulcate, sparingly branched; tendrils usually 3-parted, the peduncle about twice as long as the neighboring petiole; leaves on petioles 2-8 cm. long, the blades membranaceous, suborbicular to ovate in outline, mostly 8-15 cm. long, usually trilobate, the lobes triangular, the lateral ones smaller and subauriculate, the basal sinus deeply rectangular to ovate, the upper surface deep green, scabridulous, the lower surface paler and smooth, margins commonly denticulate, rarely entire; staminate flowers: disposed in panicles 10-20 cm. long, the branches few, borne on the upper third or less, pedicels 5-9 mm. long, slender, long-persistent after anthesis, receptacle tube campanulate, not pitted with nectariferous foveolae, sepals 5, dentiform, corolla pale yellowish green, to about 8 mm. across, the petals triangular to ovate-triangular, separate to the base, more or less patent, minutely papillose inside, stamens with sigmoid-flexuous thecae; pistillate flowers not seen; fruits 2-several on a common peduncle 3-5 cm. long, the fruits unarmed, fleshy, 3-4.5 cm. long, first green with light green spots, later brick- red with green spots, indehiscent, one-seeded; seed 1.5-2 cm. long, elliptic, compressed.
POLYCLATHRA Bertoloni Reference: C. Jeffrey, Kew Bull. 25: 196-197. 1971.
Annual, herbaceous vines, more or less glandular-pubescent, scandent, monoe- cious; stems rather slender, often several m. long; leaves petiolate, the blades thin, softly hairy to short-hispid or scabridulous, cordate in outline, simple to angular or deeply trilobate, the basal sinus deep and broad, the margins entire to denticulate; tendrils mostly 4-parted, the branching proximal; flowers white, rather large, night- blooming, all solitary, the staminate and pistillate arising from separate or from common axils; staminate flowers: very long-pedicellate, receptacle tube funnelform or narrowly campanulate, sepals long-triangular, acute, corolla divided to below the middle, the lobes 5, broadly rounded and abruptly apiculate, spreading, green-veined on the outer surface, stamens 3, inserted on wall of receptacle tube, filaments free, threadlike, anthers connate, partially exserted, connective narrow, not produced,
FIG. 24. Parasicyos maculatus. A, habit, x Vr, B, branchlet of staminate inflorescence, x 2'/2; C, staminate flower, spread open to show androecium, x 5; D, halved fruit, X Vi.
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thecae tightly triplicate; pistillate flowers: perianth similar to that of staminate flowers but with markedly narrower sepals, receptacle-tube also similar but somewhat shorter, peduncle short and sturdy, ovary ellipsoid to globose, ovules numerous, horizontal, style slender, elongated, stigmas 3, bilobate, more or less exserted from the corolla tube; fruit moderate-sized, broadly ellipsoid or oblong, dry at maturity, the thin pericarp splitting from the apex and revealing tiers of horizontal, ligneous-walled locelli, each containing one seed; seeds broadly ovate in outline, strongly compressed, dull brown.
Jeffrey seems to think there are at least three species in Guatemala, but I have not been able to separate them.
Polyclathra cucumerina Bertol. Nov. Comm. Acad. Bonon. 4: 438, t. 36. 1840; Fl. Guat.: 38, t. 12. 1840 (type from Escuintla, Velasquez s.n.) Cayaponia grandiflora Cogn. in DC. Monogr. Phan. 3: 779. 1881. Pittiera longipedunculata Cogn. Bull. Soc. Bot. Belg. 30: 271. 1892. P. trilobata Cogn. in Coult. Bot. Gaz. 20: 289. 1895 (type from Guatemala, Buena Vista, Heyde & Lux 4189). Roseanthus albiflorus Cogn. Contr. U. S. Nat. Herb. 3: 578. 1896. Pittiera grandiflora Cogn. ex Rose, op. cit. 5: 123. 1897. P. parvifolia Cogn. & Rose ex Rose, loc. cit. (type from Guatemala, sin loc., Nelson 3532). Roseanthus elongatus Rose, op. cit. 8: 337. 1905. R. heterophyllus Brandg. Univ. Calif. Publ. Bot. 10: 188. 1922. Polyclathra longipedunculata (Cogn.) C. Jeffrey, Kew Bull. 25: 196. 1971. P. grandiflora (Cogn.) C. Jeffrey, torn. cit. 197. P. albiflora (Cogn.) C. Jeffrey, loc. cit.
Damp or dry thickets, brushy, rocky slopes, 300-1,360 m.; Chiquimula; Jalapa; Retalhuleu. Mexico; Honduras; El Salvador; Nicaragua; Costa Rica.
Slender vines, the stems glandular-puberulent, glandular-pubescent, short- pilosulous, or glabrate; tendrils 4-parted; leaves on slender petioles mostly 2-8 cm. long, the blades thin, rounded-cordate or broadly ovate-cordate in outline, mostly 4- 12 cm. long, sometimes merely angulate, sometimes shallowly or deeply trilobate, usually deeply cordate at the base, the margins subentire to more or less denticulate, the upper surface puberulent or scabrous or sometimes glabrate, the lower surface usually densely and softly pubescent or puberulent, sometimes scaberulous; staminate flowers: on slender peduncles mostly 7-25 cm. long, the receptacle narrowly or broadly cylindrical, 1-2 cm. long, densely puberulent, viscid-puberulent, villous, or glabrate, as much as 1 cm. broad at the dilated apex, the sepals linear-subulate, 8-18 mm. long, corolla white, 2-2.5 cm. long, sometimes 6-8 cm. across, the lobes obovate- oblong, obtuse or acute, filaments pilose at the base, 11-12 mm. long, the anthers connate, thecae triplicate; pistillate flowers: on short, sturdy peduncles 0.5-1.5 cm. long, perianth similar to that of the staminate flowers but the receptacle tube shorter and the sepals narrower, ovary ellipsoid to subglobose, style slender, stigmas 3; fruit broadly ellipsoid or oblong, 4-6 (-8) cm. long, 3-4 cm. wide, green with white stripes, yellowing in age; seeds numerous, horizontal, broadly ovate, compressed, about 6 mm. long.
FIG. 25. Polyclathra cucumerina. A, habit, X W, B, leaf, x Vr, C, atypical leaf form, x Vr, D, staminate flower, x !/2, perianth laid back to display filaments; E, pistillate flower, corolla removed, x l'/2, another view of stigmas above; F, fruit (part of wall removed to uncover tiers of seed chambers), X Vr, G, seed, X '/2 (left), X 1'4 (right). See also Figure 45.
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POSADAEA Cogniaux
Prostrate or climbing herbs, monoecious, somewhat pilose on petioles and younger growth; leaves petiolate, the blades rough-membranous, dull green above, lighter below, shallowly angular to deeply trilobate, cordate at base, the lobes rounded and apiculate at apex, margins distantly and minutely toothed; tendrils simple or bifid; flowers rather small, yellow, the staminate in short, slender racemes, the pistillate single, often coaxillary with the staminate; staminate flowers: receptacle short-campanulate, sepals 5, ovate-lanceolate, corolla spreading, 5-parted to the base, segments somewhat reflexed, more or less notched at apex, stamens 3, one monothecous, the others dithecous, free, dorsally affixed midway on receptacle, filaments short, connectives narrow, not prolonged beyond the thecae, thecae linear, recurved at apex, pollen grains 3-colpate, reticulate, espinulose, pistillodium cupuliform, obscure; pistillate flowers: perianth as in staminate flowers; fide Cogniaux: staminodia 3, short, ligulate, ovary elliptic-ovoid, triplacentiferous, style short, thick, stigmas 3, obcordate, dilated, subreflexed; ovules numerous, horizontal; fruit spherical, indehiscent, densely and minutely verruculose, pericarp woody, flesh pulpy, peduncle long, tapering from a slender base to a broadly thickened apex confluent with curve of fruit; seeds narrowly obovate, compressed, immarginate, smooth.
The genus, represented by a single species, is named in honor of the Colombian botanist, Dr. Andres Posada-Arango.
Posadaea sphaerocarpa Cogn. Bull. Acad. Belg. 3 ser. 20: 477. 1890; Cogn. in Pflanzenr. IV, 275, I: 252. 1916; Posada-Arango in Bull. Soc. Bot. France 38: 243. 1891.
Woods or thickets, scrambling over other plants or on the ground, 100-350 m.; Alta Verapaz; Peten. Panama (fide Steyer- mark); Colombia; Trinidad (fide Hutchinson).
Vines up to several m. long, the branches rather slender, striate, with flexuous, multicellular hairs on petioles and newer growth; leaves on strong petioles 3-9 cm. long, the blades 11-15 (-18) cm. long, 8-14 (-19) cm. broad, sparsely pilose on both surfaces at first, then punctate-scabrous, especially on upper surface, basal sinus subrectangular, 3-4.5 cm. deep, 1.5-2.5 cm. wide, partially bordered by the principal basal veins; tendrils rather long and robust, thickened toward the base; staminate flowers: peduncles of inflorescences slender, (2-) 3-4 (-6) cm. long, sparsely pilose, the racemes 3-4(-5)-flowered at apex, pedicels densely pilosulous, 8-14 mm. long, sometimes minutely bracteolate, receptacle glabrous to sparsely villosulous outside, short-pilose on upper half inside, 6-8 (-9) mm. long, 8-12 mm. in diameter at apex, sepals reflexed, acute, 3-5.5 mm. long, 1 mm. wide, glabrous, corolla segments dull yellow, obovate-oblong, 8-9 mm. long and wide, 7-9-nerved, sparsely short-villous on both surfaces, anthers white, 1.5-2 mm. long; pistillate flowers: peduncles robust, densely short-pilose, 2-3 cm. long, sepals 2 mm. long, corolla segments ovate-oblong, rounded at apex, 1 cm. long, 4-5 mm. wide, staminodia obtuse, 1.5-2 mm. long, ovary sparsely pilose, disc at base of style none or obscure, style 3 mm. long, stigmas said to be canescent, 5 mm. broad; fruit 8-10 cm. in diameter, peduncle 9-15 cm. long, 3-5 mm. thick at base, 15-17 mm. thick at apex, glabrous; seeds narrowly obovate, 12-14 mm. long, 7-8 mm. wide, 2-2.5 mm. thick.
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FIG. 26. Posadaea sphaerocarpa. A, habit (section of vine with staminate inflorescence), x 1/2; B, another leaf form, X 14; C, fruit, about half-mature, x Vz.
Guatemalan collections of this species are few. Details of the pistillodium and female flower structure are taken from descriptions by Cogniaux. The fruits resemble those of Lagenaria (tecomate). Steyermark (No. 45220) and Posada- Arango report that they are useful for holding water.
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PTEROSICYOS Brandegee
Vigorous, monoecious vines, climbing or trailing; leaves membranaceous; tendrils proximally trifid; staminate inflorescences paniculate or paniculate-racemose; perianth of pistillate and staminate flowers similar; receptacle shallowly cup-shaped, with 10 nectaries making a circle of pouchlike pits at base, corolla deeply 5-parted, stamens united, the filaments forming a central column, the anthers paired at apex of filament column, in a 2-2-1 pattern, thecae vertically flexuous-replicate, connectives not produced; pistillate flowers in an umbellate or subumbellate inflorescence on a peduncle co-axillary with staminate inflorescence: mature pistillate flowers not seen, stigmas said to be 2; fruits samaroid, 1-seeded, the seed pendulous from the apex of the locule.
Very similar to Sechiopsis, differing chiefly in producing two- winged rather than three-winged fruits. The genus consists of a single species.
Pterosicyos laciniatus Brandegee, Univ. Calif. Publ. Bot. 6: 72. 1914.
Vines in thickets, 1,200-1,500 m.; specimens without flowers from Quezaltenango, Suchitepequez, and San Marcos assumed to be this species. Mexico (type from Cerro del Boqueron, Chiapas).
Herbaceous vines, glabrous or nearly so; stems striate; leaves long-petiolate, the blades elliptic or orbicular in outline, 10-17 cm. long, rich green and often white- punctate above, lighter green and glabrous below, deeply 3-7-lobed, the lobes lanceolate or narrowly obovate, somewhat constricted at base, the lateral sinuses rounded at base, open, the outermost lobes shorter, often auriculate, the basal sinus broadly arched, blade margins shallowly and remotely toothed, or entire, petioles 2-4 cm. long; tendrils trifid, on peduncles shorter than the adjacent petioles; staminate flowers: in few-branched panicles shorter than the leaves, pedicels capillary, 12-20 mm. long, receptacle shallowly cup-shaped, 10-foveolate, the sepals about 0.75 mm. long, broadly triangular, obtuse, remote, corolla about 2 cm. broad, deeply 5-parted, the segments acute, 5-7 mm. long, 7-9-nerved, filament column 1.5-2 mm. long, anthers about 2 mm. long; pistillate flowers: about 10 in subumbellate inflorescences (mature flowers not seen); fruit membranous, strongly compressed, alate, indehiscent, narrowly elliptic-rhombic in outline, unequally cordate at base, emarginate at apex, 5.5-6 cm. long, 2 cm. broad at midlength, one prominent vein extending from base to apex, pedicels about 2 cm. long, peduncle about 3 cm. long; seed solitary, pendulous from apex of locule.
RYTIDOSTYLIS Hooker & Arnott
Herbaceous vines, probably annual, monoecious, the stems usually glabrous, slender; leaves long-petiolate (in ours), at least on mature stems, the blades thin, cordate, simple or 3 (-5) lobate; flowers small, yellow, white, or green; staminate flowers: (in ours) crowded at the tips of long-pedunculate racemes, the pedicels filiform, receptacle elongate-tubular, sepals 5, dentate or filiform, commonly minute, corolla 5-parted almost to the base, the lobes narrowly linear, tapering to a subobtuse
FIG. 27. fterosicyos laciniatus. A, habit (section of a vine with 3-lobed leaves), X Vr, B, habit (leaf, tendril, and staminate inflorescence), x 1; C, staminate flower, x 5; D, staminate flower, underside, x 5; E, staminate bud, x 5; F, fruit, X Vz.
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DIETERLE: FLORA OF GUATEMALA 371
apex, usually reflexed in full anthesis, filaments united in an elongate central column, anthers connate into an oblong or globose head, thecae linear, replicate, joined end- to-end in a continuous series; pistillate flowers: solitary, short-pedunculate, the perianth like that of the staminate, ovary ovoid, usually densely hirsute or setulose, ovules numerous, ascending, style elongate, slender, stigma cylindric-globose; fruit gibbous, somewhat compressed, fleshy, rupturing explosively to release an elongate placental column which throws out the seeds; seeds flattened, more or less turtle- shaped in outline, less than 1 cm. long, the margins appendaged.
The name widely accepted for this genus, Elaterium Jacquin, is a homonym of Elaterium P. Miller, a name rejected in favor of the conserved name Ecballium A. Richard. I have therefore employed Rytidostylis Hooker & Arnott, which is taxonomically synonymous with Elaterium Jacq., and the choice dictated by nomenclatural rules of precedence. Abandoning the highly appropriate name Elaterium seems regrettable, however.
About a dozen species have been described (perhaps not more than half of them valid), all in tropical America, with two in Guatemala.
Petioles essentially glabrous; upper leaves subsessile or short-petiolate; lower leaf surface minutely scabrous or essentially glabrous; basal sinus commonly densely ciliate; pedicels and receptacles glabrous; sepals minute, usually less than 0.5 mm. long (rarely to 1 mm. long) R. gracilis.
Petioles villosulous; upper leaves seldom if ever subsessile or conspicuously short- petiolate; lower leaf surface more or less villosulous to subtomentose; basal sinus not ciliate; pedicels villous, receptacles usually sparsely pubescent to villosulous; sepals 1-3 mm. long R. macrophyllus.
Rytidostylis gracilis Hook. & Arn. Bot. Beechey Voy. 424, t. 77, f. A. 1841. Elaterium gracile Cogn. Diagn. Cucurb. 51. 1877. E. ciliatum Cogn. op. cit. 54. E. ciliatum var. major Cogn. op. cit. 866. E. saepicola Brandg. Univ. Calif. Publ. Bot. 6: 72. 1914. Cochinito (Chiquimula); quiamul (Retalhuleu); sandia de raton (Jutiapa).
Damp or wet thickets, forest, weedy clearings, sometimes on open banks, sea level to 1,400 m.; Chimaltenango; Chiquimula; Escuintla; Guatemala; Izabal; Jutiapa; Peten; Retalhuleu; Sacate- pequez; Zacapa. Mexico; British Honduras; Honduras and El Salvador to Panama. Said to be in Colombia and Venezuela.
Herbaceous vines, small or large, usually much branched, climbing over bushes or rarely prostrate, the stems slender, essentially glabrous or bifariously short- pubescent, villosulous, or puberulent; leaves on slender, glabrous (rarely somewhat pubescent) petioles mostly 1-4 cm. long, or the upper leaves sessile or nearly so, the blades ovate-rounded to rather broadly ovate-angulate, mostly 4-8 cm. long, 3-6 cm. wide, obtuse or acute, shallowly lobate or merely angulate, both surfaces minutely scabrous or in age almost glabrous, the margins minutely and rather remotely
372 FIELDIANA: BOTANY, VOLUME 24
denticulate, basal sinus broad and open or, when blades are lobate, the auriculate lower lobes sometimes almost closing the sinus, margin of sinus usually more or less conspicuously white-ciliate, usually densely so but sometimes only sparsely ciliate, or some leaves with ciliate sinuses and others eciliate; tendrils usually bifid; staminate flowers: peduncles commonly glabrous, 5-20 cm. long, pedicels glabrous (rarely more or less villosulous), filiform, 3-8 mm. long, subumbellate at the apex of the peduncle, receptacle commonly glabrous (rarely somewhat villosulous), mostly 1.5-2 cm. long, sepals usually minute, almost obsolete, less than 0.5 mm. long, rarely linear and as much as 1 mm. long, corolla white, cream, or greenish, petals narrowly linear, tapering gradually into the subobtuse apex, 8-12 (-15) mm. long, spreading, stamens about equalling the receptacle, androecial head small, oblong, usually not more than 2 mm. long; pistillate flowers: subsessile, perianth similar to that of the staminate flowers, ovary densely hirsute; fruit subreniform, 1.5-2.5 cm. long, 9-12 mm. thick, containing 12-16 seeds 5-7 mm. long.
Rytidostylis macrophyllus (Standl. & Steyerm.) Dieterle, comb. nov. Elaterium macrophyllum Standl. & Steyerm. Field Mus. Bot. 23: 94. 1944 (type from Baja Verapaz, von Tuerckheim II. 1728).
Damp or wet thickets or mixed forest, 500-2,300 m.; Baja Verapaz; Quezaltenango; Retalhuleu; San Marcos; Suchitepequez.
Small or large vines, scandent, much branched, the stems slender, densely villosulous or glabrate; leaves on slender, villosulous petioles mostly 2-6 cm. long, the uppermost leaves petiolate, blades rounded-ovate or triangular-cordate, mostly 7-12 cm. long and 6-9.5 cm. wide, acute or acuminate, usually obtusely angulate, rarely lobate, the margins rather remotely denticulate or subentire, the upper surface scaberulous or almost glabrous, the lower more or less villosulous to subtomentose, basal sinus rounded, often closed; staminate flowers: peduncles and pedicels more or less villous, receptacle usually villosulous to sparsely pubescent, rarely glabrate, sepals 1-3 mm. long, spreading, corolla greenish, the lobes 10-15 mm. long; pistillate flowers and fruits not seen.
SCHIZOCARPUM Schrader
Herbaceous vines, rather slender to moderately strong-growing, monoecious, scandent, annual; leaves petiolate, the blades cordate-ovate, entire or trilobate; tendrils proximally bifid; flowers rather large, solitary, yellow, usually with greenish black or dark purplish blotches or shading inside the corolla; staminate flowers: receptacle narrowly campanulate to shallowly cup-shaped, more or less constricted below insertion of stamens and slightly inflated at the base by a fleshy, globose- cylindrical, hollow-cored gland, the 5 sepals elongate, corolla campanulate to tubular or funnelform, 5-lobed to about midlength, the lobes ovate-oblong, acute, stamens 3, inserted on the wall of the receptacle tube, filaments free, elongate, anthers coherent in an elongate head, one of them monothecous, the others dithecous, thecae longitudinally tightly triplicate, the connective narrow and not produced; pistillate flowers: perianth like that of the staminate flowers but usually smaller, the petals smaller, the sepals shorter, receptacle shallowly cup-shaped, ovary ovoid, ovules numerous, horizontal to ascending, base of style surrounded by a fleshy nectary
FIG. 28. Rytidostylis macrophyllus. A, Tajumulco Volcano (from the slopes of which the plant illustrated was collected, Williams, Molina, & Williams 26234, 4220 m. alt.); B, habit, x l/z; C, staminate flower dissected to expose androecium, x 1%; D, peduncle and ovary of pistillate flower, X 2Vfc; E, two views of androecial head, X
7'/2.
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374 FIELDIANA: BOTANY, VOLUME 24
overlaid by scales from the base of the corolla, stigmas 3, V-shaped; fruit of medium size, pyriform, ovoid, or fusiform, terete to 3-sided, dry, pendulous by a short stalk, pericarp thin, dehiscence by partial separation from the apex downward or the pericarp shattering irregularly from the base up, exposing firm-walled, one-seeded, upwardly inclined chambers, with release of the seeds following the shattering away of the pericarp along the rows of chambers or upward from the base; seeds numerous, smooth, oblong-ovate to triangular in outline, flattened, the margins irregular.
About seven species in Mexico; one in Guatemala, rarely collected. The only Guatemalan material I have seen is two sheets of the type collection of S. guatemalense Cogn. ( = S. filiforme Schrader), Heyde & Lux 4186.
Schizocarpum filiforme Schrader, Linnaea 6, Litt. Ber. 73. 1831. S. guatemalense Cogn. in J. D. Smith, Enum. PI. Guat. 3: 34. 1893, and in Coult. Bot. Gaz. 20: 290. 1895 (type from Santa Rosa, Heyde & Lux 4186).
In Guatemala, known only from the type material of S. guatemalense collected at Carrizal, Santa Rosa, 1,500 m., Heyde & Lux 4186. Mexico.
Rather slender vines, 1-4 m. long, very variable in pubescence and in degree of lobing in the leaf blades; petioles 1.5-5 (-8) cm. long, leaf blades more or less ovate in outline, shallowly or deeply lobate, mostly 5-10 cm. long and 4-7 cm. wide, usually glabrate above and rather densely puberulent beneath; staminate flowers: peduncles slender, 4-10 cm. long, receptacle campanulate, broad and short to narrow and long, the sepals filiform or long-acuminate, 5-6 mm. long, corolla 2.5-5 cm. long; pistillate flowers: peduncles 0.2-1 cm. long, ovary ovoid, villous, corolla 1.5-2.5 cm. long; fruit ovoid-conical, 3-5 cm. long, lightly 10-costate, rounded at the base, gradually constricted toward the obtuse apex, more or less pilose, the pericarp shattering irregularly from the base upward at maturity; seeds obovate to ovate-triangular in outline, 5-6 mm. long, strongly compressed, dark.
SECHIUM P. Browne. Chayote
Rampant, perennial, monoecious vines arising from very large, tuberous roots, more or less hispidulous or scabrous, high-scandent; leaves long-petiolate, membrana- ceous, deeply cordate at the base, palmately angulate or lobate; tendrils 3-5-parted; flowers small, the staminate in elongate racemes or elongate, very short-branched panicles, the pistillate flowers usually single in the same axil as the staminate; staminate flowers: receptacle shallowly cup-shaped with a circle of 10 pouchlike nectaries in the bottom; corolla 5-parted to the base or nearly so; filaments of stamens united in a central column with 5 short, spreading arms at the apex, the arms obscurely paired at their bases in a 2-2-1 combination, each arm bearing an anther locule on its dorsal surface, the locules sigmoid-flexuous; pistillate flowers: somewhat smaller than the staminate but with similar perianths, the ovary obovoid, unilocular, the solitary ovule pendulous from the apex; style short, slender, the stigma subglobose; fruit large, fleshy, pyriform, somewhat flattened, armed with stiff or fleshy spines, or unarmed, indehiscent, the single seed germinating within the fruit; seed large, solitary, compressed, the testa ligneous, smooth.
DIETERLE: FLORA OF GUATEMALA
375
FlG. 29. Schizocarpum filiforme. A, habit (section of a fruiting vine), X !/2j B, staminate flower, partially dissected to expose stamens, X 2; C, pistillate flower opened to show style and stigma, X 2; D, seeds and dehiscing fruit, X '/2. See also Figure 46.
The genus consists of a single species. However, Ahzolia composite (Donn.-Sm.) Standl. & Steyerm. may not be generically distinct from Sechium edule (Jacq.) Swartz.
Sechium edule (Jacq.) Swartz, Fl. Ind. Occ. 1150. 1800. Sicyos edulis Jacq. Enum. PI. Carib. 32. 1760. Huisquil; quisquil; chayote
FIG. 30. Sechium edule. A, habit, x Vfe; B, staminate flower with perianth spread open (10 foveolae at base), X ZVz; C, staminate buds, X 1; D, pistillate flower, X IVz; E, pistillate flower with perianth spread open, X 3; F, longitudinal section of an immature fruit, X Vz.
376
DIETERLE: FLORA OF GUATEMALA 377
(not a common name in Guatemala); chimd (Quecchi, Alta Verapaz); chimaa (El Quiche); perulero (Huehuetenango); chayo- tera (the plant); chintla, ichintal (the root); chuma (Poconchi); quisquilar (the plant).
Common in cultivation in Guatemala at all elevations except perhaps the highest and lowest; said to be cultivated commonly in tropical regions of both hemispheres.
Vigorous vines arising from large, thickened roots, often climbing to considerable heights; stems glabrous or nearly so, sulcate; leaves long-petiolate, the petiole glabrous or subglabrous, 4-15 cm. long, the blades 5-18 cm. long and broad, with a deep basal sinus, 3-5-angled or shallowly lobed, the terminal lobe often somewhat longer and narrower than the lateral ones, surface scabrous above, less so or glabrate below, margins entire or minutely and remotely denticulate; tendrils 3-5-parted, on sturdy, sulcate peduncles, glabrous; staminate flowers: inflorescences 10-30 (-50) cm. long, much exceeding the leaves, flowers clustered in a few fascicles or on short branches along the rachis, the pedicels 1.5-2 mm. long, tomentose, receptacle broadly and shallowly cup-shaped, about 5 mm. broad, sepals narrowly triangular, 5-6 mm. long, about 1-1.5 mm. at base, well-developed before the petals, corolla subrotate, pale green or yellowish, glabrate to finely papillose-glandular, petals triangular to oblong-triangular, acute, 6-7 mm. long, filament column slender, glabrous, pollen subglobose, flattened at the poles, 8-colpate, spiny, about 60 n in diameter; pistillate flowers: greenish, somewhat smaller than the staminate, peduncle short, less than 5 mm. long, ovary obovoid, constricted below the receptacle, style columnar, glabrous, 2-4 mm. long, stigma subglobose, about 4 mm. broad; fruit green, yellow, or nearly white, fleshy, spiny or unarmed, one-seeded, edible, varying in size from that of a small pear to about 18 cm. in length, sometimes splitting in age at the apex and permitting viviparity or germination before planting.
This is one of the most important and favorite vegetables of Guatemala and of most other parts of Central America. The plant is perhaps unique in that every part of it may be eaten at some stage or another, roots, leaves, flowers, fruits, and seeds. It thrives best in mountain regions, and is little grown in the lowlands. The vines are very productive. Martinez states that in Mexico a vine may produce in a year 80-100 fruits and 20-25 kilos of roots. He reports also that the flowers are much visited by bees.
SICANA Naudin
Large, herbaceous, monoecious vines, almost glabrous, arising from a perennial root, often climbing to the tops of tall trees; leaves long-petiolate, palmately 5-9- lobate, glabrous and glossy, the lobes triangular, acute; tendrils 3-5-parted; flowers all solitary, rather large, yellow; staminate flowers: receptacle short-campanulate, sepals 5, triangular-ovate, reflexed, corolla 5-lobate to below the middle, the lobes ovate-oblong, acute, stamens 3, inserted in the mouth of the receptacle, the filaments short, anthers free, connivent into a head, the thecae sigmoid-flexuous; pistillate flowers: perianth like that of the staminate flowers, staminodia said to be 3,
378 FIELDIANA: BOTANY, VOLUME 24
elongated, linear, ovary oblong-ovoid or cylindric, with 3 placentae, style said to be short, obconic, the 3 stigmas thick, obscurely bilobate, papillose, ovules very numerous, horizontal; fruit large, dark red, ellipsoid, fleshy, indehiscent, many- seeded; seeds oblong-ovate, flattened, smooth, conspicuously marginate.
The genus consists of a single species.
Sicana odorifera (Veil.) Naud., Ann. Sci. Nat. IV. 18: 181, t. 8. 1862. Cucurbita odorifera Veil., Fl. Flum. 10, t. 99. 1827. Melocotdn; melon de olor.
Native perhaps of Brazil (fide Standley) but often cultivated in other tropical regions for its edible fruit; planted commonly in the lowlands of Guatemala, especially along the Pacific foothills, also about Coban in Alta Verapaz, in Guatemala, and probably elsewhere.
Young stems puberulent; petioles 4-12 cm. long; leaf blades suborbicular, mostly 10-20 cm. long, nearly smooth, with a large, open sinus at the base, lobate almost to the middle, the lobes acute or acuminate, the margins undulate or denticulate, the basal sinus closely bordered by the lowermost lateral nerves; tendrils mostly 4- parted; staminate flowers: on sturdy, striate, minutely puberulent peduncles 3-5 cm. long, receptacle 5-8 mm. long, sepals broadly triangular, 9-12 mm. long, about 5 mm. broad, reflexed even in bud, corolla said to be subcarnose and tomentose inside, the segments 5-nerved, 2-3.5 cm. long, filaments glabrous, 3-5 mm. long, the anthers 6-7 mm. long; pistillate flowers: on puberulent peduncles 0.5-2 cm. long, ovary oblong- ovoid or cylindric, somewhat puberulent; fruit ellipsoid, smooth, to 30 cm. long, dark red or purple, the flesh pale yellow, juicy, sweet-scented; seeds conspicuously marginate, 10-12 mm. long.
The vines are sometimes trained over trellises but more often are allowed to climb over trees and frequently go to the tops of trees 15 m. high or more. According to Standley, the fruits are sometimes eaten raw, often flavored with lime juice. It cannot be said that the flesh, in spite of its delicious odor, is very good to eat for it is so tough that it is difficult to chew and swallow it. It is frequently employed for making preserves and other sweets. Although produced mostly in the lowlands, the fruits are often carried up to the markets of such places as Quezaltenango and Huehuetenango.
SICYDIUM Schlechtendahl
Scandent, dioecious herbs, tomentose or glabrate; leaves petiolate, the blades ovate or oblong-ovate, entire, deeply emarginate or cordate at the base; tendrils bifid at the apex; inflorescences paniculate; flowers minute, whitish, the pedicels slender, bracteate; staminate flowers: receptacle pateriform, the 5 sepals ovate, triangular, or linear, corolla deeply 5-parted, the petals lanceolate or ovate-triangular, stamens 3, free, the filaments short, 2 of the anthers dithecous and didymous, the other
FIG. 31. Sicana odorifera. A, habit (part of a stem with staminate flowers), X Vz; B, staminate flower with corolla laid open to expose anthers, X iVz; C, very immature fruit, X Vz.
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380 FIELDIANA: BOTANY, VOLUME 24
monothecous, the thecae short and straight, pollen ellipsoid, 3-colporate, longitud- inally striate, pistillodium none; pistillate flowers: perianth like that of the staminate flowers, staminodia 3, often antheriferous, ovary ovoid, unilocular, styles 3, spreading, the stigmas linear, ovule one, pendulous from the apex of the locule; fruit small, globose, fleshy, one-seeded, indehiscent; seed pendulous, globose, the testa crusta- ceous, rough with sharp peaks and ridges.
Six species; southern Mexico, Central America, and tropical South America, with three in Guatemala.
Filaments very short, anthers nearly sessile; leaves softly and densely villous-
tomentose on both surfaces S. tamnifolium.
Filaments as long as anther cells, or longer; leaves glabrate, or if pubescent, chiefly so on lower surface.
Leaves membranaceous, glabrous or glabrate S. glabrum.
Leaves coriaceous, scabridulous or thinly pilose above, densely pilose below.
S. tuerckheimii.
Sicydium glabrum Standl. & Steyerm. Field Mus. Bot. 23: 95. 1944. Hoja de esperanza; murcielago triangulo; paxte de montana.
Thickets or mixed forest, 1,200-1,800 m.; Quezaltenango (type from Montana Chicharro, lower southeastern slopes of Volcan de Santa Maria, Steyermark 34273); Solola; Suchitepequez. Mexico (Chiapas).
Scandent herbs, the slender stems obtusely angulate, glabrous or essentially so; tendrils glabrous; petioles slender, 0.5-3 cm. long, glabrous; leaves firm-membrana- ceous, the blades ovate or oblong-ovate, 7-11 cm. long, 3-6 cm. wide, long-acuminate, deeply cordate at the base with a broad, open sinus, entire, the nerves prominent beneath, smooth on both surfaces, glabrous or very minutely puberulent; inflorescences paniculate; staminate flowers: the panicles laxly and diffusely branched, 8-14 cm. long, the branches very slender, often reflexed, the pedicels capillary, 3 mm. long or less, glabrous or very minutely puberulent, articulated above the middle, receptacle saucer-shaped, the outside sparsely puberulent, the sepals oblong-triangular, acute, 0.5 mm. long, puberulent, petals ovate-triangular, glabrous, 1-1.5 mm. long, filaments equal to the thecae in length or a little longer; inflorescence of pistillate flowers lax, about 15 cm. long, the pedicels short and thick, the flowers not seen; fruit globose, purple-black, about 7 mm. in diameter, glabrous or scantily and minutely puberulent; mature seeds not seen, immature seeds spherical, 2-3 mm. in diameter, the testa rough with sharp peaks and ridges, and raised-crateriform around the hilum.
Sicydium tamnifolium (HBK.) Cogn. in DC. Monogr. Phan. 3: 905. 1881. Fevillea tamnifolia HBK. Nov. Gen. & Sp. 7: 175. t 640. 1825.
Lowland thickets or forest; Escuintla; Peten; Quezaltenango; Retalhuleu; Suchitepequez. Southern Mexico; British Honduras to Panama, southward to Ecuador and Bolivia; West Indies.
FlG. 32. Sicydium glabrum. A, habit (part of a fruiting vine), X Vz; B, staminate flower, X 12'/2; C, staminate flower, lower side, X l2Vz; D, mature fruit, seed, and section of seed, X 3.
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382 FIELDIANA: BOTANY, VOLUME 24
Rather slender, much-branched, softly glandular-hairy vines, the stems short- villous, sulcate; petioles slender, villous, 1.5-4.5 cm. long; tendrils short-pubescent; leaf blades thin and soft, oblong-ovate to broadly ovate, 5-10 cm. long, 4-9 cm. broad, acute or acuminate, entire or rarely somewhat angulate-lobate, deeply and openly cordate at the base, short-villous above and soft to the touch, densely villous or subtomentose beneath; staminate flowers: panicles mostly 6-12 cm. long; the branches divaricate, bracteolate, densely hirtellous, the pedicels capillary, short- pilose, articulated in the lower half, 2-3 mm. long, receptacle saucer-shaped, somewhat thickened, short-villous outside, sepals ovate-triangular, 0.5 mm. long, reflexed, short-villous, corolla greenish white, the lobes lanceolate-linear or narrowly long-triangular, spreading, 1-1.5 mm. long, filaments very short, anthers nearly sessile, thecae 0.2 mm. long, pollen ellipsoid, 3-colpate, closely striate longitudinally; pilstillate flowers: panicles 4-15 (-18) cm. long, the branches divaricate, densely pilose, pedicels short-villous, about 1.5 mm. long at anthesis, 6-12 mm. on fruits, ovary ovoid, densely pilose, constricted at the apex, less than 2 mm. long, the shedding of the perianth long delayed; fruit globose, fleshy, one-seeded, 6-7 mm. in diameter, nearly black when mature, sparsely pilose; seed globose, 4-4.5 mm. in diameter, the testa very rough but less so around the hilum-micropyle area.
Sicydium tuerckheimii Donn.-Sm., Bot Gaz. 52: 49. 1911.
Lowland thickets or mixed forest, 350 m. or less; Alta Verapaz (type from Cubilgiiitz, Tuerckheim II. 1914); Izabal; Peten. British Honduras.
Herbaceous vines, more or less glandular-pilose; leaves on petioles mostly 2-3 cm. long, the blades subcoriaceous, oblong-ovate to broadly ovate, 12-16.5 cm. long, 4.5-10 cm. broad, acuminate, deeply cordate at the base with a broad, open sinus, conspicuously transversely veined, scaberulous or somewhat pilose on the upper surface, rather densely and softly pilose on the lower; tendrils short-hairy; staminate flowers: panicles diffusely branched, very lax, up to 40 cm. long, pedicels capillary, 1- 2.5 mm. long, articulated near the middle or in the upper half, receptacle somewhat pilose outside, the sepals about 0.5 mm. long, corolla lobes ovate-lanceolate, about 1 mm. long, filaments and thecae equal in length or filaments a little longer; pistillate flowers not seen; fruit globose or sub-globose, blue-black at maturity, about 8 mm. long, fleshy, sparsely pilose; seed spherical, 5-6 mm. in diameter, brown, the surface rough with sharply peaked or ridged excrescences, the hilum elevated between two narrow and rounded peaks.
SICYOS Linnaeus
Delicate to coarse, herbaceous vines, scandent or prostrate, annual or perhaps longer enduring, glabrous, pubescent, or glandular-pubescent, monoecious; leaves usually petiolate, the blades thin, angular or lobate; flowers small, shallowly cup- shaped or subrotate, the corolla 5-parted, dull white, cream, yellowish or green; staminate flowers: in simple racemes or few-branched panicles, stamens united, the filaments forming a central column, the anthers sessile or nearly so, at its apex; pistillate flowers: capitate on a common peduncle; ovary ovoid to fusiform, ovule one, pendulous from the top of the locule, style slender, stigmas 3, more or less dilated, spreading, often reflexed; fruits clustered on a common peduncle, small,
DIETERLE: FLORA OF GUATEMALA 383
ovoid, 1-seeded, armed with barbed setae or unarmed, glabrous or villous or densely very short-hispidulous, pericarp thin, fibrous, indehiscent, more or less tuberculate; seeds solitary, closely enveloped by the pericarp, tumidulous or somewhat compressed, smooth, shiny, not margined, but ornamented at the apex (embryo end) with two cushionlike protuberances.
Numerous species in tropical America with three or four more in Texas. Although only four are known in Guatemala, we have treated five here, as the fifth, S. parviflorus Willd., has been collected in nearby Chiapas, Mexico, and can be expected in Guatemala. Common in disturbed areas, Sicyos species often become troublesome weeds.
Fruits setose.
Inflorescences racemose; setae numerous, slender, easily detached.... S. longisepalus.
Inflorescences paniculate; setae few, sturdy, firmly attached S. kunthii.
Fruits unarmed or setae poorly developed.
All leaves petiolate, erect, none enfolding the fruit clusters S. galeottii.
Upper leaves sessile or very short-petiolate, drooping, enfolding adjacent fruit
clusters. Staminate racemes long, much exceeding the leaves; flowers yellowish or
greenish yellow S. guatemalensis.
Staminate racemes minute, much shorter than the leaves; flowers white to pale green or cream S. parviflorus.
Sicyos galeottii Cogn. in DC. Monogr. Phan. 3: 881. 1881.
Moist forest, 1,800-3,000 m.; Huehuetenango; El Quiche; Sacatepequez; San Marcos. Southern Mexico.
Slender, herbaceous vines, small to often extensive and freely branching; leaves petiolate, the blades thinly membranaceous, mostly 3-12 (-15) cm. long, deep green above, paler beneath, angulate-ovate in outline, 3-5-angulate or lobed, with a deep basal sinus, the terminal lobe larger and longer, both blade surfaces more or less scabridulous or hispidulous, margins shallowly and distantly denticulate or subentire; tendrils usually bifid; flowers very small, white; Staminate inflorescence a slender raceme, 8-12 cm. long, exceeding the leaves, pedicels capillary, erect or spreading, 5-12 mm. long, reflexed and long-persisting after anthesis; staminate flowers: narrowly campanulate to funnelform, with a narrow subdisclike, slight thickening at the base, petals narrowly ovate, 2.5-4 mm. long, glandular on inner or both surfaces, sepals subulate, 0.5 mm. long or shorter, filament column slender, slightly broader toward the apex, thecae usually reduced in number, often to only 2, each of these borne ribbonlike on a curved lobe of connective tissue at the apex of the filament column; pistillate inflorescence delicate, few-flowered, about 15 mm. long, rarely longer than the subtending petiole; pistillate flowers: sessile or subsessile at the apex of a slender peduncle, the perianth similar to that of the staminate flowers but smaller, and sometimes lacking sepals, ovary ovoid, glabrous, style linear, stigmas 3, spreading, somewhat dilated; fruiting peduncle to about 4 cm. long, fruits ovoid, glabrous, about 6 mm. long, tan, unarmed, but bearing a few tuberculae that are sometimes tipped with reduced aculei.
FIG. 33. Sicyos galeottii. A, portion of a stem with immature pistillate inflorescences, X Vr, B, pistillate flower, x 8; C, perianth of pistillate flower laid open to display pigmented area near base, x 8; D, fruit, X 4; E, young staminate flower, x 5; F, pistillate bud, x 12!/2j G, immature stigma, X 25; H, pistillate perianth exterior, x 10.
384
DIETERLE: FLORA OF GUATEMALA 385
Sicyos guatemalens is Standl. & Steyerm. Field Mus. Bot. 23: 96. 1944.
Thickets or disturbed areas, 1,300-1,800 m.; Jalapa (type, Steyermark 32140); Zacapa. Mexico (Oaxaca, Hidalgo).
Scandent vines, the stems slender, elongate, sparsely white-villous, branched; upper leaves sessile or very short-petiolate, reduced in size, the veins pale and prominent, each blade enfolding a fruit cluster, lower leaves long-petiolate, the blades erect and spreading; leaf blades ovate-triangular to suborbicular in outline, 4-10 cm. long, more or less hispidulous or scabrous on both surfaces, 5-lobed, the lobes ovate- triangular, the terminal lobe larger and longer, often when young appearing caudate- acuminate, basal sinus rather deep, and broadly arched, leaf margins entire or shallowly denticulate; flowers yellow-green, corolla lobes ovate-triangular, obtuse, sepals minute, dentiform or subulate, or lacking; staminate flowers: shallowly cupular to subrotate, 6-10 mm. in diameter, borne in simple racemes exceeding the leaves; pistillate flowers: 2-3 mm. in diameter, nearly funnelform, clustered at the apex of short peduncles, on young vine tips or branchlets; fruits small, ovoid, 4-6 mm. long, more or less rough-tuberculate, without setae, sometimes sparsely villous, dull brown when ripe, the clusters enveloped by blade of nearest leaf.
Sicyos kunthii Cogn. in DC. Monogr. Phan. 3: 888. 1881.
Damp, brushy or thinly forested slopes, 1,200-2,450 m.; Quezaltenango; El Quiche; Solola. Mexico.
Small to extensive vines, usually scandent over shrubbery; stems sulcate, mostly glabrous; leaves on petioles 2-10 cm. long, these sulcate, glabrous, but bearing a narrow, ventral stripe of close hairs, the blades thin, ovate in outline, shallowly 5- lobed, mostly 3-10 cm. long, acuminate, the lateral lobes smaller than the central one, basal sinus deep, varying from rectangular to broadly rounded, upper surface scabrous or scabridulous, the lower surface paler, usually densely short-hisped; flowers small, 5-6 mm. across, white or greenish, with a green center; perianths of staminate and pistillate flowers alike but those of the pistillate smaller; receptacle shallowly cupular to subrotate, with a firmer, disclike (perhaps glandular) base or center; sepals minute, subulate, much shorter than the petals; petals ovate- triangular, obtuse, about 2 mm. long; staminate flowers: numerous, a few at a time in anthesis at the ends of few-branched panicles, the panicles mostly glabrous, 8-15 cm. long, much exceeding the leaves; pistillate flowers: subcapitate on slender peduncles 0.5-3 cm. long, ovaries sparsely villous and with a few setae; stigmas dilated, rounded, reflexed; fruits ovoid, 6-9 mm. long, somewhat villous, sparingly setose, the setae ascending, sturdy, broadened at the base, firmly attached, about 3.5 mm. long, densely beset with retrorse barbs on the distal half or two-thirds; seeds 4-6 mm. long.
The name Sicyos kunthii Cogn. is the one selected by Standley for this taxon of Guatemalan Cucurbits. I am continuing his usage, even though Cogniaux's description, written for Peruvian collec- tions, does not fit our material in all respects. Perhaps the Guatemalan specimens represent an undescribed species.
386 FIELDIANA: BOTANY, VOLUME 24
Sicyos longisepalus Cogn. in Donn.-Sm. Bot. Gaz. 16: 11. 1891. Quimiche; huisquil de raton.
Moist thickets or fields, frequently in hedges, a common weed in cornfields and on waste ground, often abundant, 1,200-1,800 m.; Chimaltenango; Jalapa; Sacatepequez (type from Antigua, J. D. Smith 2202); Santa Rosa.
Small or large vines, much branched, the slender branches usually densely short - villous and more or less viscid; leaves long-petiolate, the blades thin, cordate-ovate in outline, shallowly or deeply 5-lobate, the lateral lobes shorter, basal sinus broad, margins shallowly denticulate to entire, upper surfaces scabrous or short-hispid, the lower densely short-hispid and not as green; tendrils 3-5-parted; staminate flowers: inflorescence usually a simple raceme, 10-15 cm. long, much exceeding the leaves, the numerous flowers borne on the upper third or higher, a few at a time in full anthesis near the apex, perianth subrotate, petals ovate-triangular, obtuse, greenish white, green-nerved, sepals leaflike, elongate, attenuate, much longer than the petals in bud; pistillate flowers: perianth similar to that of staminate but much smaller, peduncle short, about 1.5 cm. long or less, ovary densely setose, stigmas dilated, reflexed, borne globelike on a slender style about as long as the radius of the perianth; fruits ovoid, obtuse, slightly compressed, 7-9 mm. long, dull brown when mature, villous and densely setose, the setae slender, easily detached, 4.5-5.5 (-9) mm. long, their barbs retrorse, short and uncrowded.
Sicyos parviflorus Willd. Sp. PL 4: 626, 1805, non Kunth. S. depauperatus Naud. Ann. Sci. Nat. ser. 5, 6: 23. 1866.
Not reported from Guatemala, but to be expected, as several collections have been made in nearby Chiapas, Mexico. Mexico; Ecuador.
Slender, herbaceous, annual vines 1-4 (-5) m. long, the stems branching, more or less short-villous; tendrils bifid; upper leaves sessile or very short-petiolate, drooping and enfolding the adjacent clusters of fruits, the lower leaves on petioles mostly 1-9 cm. long, the blades thin, more or less ovate-triangular in outline, 3-5-angulate or angulate-lobate, with a deep basal sinus, the central lobe triangular to ovate - triangular, the apex obtuse and apiculate; staminate flowers: inflorescences racemose, the racemes minute, shorter than the mature petioles; pistillate flowers: in short clusters less than 0.5 cm. long, receptacles broadly cup-shaped or campanulate, white to pale green or cream, 1-1.5 mm. across; fruiting peduncles 1-2 cm. long, the fruits shield-shaped or ovoid-triangular, often with a median ridge on one face, somewhat compressed, unarmed, finely and densely short-hispidulous, first cream-colored but nearly black when mature.
TECUNUMANIA Standley & Steyermark
Herbaceous, dioeceous vines, more or less scaberulous; leaves long-petiolate, simple, the blades densely villous at apex of petiole; tendrils proximally 2-4-parted; flowers yellow, moderately large, all solitary, perianths similar in staminate and pistillate flowers; receptacle tube long-campanulate, the lower half or third somewhat expanded and fleshy; sepals ligulate, much elongated; corolla deeply 5-parted;
DIETERLE: FLORA OF GUATEMALA 387
stamens 3, inserted on receptacle tube; filaments discrete but closely connivent, glabrous; anthers coherent in a subglobose head above the receptacle tube; thecae elongate- linear, vertically replicate; pistil said to be long-columnar, 3-branched in upper part; stigmas 3, bilobate; staminodia 3, ligulate; fruit rather large, subglobose, smooth, unarmed, indehiscent; seeds medium-sized, numerous, oval, compressed, not marginate, pale ochraceous, smooth, the outermost layer of seed coat made up of tightly appressed, furlike hairs.
Standley wrote: "The genus is dedicated to Tucun Uman (the name is spelled in various ways), hero and leader of the Indians of western Guatemala, who fought heroically to resist the Spanish invaders under Pedro de Alvarado, but were finally overthrown and dispersed in a bloody battle near the present city of Quezal- tenango."
Tecunumania quetzalteca Standl. & Steyerm. Field Mus. Bot. 23: 97. 1944. Melocoton; melocoton de monte.
Moist or wet, dense, mixed forest, often in ravines, 800-1,750 m.; Quezaltenango (lower slopes of Volcan de Santa Maria); San Marcos (type from Finca Vergel, near Rodeo, Standley 68931); Suchitepequez. Mexico (Chiapas); Costa Rica.
Large, herbaceous, climbing vines; stems rather stout, sulcate, glabrous, often villous at nodes; petioles (4-) 7-12 cm. long, glabrous, but densely long-villous at apex, and sometimes bearing 2 rows of closely-set villae along the ventral length; leaf blades membranaceous, 10-17 cm. long, smooth or finely scaberulous, dark green above, lighter green below, often bearing scattered, small, flat, round glands on lower surface near base and apex, margins remotely and minutely toothed, or subentire, basal sinus 2-3 cm. deep, broadly arching, bordered in part by basal lateral veins; tendrils proximally 2-3 (-4) parted, glabrous, long-pedunculate; staminate flowers: solitary, peduncle 20-25 mm. long, glabrous, receptacle tube long-campanulate, 10-14 mm. long, 8 mm. broad, glabrous, the lower half or third somewhat fleshy, sepals linear, ligulate, usually longer than the receptacle, 15-20 mm. long, about 1 mm. broad, spreading or reflexed, often bearing round, flat glands on dorsal surface of upper third, corolla deeply 5-lobed, spreading, the lobes broadly obovate, abruptly acuminate, 5-nerved, 20-25 mm. long, stamens 3, filaments free but closely connivent, 6 mm. long, 1.75 mm. broad at base, inserted on receptacle tube, anthers united in a subglobose and exserted head 8 mm. long, thecae elongate-linear, vertically replicate; pistillate flowers: solitary, perianth similar to that of staminate flower, morphological structure poorly known; fruit subglobose, indehiscent, about 7.5 cm. long, smooth, many-seeded; seeds flat, rounded at base and apex, 6-7 mm. long, 4-5 mm. broad, 2 mm. thick, ivory-colored.
In general appearance the vine suggests Luffa cylindrica, but in the latter the fruits and flowers are very different. Two of the distinctive, conspicuous characters of this cucurbit are the tufts of white hairs or wool at the apex of the petiole, and the much elongated, narrow sepals.
FIG. 34. Tecunumania quetzalteca. A, habit (section of stem with a staminate flower), x Vfc; B, apex of petiole and part of blade, underside, x l'/2 (greatly enlarged gland at left); C, staminate bud, x 2; D, stamens, x 2; E, diagrammatic sketch of stigmas, style, and staminodia, x 2'/2.
388
DIETERLE: FLORA OF GUATEMALA 389
Two species in other families, often mistaken for Cucurbitaceae, may be mentioned here:
Gronovia scandens L., a slender vine in the Loasaceae, with deeply lobed leaves, yellow flowers, and unpleasant texture, closely resembles cucurbits in general habit. It even has some of the same habitat preferences. Its bisexual flowers and lack of tendrils distinguish it readily from the Cucurbitaceae.
Tourrettia lappacea (L. 'Her.) Willd., a vine in the Bigno- niaceae, resembles cucurbits in having tendrils and spiny fruits. However, the tendrils are only modified foliar lobes, which together with the zygomorphic, bisexual, red flowers quickly set it apart from the Cucurbitaceae.
SCANNING ELECTRON MICROSCOPE PHOTOGRAPHS (Figures 35-46)
Pollen grains removed from herbarium specimens were placed on aluminum stubs by means of double-stick Scotch Tape. The stubs were then gold-coated for about 3 min. using a glow-discharge coater, and viewed with a JEOL Model JSM-U3 scanning electron microscope with an accelerating voltage of 15 KV. Secondary electron images were recorded with Polaroid Type 55 P/N 4 X 5 film.
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395
396 FIELDIANA: BOTANY, VOLUME 24
CAMPANULACEAE. Bluebell Family By DOROTHY L. NASH
Annual or perennial herbs, shrubs, or small trees; leaves alternate, estipulate; inflorescences basically cymose or the flowers solitary in the leaf axils, but often appearing racemose or paniculate; flowers perfect, epigynous, gamopetalous, usually bracteate; cleistogamous flowers sometimes produced; calyx lobes usually 5 (3-10); corolla regular and more or less campanulate or tubular and bilabiate, and sometimes cleft on one side, valvate, the lobes usually 5 (3-6); stamens as many as the corolla lobes and alternate with them, distinct or united into a tube, free or the filaments inserted at the base of the corolla; anthers bithecous, longitudinally dehiscent, introrse, distinct or connate; ovary commonly one-third or more inferior (rarely less than one-third inferior and appearing nearly superior in Diastatea and some Lobelia species), 2-5-carpellate, 2-5-locular, the placentation axile (in ours), the ovules anatropous; style simple, the stigma 2-5-lobate; fruit capsular or berrylike; seeds numerous, minute; embryo straight; endosperm carnose.
The family is treated here as consisting of two subfamilies, the Campanuloideae and the Lobelioideae, with a total of about 65 genera and more than 1,500 species, widely distributed in both hemispheres.
Corolla actinomorphic, subrotate to campanulate or campanulate-urceolate; stamens distinct (in ours) Campanuloideae.
Corolla zygomorphic, the limb more or less bilabiate (except in Hippobroma which has a salverform corolla, the tube elongated and very narrow, the subequal lobes spreading); stamens united by their anthers into a tube around the style (in ours) ....Lobelioideae.
Subfamily CAMPANULOIDEAE
Mostly annual or perennial herbs, the sap often milky; leaves alternate, simple, estipulate; flowers perfect, solitary in the leaf axils or disposed in cymes, spikes, or racemes; calyx tube or hypanthium adnate to the ovary, the limb 5-lobate (in ours), the lobes equal or nearly so, valvate or imbricate in bud, commonly persistent in fruit; corolla sympetalous, regular, subrotate to campanulate, inserted at the line where the calyx becomes free from the ovary, the tube entire or deeply cleft on one side, the limb 5-lobate (in ours); stamens 5 (in ours), alternate with the corolla lobes, the filaments free or inserted at the base of the corolla, distinct, the anthers bithecous, introrse, distinct (in ours); ovary 2-5-locular, the placentation axile (rarely unilocular and with 2 parietal placentae); ovules anatropous; style simple, the stigma 2-5-lobate; fruit capsular; seeds numerous, minute.
Only one of the 35 or more genera, Triodanis, is native in Central America; the pantropical Sphenoclea, introduced long ago, is now fairly widespread, and Campanula is widely planted, with one species sometimes naturalized.
NASH: FLORA OF GUATEMALA 397
Corolla subrotate; earliest flowers cleistogamous Triodanis.
Corolla campanulate to campanulate-urceolate; flowers commonly all chasmo-
gamous.
Flowers crowded in a dense, many-flowered, cylindrical spike, the corollas very small, only 2-3 mm. broad, campanulate-urceolate; capsule circumscissile.
Sphenoclea.
Flowers solitary in the upper leaf axils or disposed in cymes, the inflorescence sometimes appearing racemose or spicate but never crowded or cylindrical, the corollas relatively large, commonly 1-4 cm. broad, campanulate; capsule variously dehiscent by valves or perforations but never circumscissile.
Campanula.
CAMPANULA Linnaeus
Perennial, biennial, or annual herbs, pubescent or glabrous, simple or branching; leaves alternate; inflorescence basically cymose but sometimes appearing racemose, spicate, or the flowers solitary in the leaf axils; flowers conspicuous, often rather large; hypanthium hemispheric to turbinate; calyx deeply parted, the 5 segments linear to triangular or lanceolate, often acuminate; corolla commonly blue, violet, or white, usually broadly campanulate, 5-lobate, valvate in bud, the flowers normally all chasmogamous, rarely partly cleistogamous; stamens 5, alternate with corolla lobes, the filaments abruptly or gradually dilated at the ba^e, inserted at the base of the corolla, the anthers distinct; ovary 3-5-locular, the stigma 3-5-lobate; capsule wholly or partly inferior, variously dehiscent by lateral valves or by 3-5 small valves or perforations at the base, near the middle, or near the apex.
A large genus of perhaps 250 species, widely distributed in the northern hemisphere, mostly in Europe and Asia. One species is apparently native in Mexico but none extend to Central America. Two are treated here as they are frequently cultivated in Guatemala.
Calyx lobes hispid, aunculate-appendaged at the base; corolla lobes short, usually less than one-fourth the length of the entire corolla; widely cultivated but not naturalized C. medium.
Calyx lobes scabrous, not appendaged; corolla lobes one-third to one-half the length of the corolla; sometimes naturalized C. rapunculoides.
Campanula medium L. Sp. PI. 167. 1753. Campana; Can- terbury bells.
Native of southern Europe; cultivated for ornament in many parts of the world, especially in temperate regions; often in cultivation in Guatemala.
Erect biennials, stout, mostly 30-60 cm. tall, pilose or hispid; leaves sessile and more or less clasping, lanceolate or ovate-lanceolate, mostly 3-7 cm. long, the margins crenate-dentate; flowers solitary in the leaf axils, pedicellate, the area of inflorescence sometimes appearing laxly racemose; calyx 1.5-3 cm. long, the lobes foliaceous, ovate to lanceolate, more or less hispid, auriculate-appendaged; corolla
398 FIELDIANA: BOTANY, VOLUME 24
commonly blue to purple, sometimes white, rarely pink, broadly campanulate, 4-5 cm. long, the broad lobes less than 1 cm. long; capsule dehiscent at base.
Often sold in markets, especially those of Quezaltenango and Guatemala.
Campanula rapunculoides L. Sp. PI. 165. 1753. Campanola; bellf lower. ( Fig. 48, right. )
Native of Europe, sometimes planted for ornament in other regions; often naturalized in temperate North America; frequently planted in gardens in the uplands of Guatemala and sometimes naturalized, especially along roadsides.
Rather stout perennials, sparsely branched, usually less than 1 m. tall, thinly pubescent or glabrate; basal leaves ovate to reniform, petiolate, cordate at the base, the margins coarsely and irregularly serrate; cauline leaves mostly 6-12 cm. long, ovate-lanceolate or lanceolate, acute or acuminate, narrowed at the base, short- petiolate or the upper ones sessile; flowers nutant, pedicellate, solitary in the leaf axils, the flowering stem appearing as a terminal, leafy-bracteate raceme; calyx lobes linear; corolla blue or purple, 2.5-3.5 cm. long, the lobes mostly 1-1.5 cm. long; capsule opening by basal pores.
SPHENOCLEA Gaertner
Erect, glabrous annuals, the stems somewhat succulent; leaves alternate, short - petiolate, estipulate, the blades usually narrow, the margins entire; inflorescence a densely-crowded, elongated, terminal spike; flowers perfect, sessile, subtended by a bract and bracteoles; calyx hemispheric, the tube adnate to the ovary, the limb 5- lobate, the segments rounded, imbricate in bud; corolla campanulate-urceolate, the 5 lobes imbricate; stamens 5, alternate with the corolla lobes, the short filaments inserted at about the middle of the short corolla tube, the anthers bithecous, longitudinally dehiscent; ovary semi-inferior, bilocular, the placentation axile, the ovules numerous; style short, the stigma capitate; capsule more or less obconic, bilocular, circumscissile; seeds numerous, minute; embryo straight.
Airy-Shaw (Flora Malesiana, ser. 1, 4(1): 27. 1948, and in Willis, A Dictionary of the Flowering Plants and Ferns, 7th ed.: 1058. 1966) maintains Sphenocleaceae as a separate family. However, Hut- chinson (The Families of Flowering Plants, ed. 2, 1: 476. 1959) retains Sphenoclea in the Campanulaceae, as have others. Although superficially Sphenoclea does not resemble other genera in the family, there seems to be no real evidence, anatomical or otherwise, to justify its separation.
Two species have been reported, the pantropical one treated here, and one said to be endemic in West Africa.
Sphenoclea zeylonica Gaertn. Fruct & Sem. 1: 113. 1788.
NASH: FLORA OF GUATEMALA
399
FIG. 47. Sphenoclea zeylonica. A, habit, x Vfe; B, detail of inflorescence, X 3; C, flower opened to show stamens and pistil, x 5; D, capsule, opened, and cross-section of immature ovary, X 5.
In wet fields, marshes, or ditches, sometimes in Manicaria swamps, sea level to 800 m.; Izabal; Jutiapa; Peten; San Marcos. Southern United States; Mexico; British Honduras to Panama; West Indies; South America; Old World Tropics.
Erect, usually branching, somewhat glaucous plants, glabrous throughout, mostly 20-100 cm. tall, the stems hollow, somewhat succulent, often thickened at the
400 FIELDIANA: BOTANY, VOLUME 24
base; leaves short-petiolate, the blades oblong-lanceolate to elliptic, mostly 4-12 cm. long, 1-3 cm. broad, acute or obtuse, acute or attenuate to the base; flowering spike cylindrical, very densely flowered, mostly 3-10 cm. long, in fruit about 1 cm. thick; calyx segments broadly triangular to ovate, rounded; corolla white or greenish white, 2.5-4 mm. long, 2-3 mm. broad, caducous, the 5 lobes broad, ovate-triangular, obtuse or acute; stamens about as long as the corolla lobes, subsessile, the short filaments slightly dilated at base; capsule 3-5 mm. in diameter, circumscissile; seeds light brown, rugose.
TRIODANIS Rafinesque
Reference: Rogers McVaugh, The genus Triodanis Rafinesque and its relationships to Specularia and Campanula, Wrightia 1(1): 13-52. 1945.
Erect or reclining annuals, simple or branched, the leafy stems more or less 5- angulate, usually pubescent on the angles; leaves alternate; flowers sessile or nearly so, bracteolate, in the axils of the middle and upper leaves (bracts), the lower flowers commonly cleistogamous and much smaller than the chasmogamous ones; hypan- thium narrow, the calyx lobes 5 or 4 in chasmogamous flowers, 3-5 in cleistogamous ones; corolla subrotate, the 5 lobes abruptly acuminate, imbricate in bud, 2-3 times longer than the tube; stamens 5, the filaments abruptly dilated and ciliate at the base, the linear anthers distinct, longer than the filaments; ovary of chasmogamous flowers commonly trilocular, but bilocular or trilocular in cleistogamous flowers (rarely unilocular), the placentation axile (in ours); style more or less pubescent, the stigmatic branches equalling the number of locules in the ovary; capsule linear, oblong, or ellipsoidal to obovoid or clavate, with commonly 3 openings near the apex or about the middle; seeds numerous, minute, ovoid, oblong or lenticular.
Seven of the eight species are American, the other chiefly Mediterranean. Only one is found in Central America.
Triodanis perfoliata (L.) Nieuwl. Am. Midi. Nat. 3: 192. 1914. Campanula perfoliata L. Sp. PL 169. 1753. Specularia perfoliata A. DC. Monogr. Campan. 351. 1830.
Dry hillsides, open wet banks, or a weed in or near cornfields, 2,200-2,500 m.; Quezaltenango. Widely distributed in the United States (most common in the east); British Columbia to southern Ontario; southern Mexico; Jamaica; Hispaniola; Ecuador.
Erect or sometimes decumbent plants, mostly 10-70 cm. tall, the stems simple or sparsely branched, densely leafy, somewhat pubescent near the base or retro rse- hispidulous on the angles or almost glabrous; upper and median leaves sessile, caudate-clasping, the lower ones sometimes cuneate-clasping, but the lowermost leaves often petiolate, the lower leaf blades orbicular or ovate to elliptic or obovate, rounded or obtuse at the apex, mostly 0.5-3.5 cm. long, often undulate, the margins more or less crenate-dentate, ciliate, glabrous above, more or less scabrous beneath or the hairs confined to costae and veins, The upper leaf blades (bracts) similar but relatively broader, ovate to reniform, to 2.5 cm. long and about 3 cm. wide, obtuse at
NASH: FLORA OF GUATEMALA
401
FIG. 48. (Left) Triodanis perfoliata. A, habit, x Vr, B, habit showing flowers, from a photograph, X 2. (Right) Campanula rapunculoides . C, habit, X Vz; D, flowering stem, X 1; E, cross-section of ovary, X 3; F, seed, much enlarged; G, mature capsule showing method of dehiscence, X 2. (D-G, after Hegi, Flora von Mittel Europa.)
the apex or acute and mucronate; chasmogamous flowers 1-3 (-8), sessile in the upper leaf axils, cleistogamous flowers usually in the lower axils; calyx lobes 4-5 (3-5 in cleistogamous flowers), triangular to lanceolate or ovate, acuminate, the midrib prominent, glabrous, sometimes ciliate; corollas blue, violet, or rarely white, those of the open flowers 8-13 mm. long, those of the cleistogamous ones less than 0.5 mm. long; stamens included; style pubescent; capsule oblong or obovoid, about 10 mm. long in the open flowers, 4-6 (-8) mm. long in the cleistogamous flowers, opening at about the middle; seeds minute, lustrous.
Subfamily LOBELIOIDEAE
Mostly annual or perennial herbs, the sap usually milky; leaves alternate, cauline or basal, estipulate, inflorescences often appearing racemose, the flowers solitary in the axils of the upper leaves or bracts; pedicels usually bracteate; calyx tube or hypanthium adnate to the ovary, usually attached well beyond the base of the ovary (rarely attached only to the base, in Diastatea), commonly persistent in fruit, the limb 5-lobate; flowers perfect; corolla sympetalous, withering-persistent in fruit, inserted at the line where the calyx becomes free from the ovary, the tube entire or cleft on one side and often laterally fenestrate, the limb commonly more or less bilabiate (rarely the corolla salverform with a long, narrow tube and spreading, subequal lobes in Hippobroma); stamens 5, alternate with the corolla lobes, the filaments flattened, free or loosely attached to the corolla, united into a tube or distinct at the base, the anthers bithecous, introrse, united into a tube around the style (in ours); ovary usually bilocular, the placentation axile (in ours); ovules
402 FIELDIANA: BOTANY, VOLUME 24
anatropous; style one, the stigma bilobate; fruit capsuler or berrylike; seeds numerous, minute (in ours).
About 30 genera, with seven in Guatemala. Except for the genus Lobelia, the family is much better represented in Costa Rica and Panama.
Hypanthium irregular, the lower side forming a conic or linear spur; corolla tube
prolonged on the lower side into a spur Heterotoma.
Hypanthium essentially regular, not spurred; corolla tube not prolonged into a spur. Corolla salverform, the narrow tube much elongated, 5-13.5 cm. long.
Hippobroma.
Corolla never salverform, the tube never more than 4 cm. long. Ovary nearly superior, the hypanthium attached only to the base of the ovary; corolla much distended by the maturing capsule and finally becoming
hyaline Diastatea.
Ovary partially or wholly inferior, the hypanthium always attached beyond the
base of the ovary; corolla not or scarcely distended, not becoming hyaline. Corolla tube cleft dorsally.
Style jointed above the ovary and deciduous with corolla and stamens from
the fruit; fruit berrylike, somewhat fleshy, indehiscent Pratia.
Style not jointed above the ovary, persistent with corolla and stamens on
the fruit; fruit capsular, dehiscent by apical valves Lobelia.
Corolla tube entire.
Corolla green, the tube usually broadest at the base; anther tube obliquely enlarged and open at the top; style jointed above the ovary and
deciduous with corolla and stamens from the fruit Burmeistera.
Corolla red to orange (in ours), the tube usually narrow at the base; anther tube not enlarged, partially closed at the top; style not jointed, persistent with corolla and stamens on the fruit Centropogon.
BURMEISTERA Triana
Herbaceous or somewhat suffruticose plants, terrestrial or sometimes epiphytic; leaves cauline, alternate; flowers borne in the upper leaf axils, the floral leaves usually little reduced, the pedicels ebracteolate; calyx adnate to the ovary, 5-lobate, the segments subequal; corolla epigynous, green, yellow, or purplish to reddish brown, the tube entire, cylindric, usually broadest at the base and somewhat narrowed toward the base of the lower lobes, there abruptly dilated, the limb obliquely 5- lobate, the lobes falcate, turned downward away from the dorsal side of the flower, the 2 upper lobes much longer than the others, the ventral one shorter than the lateral ones; stamens 5, the filaments distinct only at the base, adherent to the corolla or sometimes essentially free from it, the anther tube usually abruptly curved at the base, obliquely open at the apex, in age the anther tips separating and curving outward, the 2 shorter anthers not barbate at the apex but all the anthers or sometimes only the 2 shorter ones often sparsely pilose at the apex; ovary inferior, bilocular; style jointed just above the ovary, deciduous with the corolla and stamens from the fruit; fruit fleshy, indehiscent, nearly globose or somewhat elongated, frequently inflated at maturity; seeds numerous, small, oblong to ovoid or fusiform, minutely foveolate-reticulate.
NASH: FLORA OF GUATEMALA 403
More than 70 species have been reported from the mountains of tropical America. The genus apparently reaches its northern limit with a single species in Guatemala. At least nine other species are known from Costa Rica and Panama.
Burmeistera virescens (Benth.) Benth. & Hook, ex Hemsl. Biol. Centr. Am. Bot. 2: 263. 1881. Lobelia virescens Benth. PL Hartweg. 77. 1841. Centropogon virescens Planch. & Oerst. Vidensk. Meddel. 157. 1857. C. lignescens Wimmer in Fedde, Rep. Sp. Nov. 19: 252. 1924 (type from Purulha, Baja Verapaz, Tuerckheim II. 1743). B. lignescens Wimmer, op. cit. 30: 34. 1932. B. lignescens var. martialis Wimmer, op. cit. 30: 34. t. 126, f. 53. 1932 (type from Finca Sepacuite, Alta Verapaz, Cook & Gilbert 234). B. virescens var. subdentata Wimmer, Pflanzenr. 106(4), Fam. 276b: 135. 1943 (type from Chihob, Alta Verapaz, Harry Johnson 916).
Dense, wet, mixed forest, 900-2,500 m.; Alta Verapaz; Baja Verapaz; Huehuetenango; Quezaltenango (type from Rancho de Santa Rita, near Quezaltenango, Hartweg 540); San Marcos; Suchitepequez.
Terrestrial plants, erect, suffruticose or coarsely herbaceous, the stems usually glabrous, sometimes puberulent, to about 1.5 m. tall; leaves on petioles mostly 1-3 cm. long, the blades thin, oblong-elliptic to ovate, mostly 7-20 cm. long, acuminate to long-acuminate, subacute or attenuate to the base, the margins subentire to coarsely and irregularly laciniate-dentate, usually glabrous above or nearly so, almost glabrous beneath or scaberulous to strigose on the costae and veins; area of inflorescence mostly 10-16 cm. long, the pedicels in fruit 3-8 cm. long, glabrous; hypanthium glabrous, obconic in anthesis, The calyx lobes narrowly triangular, 3.5-9 mm. long, obtuse, essentially entire or very obscurely dentate; corolla green or rarely greenish tinged with pink, glabrous or minutely puberulent, about 3 cm. long, the lobes acute or subacute; filament tube 2-2.5 cm. long, the anther tube 4-6 mm. long; fruit berry like, scarcely inflated, white or pink, 7-10 mm. long; seeds numerous, oblong, somewhat flattened or centrally depressed, about 1 mm. long.
CENTROPOGON Presl
Herbaceous or suffrutescent plants; leaves cauline, alternate; flowers long- pedicellate, borne singly in the axils of the uppermost leaves or bracts; calyx tube adnate to the ovary, the limb 5-lobate, the segments subequal, persistent; corolla tubular, usually red or orange, varying to pink or yellowish, the tube straight or curved, usually entire, contracted just above the base and gradually dilated upward; filaments distinct at the base but adherent to the corolla above its narrow basal portion, the anther tube exserted, the orifice partially closed by the incurved tips of the longer anthers, the 2 shorter anthers with tufts of stiff, white hairs at the apex (in ours); style included in the staminal tube, not jointed, persistent with the corolla on the fruit; ovary inferior, bilocular, the summit flat or nearly so; fruit fleshy, globose or subglobose, often inflated at maturity, indehiscent; seeds minute, broadly ovoid, minutely foveolate-reticulate.
FIG. 49. Burmeistera virescens. A, branch and inflorescence, x Vz; B, leaves and stem of another plant, X W, C, flower, x iVr, D, flower dissected, x 1V&; E, anther column and stigma, x 3Vr, F, fruit, x 2; G, seeds, x 10.
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NASH: FLORA OF GUATEMALA 405
A very large genus with perhaps 200 species, chiefly in mountain regions of tropical America. Only three are known in Guatemala.
Leaf blades cordate or broadly ovate, cordate or subcordate at the base.
C. cordifohus.
Leaf blades elliptic, elliptic-oblong, or ovate-oblong (rarely ovate), acute or obtuse at
the base or sometimes attenuate to the base. Indument of lower leaf surfaces and inflorescences consisting of conspicuously
branched hairs C~ f™&*"
Indument of lower leaf surfaces and inflorescences consisting of minute, crisped , . C. grandidentatus.
Centropogon cordifolius Benth. PI. Hartweg. 77. 1841 (type from Palo Hueco, Guatemala, Hartweg 539). C. cordatus Mart. & Gal. Bull. Acad. Brux. 9(2): 40. 1842. Siphocampylus guatemalensis Vatke, Linnaea 38: 730. 1874. Lechera (Quezaltenango); platanillo (San Marcos).
Damp or wet, mixed forest and thickets, sometimes in pine and Liquidambar forest, 900-2,500 m.; Alta Verapaz; Baja Verapaz; Chimaltenango; Chiquimula; Huehuetenango; Quezaltenango; El Quiche; San Marcos; Zacapa. Southern Mexico.
Herbaceous or suffrutescent plants, the weak stems usually suberect, sometimes decumbent, branching, often dark red, glabrate; leaves on petioles mostly 2-4 cm. long the blades broadly ovate to cordate, mostly 5-14 cm. long, acute or short- acuminate with an obtuse or sometimes minutely apiculate tip, cordate or subcordate at the base, glabrous above or nearly so, paler beneath and puberulent, at least on the veins, the margins irregularly repand-dentate or remotely denticulate; pedicels mostly 3-8 cm. long, bibracteolate; hypanthium hemispheric in anthesis, sparsely puberulent, enlarging in fruit to about 10 mm. in diameter, the calyx lobes linear or subulate, obtuse at the tip, 5-13 mm. long, puberulent, usually somewhat denticulate, often reflexing, corolla bright, deep red or rose, rarely pink, mostly 3-4.5 cm. ong, minutely puberulent outside, the lobes erect or only slightly recurved, 6-8 mm. long, the 2 upper lobes linear to elliptic, the others triangular; filament tube 30-35 mm. long, the anther tube 5-6.5 mm. long; fruit dark, purplish red, little inflated; seeds about 0.6 mm. long.
Centropogon ferrugineus (L. f.) Gleason, Bull. Torrey Bot. Club 52: 11. 1925. Lobelia ferruginea L. f. Suppl. PI. 394. 1781. C. costaricanus Planch. & Oerst. Vidensk. Meddel. 1857: 156. 1857. C. affinis var. costaricanus Zahlbr. Ann. Naturh. Mus. Wien 6: 437. 1891. C. poasensis Gleason, Torreya 25: 92. 1925. C. costaricanus var. tomentellus Wimmer, Ann. Naturh. Mus. Wien 46: 240. 1933. C. ferrugineus var. costaricanus McVaugh, Ann. Mo. Bot. Gard. 27: 352. 1940. C. ferrugineus var. tomentellus McVaugh, N. Am. Fl. 32A, pt. 1: 126. 1943. Lechoso (Jalapa).
FIG. 50. Centropogon cordifolius. A, habit, X Vfc; B, flower from side, X iVfc; C, flower dissected, X IW, D, two views of staminal column and stigma, x 3V2.
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NASH: FLORA OF GUATEMALA 407
Wet thickets or mixed forest, 2,000-3,000 m.; Jalapa; El Progreso. El Salvador; Costa Rica; Panama; southward to Ecuador and Venezuela.
Herbaceous of suffrutescent plants, the stems simple or branching, erect or straggling, 0.5-3 m. tall, tomentose with branched, brownish hairs, becoming glabrate, often purplish; leaves short-petiolate, the petioles more or less winged, at least on one side, and densely puberulent, the blades elliptic-oblong to ovate- oblong, mostly 6-13 cm. long, acute or abruptly short-acuminate, acute or obtuse at base, the margins serrate-denticulate, puberulent or glabrate above, densely tomentose beneath at first with branching, light or dark brown hairs later the indument much less, often largely confined to costae and veins; flowers on pedicels mostly 4-8 cm. long; hypanthium in anthesis hemispheric, rather densely pubescent, in fruit 7-12 mm. broad, the calyx lobes narrowly triangular, usually purple, denticulate, acute, pubescent, 2.5-4 mm. long; corolla orange-red, 4-5 cm. long, somewhat pubescent outside, the lobes falcate, acuminate, the 2 upper ones 5-12 mm. long, the others shorter; fruit scarcely inflated; seeds about 0.5 mm. long.
The typical form of the species is South American. McVaugh refers Central American material to three varieties separated by color of indument, said to be either light yellow-brown, light gray- brown, or rich, dark brown, but it is difficult to detect the alleged differences in herbarium material.
Centropogon grandidentatus (Schlecht.) Zahlbr. Ann. Na- turh. Mus. Wien 6: 439. 1891. Lobelia grandidentata Schlecht. Linnaea 9: 262. 1834. C. affine Mart. & Gal. Bull. Acad. Brux. 9, pt. 2: 40. 1842. Siphocampylus regeli var. umbrosus Vatke, Linnaea 38: 733. 1874. C. grandidentatus f. incisa Wimmer in Fedde Repert, Sp. Nov. 19: 244. 1924. C. grandidentatus var. diversidens Wimmer, I.e.; Siphocampylus oaxacanus Wimmer, torn. cit. 260. C. oaxacana Wimmer, op. cit. 22: 202. 1926. Arete, poc-el (Huehuetenango); chilera (San Marcos).
Damp or wet, mixed forest, often in Cupressus forest, sometimes in pine-oak forest, 1,800-3,800 m.; Chimaltenango; Huehuetenango; Quezaltenango; El Quiche; San Marcos; Solola. Southern Mexico; Honduras; El Salvador; Costa Rica; Panama; Colombia; Venezuela.
Herbaceous or suffrutescent plants, simple or sparsely branched, commonly about 1 m. tall, rarely to 2.5 or 3 m., erect or decumbent, the young growth sparsely or densely tomentulose with sordid or brownish hairs; leaves petiolate, the petioles more or less winged, the blades thin, elliptic to oblong or rarely ovate, mostly 6-20 cm. long, short-acuminate to long-acuminate, acute, obtuse, or attenuate to the base, the margins subentire or finely glandular-serrate to conspicuously and irregularly incised-serrate, sparsely puberulent above, thinly puberulent beneath, or only along costae and veins, the hairs minute, crisped; flowers on pedicels mostly 3-10 cm. long;
408 FIELDIANA: BOTANY, VOLUME 24
hypanthium puberulent, in anthesis hemispheric, the calyx lobes triangular to narrowly triangular, acute to long-acuminate, 2-4 mm. long, usually sparsely callose- denticulate; corolla 3.5-5 cm. long, sparsely puberulent outside, rose-red or pale red to orange or salmon, or the lobes yellow or paler than the tube, the lobes recurved- falcate, acuminate, the 2 upper ones longer, to 14 mm. long; filament tube 4-4.5 cm. long, usually densely hirsute, the anther tube 6.5-8 mm. long, purplish or bluish gray, the tips of the 2 shorter anthers densely white-tufted; fruit little inflated, mostly 7-10 mm. in diameter; seeds 0.5-0.7 mm. long.
DIASTATEA Scheidweiler
Small, slender, erect annuals, simple or branched; leaves cauline, alternate, often numerous; inflorescences racemose, lax, the pedicels in the axils of the upper leaves or bracts, more or less secund; hypanthium in anthesis commonly broadly cupular, varying to almost flat or obconic, in fruit unchanged or becoming turbinate, the calyx lobes 5; corolla glabrous, the tube narrowly cylindric, not dorsally cleft, dilated by the expanding capsule, finally becomine scarious and hyaline, the 3 lower lobes united to form a distinct lip, this deflexed and bituberculate at the base; filaments about as long as or slightly exceeding the corolla tube, connate distally for part of their length, distinct at the base, the anther tube bluish gray, 2 of the anthers shorter than the others and minutely white-barbate at the apex, the 3 longer ones usually glabrous; ovary nearly superior, the hypanthium attached only to the base; capsule bilocular, dehiscent by apical valves; seeds numerous, light brown, ellipsoidal, smooth, lustrous, 0.5-0.6 mm. long.
Five or six species in Mexico and Central America with one extending southward to Bolivia; three are known in Guatemala.
Middle cauline leaves narrowly linear, 1-2.5 mm. wide; calyx lobes mostly 1-2 mm.
long; filaments of stamens 6-7.5 mm. long D. tenera.
Middle cauline leaves linear-lanceolate to ovate, at least 3 mm. wide; calyx lobes
mostly 2-6 mm. long; filaments of stamens 3-5 mm. long. Pedicels minutely scabrous; calyx lobes narrowly triangular, 0.5-1 mm. wide at the
base, conspicuously scabrous-cilia te D. costaricensis.
Pedicels glabrous or rarely with some scattered pubescence on lower part; calyx lobes linear, rarely as much as 0.5 mm. wide at the base, eciliate or remotely ciliate D. micrantha.
Diastatea costaricensis McVaugh, Bull. Torrey Bot. Club 67: 789. 1940. Laurentia irazuensis Wimmer, Field Mus. Bot. 18: 1415. 1938 (non Lobelia irasuensis Planch. & Oerst. Vidensk. Meddel. 153. 1857).
Damp, brushy, often rocky slopes or meadows, sometimes in oak or pine-oak forest, 900-1,900 m.; Alta Verapaz; Guatemala; Huehuetenango; Jalapa; Santa Rosa. Honduras; Costa Rica.
Erect plants, mostly 10-35 cm. tall, the stems simple or branched, usually angled, often winged and more or less hispidulous on the angles; leaves sessile or subsessile, the middle and upper blades linear-lanceolate to lanceolate, acute to acuminate,
NASH: FLORA OF GUATEMALA 409
cuneate at the base, mostly 1-3 cm. long, the margins sharply and often finely serrate, usually more or less pubescent, or the indument confined to the margins and the costae and veins beneath, the lower leaf blades smaller but relatively broader than the others; racemes to 15 cm. long, few-many-flowered, the pedicels minutely scabrous, mostly 0.3-1.5 cm. long; hypanthium glabrous, the calyx lobes narrowly triangular, 3-6 mm. long, 0.5-1 mm. wide at base, conspicuously scabrous-ciliate; corolla blue, lilac, or purple, 5-7 mm. long; filaments 3-4 mm. long, the anther tube about 1 mm. long; capsule broadly ellipsoidal, 2-3 mm. in diameter, 4-6.5 mm. long.
Similar in appearance to, and often confused with, Lobelia xalapensis HBK., which has an ovary commonly less than one- third inferior; however, at least the lower leaves of the Lobelia are petiolate, and the blades are ovate to lance-ovate with sinuate or irregularly and coarsely dentate margins.
Diastatea micrantha (HBK.) McVaugh, Bull. Torrey Bot. Club 67: 143. 1940. Lobelia micrantha HBK. Nov. Gen. & Sp. 3: 316. 1819. L. ruderalis Willd. ex Roem. & Schult. Syst. Veg. 5: 56. 1819. L. tuerckheimii Vatke ex Robins., Proc. Am. Acad. 26: 167, as synonym. 1891 (Antigua, Donn.-Sm. 2106). Laurentia ovatifolia Robins, torn. cit. 166. L. michoacana Robins, torn. cit. 167. L. pedunculata Brandg. Univ. Calif. Publ. Bot. 6: 73. 1914. L. micrantha var. longibracteata Wimmer, Rev. Sudam. Bot. 2: 104. 1935. D. serrata Standl. & L. Wms., Ceiba 1: 91. 1950. D. micrantha var. ovatifolia Wimmer, Pflanznr. IV (276c): 853. 1968. Santa jovita (Chimaltenango); te silvestre (Guatemala).
Damp or wet, open forest, thickets, and brushy slopes, often in pine, pine-oak, or oak forest, sometimes on sandbars of streams, in rock crevices, and occasionally a weed in cornfields and gardens, 900-3,800 m.; Alta Verapaz; Chimaltenango; Chiquimula; Es- cuintla; Guatemala; Huehuetenango; Jalapa; Jutiapa; Quezal- tenango; El Quiche; Sacatepequez; San Marcos; Santa Rosa; Solola; Totonicapan; Zacapa. Central and southern Mexico; Honduras; Costa Rica; Panama; western South America.
Erect plants, the stems simple or branching, mostly 10-50 cm. tall, hispidulous below or throughout or nearly glabrous; leaves mostly short-petiolate, sometimes sessile, the blades mostly 1-6 cm. long, the middle and upper leaves lanceolate to ovate, acute or acuminate, cuneate or attenuate to the base or sometimes nearly rounded and then abruptly decurrent on the petiole, the lower ones smaller, ovate to orbicular, the margins commonly serrate, sometimes sinuate-dentate, more or less pubescent along the margins and beneath on the veins, or sometimes glabrous; racemes with few to 30 flowers, mostly 3-25 cm. long, the pedicels 6-25 mm. long, glabrous; hypanthium in anthesis cupular; usually glabrous, in fruit becoming conic, 0.5-1 mm. long, the calyx lobes linear, 1.5-3.5 mm. long; corolla blue or purplish, rarely white, 4.5-6.5 mm. long, the tube 2.5-4 mm. long; capsule narrowly ellipsoid, 3-6 mm. long.
FIG. 51. Diastatea micrantha. A, habit, x Vz; B, detail of stem and leaves, x 2; C, fruiting inflorescence, X 2; D, flower, x 7; E, flower dissected, x 10; F, mature capsule, x 8; G, partially dissected capsule, X 5, with seeds, greatly enlarged.
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NASH: FLORA OF GUATEMALA 411
Diastatea tenera (Gray) McVaugh, Bull. Torrey Bot. Club 67: 143. 1940. Palmerella tenera Gray, Proc. Am. Acad. 22: 433. 1887. Lobelia palmeri Greene, Pittonia 1: 297. 1889. Laurentia pinetorum Brandg. Univ. Calif. Publ. Bot. 4: 92. 1910.
Zacapa, 1,100-1,900 m. (along Rillito del Volcan de los Monos, Sierra de las Minas, Steyermark 42316). Western and southern Mexico.
Erect herbs, the stems simple or sparsely branched, mostly 8-20 cm. tall, glabrous or minutely scabrous- puberulent; leaves glabrous, sessile or nearly so, the middle and upper ones narrowly linear, acute at each end, 0.5-3.5 cm. long, 0.1-0.25 cm. wide, finely and shallowly serrate, the lower leaves smaller and relatively broader, obtuse or acute; inflorescences few-flowered, the pedicels 6-20 mm. long, glabrous; hypanthium glabrous, conic or turbinate in fruit, the calyx lobes linear to elliptic or lanceolate, acute, glabrous, with 1-2 minute teeth on each side; corolla purplish blue, 10-16 mm. long, the tube 5-7.5 mm. long; anther tube 1-1.3 mm. long; capsule 5-7 mm. long.
HETEROTOMA Zuccarini
Annual or perennial herbs, the stems erect, decumbent or repent, simple or branching; leaves alternate, the blades dentate or incised; inflorescences appearing pedunculate, the flowers pedicellate, the pedicels bracteate; calyx limb 5-lobate, the hypanthium irregular, more or less unilateral, the lower part bearing 2 calyx lobes opposite the 2 shorter anthers and forming a conic or linear spur, the calyx lobes on the spur often shorter than the other 3; corolla zygomorphic, the tube prolonged on the under side to form the spur, dorsally cleft to the base or for about two-thirds of its length; stamens 5, the filaments free from the corolla, connate distally, distinct at the base, the 2 anterior ones more or less extended into the spur, the tips of the 2 smaller anthers densely barbellate, the 3 longer ones curved apically; capsule bilocular, apparently dehiscent by apical valves, enclosed by the hypanthium tube on the upper side for two-thirds to three-fourths its length, but little or not at all on the lower side; seeds numerous, minute, ellipsoidal, smooth and shining.
Seven species, in Mexico and Central America. Although only one, H. lobeliodes Zucc., is known from Guatemala, two are treated here. H. cordifolia (Hook. & Arn.) McVaugh, ranging from Mexico (although not yet reported from Chiapas) to Costa Rica, may be expected in Guatemala.
Leaves mostly 1-4 cm. long; pedicels secund; corolla white, pale blue, or lavender, the spur commonly less than 1 mm. long H. cordifolia.
Leaves mostly 6-15 cm. long; pedicels not secund; corolla red and yellow, the spur 14- 24 mm. long H. lobelioides.
Heterotoma cordifolia (Hook. & Arn.) McVaugh, Bull. Torrey Bot. Club 67:143.1940. Lobelia cordifolia Hook. & Arn. Bot. Beechey Voy. 301. 1838. H. tenella Turcz. Bull. Soc. Nat. Mosc.
412 FIELDIANA: BOTANY, VOLUME 24
25(2), No. 3: 175. 1852. H. cordifolia var. tenella Wimmer, Pflanzenr. IV. 276b (107. Heft): 717. 1953.
Not reported from Guatemala but to be expected there. Mexico; El Salvador; Costa Rica.
Erect herbs, simple or branched, mostly 10-50 cm. tall, the stems pubescent or nearly glabrous; leaves on narrowly winged petioles 0.5-3 cm. long, the blades ovate to reniform, acute to obtuse, rounded to cordate at base, mostly 1-4 cm. long, 1-3 cm. wide, usually simple, sometimes palmately lobed or cleft, the margins irregularly serrate or crenate to subentire, sparsely pubescent or strigose or essentially glabrous above, more or less pubescent to puberulent below, the indument sometimes confined to costae and veins; area of inflorescence 3-25 cm. long, appearing pedunculate, the pedicels secund, mostly 1-3 cm. long; bracts narrowly elliptic or linear to filiform, entire or the lower ones incised; hypanthium in anthesis ellipsoidal, the spur conic, usually less than 1 mm. long, the calyx lobes narrowly triangular to linear, entire, ciliate or not, 1.5-4 mm. long; corolla white, pale blue, or lavender, 4-8 mm. long, glabrous, the tube cleft dorsally almost to the base, otherwise entire, the spur usually less than 1 mm. long, the 2 upper lobes linear to narrowly elliptic, the 3 lobes forming the lower lip elliptic to obovate; staminal column 3.5-5 mm. long; capsule about three-fourths inferior, glabrous, 3.5-6 mm. long; seeds minute, smooth and shining.
Heterotoma lobelioides Zucc. Flora 15(2): Beibl. 101. 1832. Lobelia calcarata Bertol. Fl. Guat. 409. 1840 (type from San Cristobal, probably Escuintla, Velasquez s.n.).
Damp or wet thickets or brushy banks, often in rocky ravines, 1,200-2,800 m.; Chimaltenango; Guatemala; Quezaltenango; Sacate- pequez; San Marcos; Suchitepequez. Central and southern Mexico; Honduras; El Salvador; Nicaragua; Costa Rica.
Erect, annual or perennial herbs, mostly 1-2 m. tall, the stem simple or branched, pubescent below or throughout; leaves alternate, long-petiolate, the blades thin, ovate to lance-ovate, mostly 6-15 cm. long, long-acuminate, rounded or cordate at the base, often densely pubescent beneath, sparsely pubescent or glabrate above, the margins repand-denticulate to coarsely serrate; inflorescences 7-20 cm. long, long- pedunculate, the flowers few or many, on slender pedicels 2.5-7 cm. long, pubescent or glabrous, not secund; calyx lobes 2-4.5 mm. long, entire, ciliate, the 2 lower ones at the tip of the spur slightly shorter than the other 3; corolla strongly zygomorphic, red or yellow or the spur red or rose with the upper part of the tube yellow, glabrous or puberulent, the spur 1.5-2.5 cm. long, the tube cleft dorsally to the base, sometimes fenestrate on the ventral side, the 2 upper lobes narrow, slightly recurved, the 3 lower ones partially fused; staminal column exserted, about 2 cm. long; capsule about three-fourths inferior, 8-9 mm. long; seeds minute, smooth and shining.
HIPPOBROMA G. Don
Coarse, annual or perennial herbs, erect to decumbent, simple or sparsely branched, pubescent or nearly glabrous, the stems leafy; leaves alternate, sessile or nearly so, the margins of the blades coarsely dentate or laciniate-dentate; flowers
FIG. 52. Heterotoma lobelioides. A, terminal part of stem and inflorescence, x Vz; B, leaves from another plant, X Vfe; C, flower partly opened, X 2; D, anthers, X 5; E, flower dissected to show origin of filaments and the style dissected out, X 2; F, stigma, much enlarged.
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414 FIELDIANA: BOTANY, VOLUME 24
pedicellate in the leaf axils, bracteolate; calyx adnate to the ovary, 5-lobate; corolla salverform, the tube greatly elongated, narrowly cylindrical, not cleft dorsally, the 5 lobes subequal; stamens 5, the filaments connate except near the base, the anthers connate into a tube obliquely open at the top; ovary about three-fourths inferior; style (together with corolla and stamens) persistent in fruit; fruit capsular, bilocular, apically dehiscent; seeds numerous, minute, foveolate-reticulate.
A monotypic genus, the species widely distributed.
Hippobroma longiflora (L.) G. Don, Gen. Hist. 3: 717. 1834; McVaugh, Bull. Torrey Bot. Club 67: 783. 1940. Lobelia longiflora L. Sp. PL 930. 1753. Rapuntium longiflorum Mill., Diet. ed. 8. 1768. Isotoma longiflora Presl, Prodr. Monogr. Lobel. 42. 1836. Laurentia longiflora Endl. Gen. PL 512. 1838. Flor de San Juan.
Damp thickets, banks, or fields, often a weed about dwellings or in cultivated or waste ground, sea level to 950 m.; Alta Verapaz; Escuintla; Izabal; Peten; Quezaltenango; Retalhuleu; San Marcos; Santa Rosa; Suchitepequez. Southern Florida; Mexico; British Honduras to El Salvador and Panama; West Indies; South America; introduced in the Old World tropics.
Usually erect plants, rarely decumbent, mostly 20-50 (-90) cm. tall, the stems stout, hispidulous or glabrate, the sap acrid, milky; leaves usually numerous, sessile or nearly so, the blades oblanceolate or oblong-elliptic, mostly 10-20 cm. long, acute or acuminate and mucronulate, attenuate to the base, the margins coarsely repand- dentate to laciniate-dentate and minutely callose-denticulate in the sinuses, glabrous above or nearly so, thinly pilose or coarsely villous beneath; flowers borne in the upper leaf axils, the pedicels to about 15 mm. long; hypanthium in anthesis turbinate, sparsely short-hirsute, in fruit ellipsoid or ellipsoid-campanulate, 7-9 mm. in diameter, the calyx lobes linear, 10-22 mm. long, callose-denticulate; corolla white, salverform, 8-16 cm. long, puberulent outside, the slender tube 5-13 cm. long, 1-2.5 mm. broad, the lobes subequal, spreading, 12-27 mm. long; anther tube 5-6 mm. long, all the anthers white-barbellate at the apex; capsule 12-17 mm. long, pendent at maturity, loculicidally dehiscent by 2 apical valves; seeds light brown, 0.6-0.8 mm. long, minutely foveolate-reticulate.
Said to be poisonous to livestock, and that the acrid, milky sap may cause inflammation of the skin and eyes.
LOBELIA Linnaeus
Annual or perennial herbs or sometimes shrubs, pubescent or glabrous; leaves alternate, cauline or sometimes all or mostly basal, petiolate or sessile, the margins of the blades commonly serrate or dentate or if subentire, usually with minute callosities; flowers commonly pedicellate in the axils of upper, usually reduced leaves or bracts, but the area of inflorescence often appearing pedunculate and terminal; calyx tube adherent to the ovary, the lobes 5; corolla irregular, the tube cleft dorsally to the base in most species, often fenestrate, the limb more or less bilabiate, the 2 upper lobes usually erect, the 3 lower ones spreading; stamens 5, usually free from the
FIG. 53. Hippobroma longiflora. A, habit of plant, X Vz; B, part of corolla, showing spreading limb, exserted staminal column and stigmatic portion of style, X 2Vz\ C, three views of capsule: left, opened capsule enclosed in calyx; upper right, capsule opened with seeds; lower right, apical view of opened pores, all x 2Vz; D, anther column partly dissected, X 2'/2.
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corolla, the anther tube partially closed at the orifice by the incurved tips of the longer anthers, the 2 shorter ones usually tufted at the apex; corolla, with stamens and style, withering and persistent on the fruit; ovary partially or wholly inferior; fruit capsular, dehiscent by apical valves; seeds numerous, minute.
A large genus with perhaps 200 species, widely distributed in temperate and tropical regions, in the tropics chiefly in the mountains. Of the 17 species treated here, a few have not been reported in Guatemala but may be expected as they occur in Chiapas, Mexico. I have seen no material of one of these, L. caeciliae Wimmer, as it is known only from the type collection which was deposited in Berlin and presumably destroyed.
Corollas 2-5 cm. long.
Corollas in various shades of red, or with yellow intermixed, or rarely yellow throughout.
Plants herbaceous; corollas cardinal-red L. cardinalis.
Plants suffrutescent or woody throughout; corollas red and yellow or yellow throughout, not cardinal-red.
Pedicels mostly 10-20 cm. long; corollas 4-5 cm. long L. aguana.
Pedicels mostly 1-8 cm. long; corollas 2-3 cm. long L. laxiflora.
Corollas orchid-purple (fide Steyermark) L. nubicola.
Corollas less than 1.5 cm. long.
Leaves so close together on the stem as to appear subimbricate, the blades orbicular to rounded-ovate, commonly less than 0.5 cm. long and wide (rarely to 0.7 cm.), the margins sharply and uniformly dentate to the base.
L. standleyi.
Leaves not close together on the stem (sometimes clustered near the base of the
stem but never appearing subimbricate), the blades neither orbicular nor
rounded-ovate, commonly more than 0.5 cm. long, the margins various but if
dentate, the teeth not uniform to the base.
Leaf blades mostly 6-15 cm. long, long-acuminate, the margins often doubly
serrate, short teeth alternating with the longer ones L. stenodonta.
Leaf blades mostly 0.5-6 cm. long, not long-acuminate, the margins various but
never doubly serrate.
Leaf blades, at least the lower ones, broadly ovate to lanceolate. Hypanthium almost patelliform or very shallowly cupular, the capsule less
than one-third inferior. Pedicels mostly 1.5-2.5 cm. long, the bracts 1-2 cm. long; corollas 8-9 mm.
long L. diastatoides.
Pedicels mostly 0.5-1.5 cm. long, the minute bracts 1-2 mm. long; corollas
about 4 mm. long L. xalapensis.
Hypanthium more or less campanulate, hemispheric, turbinate, or narrowly
obconic, the capsule one-half to three-fourths inferior.
Hypanthium in anthesis narrowly obconic, the mature capsule three- fourths or more inferior; corollas 3-5 mm. long L. yucatana.
Hypanthium in anthesis campanulate to turbinate, the mature capsule
one-half to two-thirds inferior; corollas 6-13 mm. long. Inflorescence appearing pedunculate, the flowers in the axils of much reduced leaves or bracts; pedicels bracteolate at base.
NASH: FLORA OF GUATEMALA 417
Leaves uniformly pubescent or at least the lower ones villous-hirsute
beneath.
Stems straggling or decumbent; leaves conspicuously cauline; leaf margins essentially entire; seeds smooth, not visibly striate with
20 X magnification L. illota.
Stems erect; leaves often crowded near the base of the stem; leaf margins subentire to denticulate or coarsely serrate; seeds with faint longitudinal lines visible with 20 X magnification.
L. sartorii.
Leaves glabrous L. caeciliae.
Inflorescence not appearing pedunculate, the lower flowers, or some- times all, in the axils of scarcely reduced upper leaves; pedicels
ebracteolate L. longicaulis.
Leaf blades, at least the lower ones, spathulate, obovate, oblanceolate, or
oblong to narrowly elliptic. Inflorescence subumbellate, the 1-6 pedicels arising almost together; corollas
laterally fenestrate L. umbellifera.
Inflorescence racemose or the flowering stem appearing scapose and the
flower solitary; corollas not laterally fenestrate. Leaf blades mostly spathulate to oblanceolate, commonly 2-4 times longer
than broad; plants of alpine or subalpine meadows, 3,000-3,500 m. Plants caespitose, with leaf blades 0.6-1.5 cm. long, or the stems sometimes creeping and rooting at the nodes with leaves usually 1-
3.5 cm. long; corollas 6-10 mm. long L. nana.
Plants not caespitose, sometimes sending out filiform, leafy stolons, these rooting at the tips; leaf blades of flowering stems mostly 1-3 cm. long but those of the stolons usually smaller, 0.5-1 cm. long.
L. stolonifera.
Leaf blades narrowly oblong to narrowly elliptic, mostly 2-7 cm. long, commonly 5-10 times longer than broad; plants of damp forest and thicket, 250-1,800 m L. mexicana.
Lobelia aguana Wimmer in Fedde, Repert. Sp. Nov. 38: 86. 1935. L. laxiflora var. insignis Donn.-Sm. Bot. Gaz. 16: 12. 1891 (type from Volcan de Agua, Sacatepequez, Donn.-Sm. 2173). Pitijo; coral (San Marcos).
Damp, brushy, mountain slopes or in damp or wet, mixed or pine-fir forest, 2,100-3,500 m.; Huehuetenango; Quezaltenango; San Marcos; Sacatepequez (type from Volcan de Agua, Kellerman 7502); Solola; Totonicapan. Mexico.
Plants herbaceous or suffrutescent, the stems erect, usually simple, 1-2 m. tall, very thick and stout, glabrous or nearly so, densely leafy; leaves sessile or subsessile, essentially glabrous or somewhat pubescent, the blades mostly 8-20 cm. long and 1.5-3 cm. wide, lance-linear, lanceolate, or elliptic, narrowly long-attenuate, narrowed to the obtuse base, the margins finely incurved-serrate or subentire, the venation beneath conspicuously elevated; flowers few to many in the uppermost leaf axils, the pedicels mostly 10-20 cm. long, glabrous; hypanthium glabrous, cupular in anthesis, broadly hemispheric in fruit and 10-13 mm. broad, the calyx lobes linear or narrowly
418 FIELDIANA: BOTANY, VOLUME 24
triangular, acute, entire, 6-14 mm. long; corolla 4-5 cm. long, glabrous or sparsely pubescent, bright red to salmon-red, often yellow at base, or the tube red and the lower lip pale orange, the tube fenestrate laterally, the lobes 2-2.5 cm. long; anther tube 7-11 mm. long; capsule about two-thirds inferior, 1-1.5 cm. long.
The showiest and most handsome of the Guatemalan Lobe- liaceae. In general appearance these plants resemble L. laxiflora HBK., but are much larger in all parts. They are seldom plentiful, occurring mostly above the highest limits at which L. laxiflora grows.
Lobelia caeciliae Wimmer, Ann. Nat. Mus. Wien. 56: 348. 1948.
Not in Guatemala; known only from the type collection from Chiapas, Mexico, C. & E. Seler 1909.
Erect plants, diffusely branched, the stems angulate, subpubescent below, glabrate above, to 30 cm. tall; leaves thin, glabrous, the blades ovate, 3-7 cm. long and 1-4 cm. broad, acute or acuminate, rounded at base or narrowing into the more or less pubescent petiole, the margins minutely denticulate; racemes terminal, lax, the pedicels 1-2 cm. long, glabrous, incurved-ascending, minutely bibracteolate at base; hypanthium before anthesis broadly turbinate, soon semi-globose, glabrous, the calyx lobes subulate, entire, about 2 mm. long; corolla white and blue, about 8 mm. long, glabrous outside, the tube puberulent within, the 2 upper lobes sublinear, the lower ones oblong-spathulate; filament tube about 3 mm. long, the anther tube about 1 mm. long, lavender-blue, the 2 lower anthers barbate at the apex; capsule nearly globose, to 3 mm. wide, half inferior; seeds oblong-ellipsoid, brownish, striate, barely 0.4 mm. long.
I have seen no material of this species. The type, deposited in the Berlin herbarium, has presumably been destroyed.
Lobelia cardinalis L. Sp. PI. 930. 1753; McVaugh, Ann. Mo. Bot. Gard. 27: 347-349. 1940; Bol. Soc. Bot. Mex. 23: 48-54. 1958. Rapuntium cardinalis Mill. Gard. Diet. ed. 8, Rapuntium, No. 1. 1768. L. graminea Lam. Encycl. 3: 583. 1791. L. fulgens Willd. Hort. Berol. t 85. 1809. L. splendens Humb. & Bonpl. ex Willd. op. cit. t. 86. Rapuntium splendens Presl, Prodr. Monog. Lobel. 26. 1836. R. gramineum Presl, I.e. R. fulgens Presl, I.e. L. mucronata Engelm. in Wisliz. Tour No. Mex. 108. 1848 (not L. mucronata Cav. 1800). L. fulgens var. multiflora Paxton, Mag. Bot. 15: 7. 1849. Dortmannia fulgens Kuntze, Rev. Gen. 972. 1891. D. graminea Kuntze, I.e. D. phyllostachya Kuntze, torn. cit. 973. D. splendens Kuntze, I.e. L. cardinalis subsp. graminea (Lam.) McVaugh, Ann. Mo. Bot. Gard. 27: 347. 1940. L. cardinalis subsp. graminea var. graminea McVaugh, torn. cit. 348. L. cardinalis subsp. graminea var. pseudolsplendens McVaugh, I.e. L. cardinalis subsp. graminea var.
NASH: FLORA OF GUATEMALA 419
multiflora McVaugh, torn. cit. 349. L. cardinalis subsp. graminea var. phyllostachya McVaugh, I.e.
Marshes, edges of streams, in damp or wet soil in forest, and in damp or wet fields, 300-2,100 m.; Alta Verapaz; Chimaltenango; Chiquimula; Huehuetenango; Izabal, Jalapa; Jutiapa; Peten; El Quiche; Santa Rosa. Southern Canada and the United States; Mexico; British Honduras to Panama; Colombia.
Erect perennials from fibrous roots, the stems usually stout, commonly 30-90 cm. tall but sometimes attaining a height of 200 cm., glabrous to densely pubescent; leaves numerous, sessile or the lower ones short-petiolate, the blades narrowly linear to narrowly elliptic-oblong or lanceolate to ovate, mostly 5-15 cm. long, acute or acuminate, attenuate to the base, the margins subentire or finely denticulate to coarsely dentate; pedicels mostly 0.5-2 cm. long, solitary in the axils of much reduced upper leaves, each pedicel subtended by a pair of bracteoles; hypanthium in anthesis obconic to short- campanula te, glabrous or pubescent, costate, becoming hemispheric in fruit, the calyx lobes linear-subulate, mostly 0.8-2 cm. long, the margins entire, sometimes ciliate near the apex; corolla cardinal-red, 2.5-4 cm. long, somewhat puberulent outside, the tube fenestrate, the 2 upper lobes erect, linear, 1-2 cm. long, the 3 lobes of the lower lip 3-5 mm. wide (in ours), spreading, narrowed at base; filament tube red, the anther tube bluish gray, 3-4 mm. long; capsule about half inferior; seeds linear-oblong, light brown, foveate-reticulate, about 1 mm. long.
A complex of several wide-ranging and very closely related populations. The Central American and Mexican material is quite variable in leaf form. McVaugh (1940) refers all tropical material to L. cardinalis subsp. graminea (Lam.) McVaugh, under which he recognizes 4 varieties. Two of these, var. graminea (essentially glabrous plants with leaves linear to narrowly linear-lanceolate, 8- 14 times as long as broad) and var. multiflora (densely short- pubescent plants with lanceolate to ovate leaves, usually 6-8 times as long as wide) occur in Guatemala. It is true that the extremes are quite distinct, but there are also various intermediate forms, some with dense, short pubescence but with narrowly linear leaves (i.e., Standley 76148 from Jutiapa and Tun Ortiz 975 from Peten), and others with more or less pubescent stems but essentially glabrous, lanceolate leaves (L. O. Williams 13540 from Alta Verapaz). It is because of the many intergrading forms that I prefer not to separate the varieties here.
Lobelia diastatoides McVaugh, Am. Midi. Nat. 24: 695. 1940.
Not reported from Guatemala, but may be expected there as it occurs in the mountains of Chiapas (around .2,100 m.) as well as in Oaxaca.
420 FIELDIANA: BOTANY, VOLUME 24
Erect or decumbent perennials, the stems one or several, purplish at base, glabrous above, more or less pubescent below; leaves cauline, mostly 1.5-3 cm. long, the upper ones reduced, sessile, the blades broadly lanceolate to narrowly elliptic, acute, the lower ones subsessile to short-petiolate, the blades rhombic-ovate, acute or obtuse, acute or cuneate at the base, more or less pubescent above, glabrous or pubescent beneath, the margins crenate or shallowly serrate; inflorescences appearing pedunculate, the slender pedicels mostly 1.5-2.5 cm. long, ascending, upcurved, glabrous or sparsely pubescent near the base, ebracteolate, solitary in the axils of bracts or reduced bractlike leaves, these mostly 1-2 cm. long; hypanthium in anthesis very shallowly cupular to almost petelliform, glabrous, the calyx lobes linear- subulate, entire, glabrous or sometimes inconspicuously ciliate; corolla light blue to violet, 8-9 mm. long, the tube 4-5 mm. long, dorsally cleft, the 3 lower lobes spreading; anther tube 1.5-2 mm. long; capsule ellipsoid, 4-6 mm. long, nearly superior; seeds ellipsoid, shining, about 0.6 mm. long.
Lobelia illota McVaugh, Am. Midi. Nat. 24: 689. 1940.
Known only from the type collection, Chiapas, Mexico, Rovirosa 900.
Herbs with weak stems, 15-20 cm. long, straggling or decumbent, more or less canescent-pubescent throughout; leaves cauline, petiolate, the blades broadly elliptic, 2-4.5 cm. long, acute or obtuse, acute or broadly cuneate at the base, the margins sinuate, essentially entire; area of inflorescence 7-9 cm. long, the pedicels ascending, flexuous, to about 1 cm. long, minutely bracteolate at the base; hypanthium in anthesis hemispheric, densely pubescent, the calyx lobes subulate, entire, 2.5-3 mm. long; corolla blue or white, somewhat pubescent, 7-9 mm. long, the tube cleft dorsally nearly to the base; filament tube glabrous, the anther tube about 1.5 mm. long; capsule about half inferior; seeds ellipsoid, brown, shining, about 0.5 mm. long, not visibly striate with 20 x magnification.
Lobelia laxiflora HBK. Nov. Gen. & Sp. 3: 311. 1819; McVaugh, Ann. Mo. Bot. Card. 27: 350. 1940. L. persicifolia Cav. Ic. 6: 12, t. 518. 1800 (not L. persicifolia Lam. 1791). L. cavanillesiana Roem. & Schult. Syst. Veg. 5: 43. 1819. L. fissa Willd. ex Roem. & Schult, torn. cit. 57. L. rigidula HBK. Nov. Gen. & Sp. 3: 311. 1819. Rapuntium laxiflorum Presl, Prodr. Monog. Lobel. 26. 1836. R. haenkeanum Presl, I.e. R. rigidulum Presl, I.e. R. cavanillesianum Presl, torn. cit. 27. L. haenkeana A. DC. in DC. Prodr. 7: 382. 1839. L. laxiflora var. angustifolia A. DC. torn. cit. 383. L. concolor Mart. & Gal. Bull. Acad. Brux. 9: 46. 1842. Siphocampylos mollis Regel, Flora 33: 353. 1850 (based on cultivated plants of Guatemalan origin). S. warszewiczii Regel, Schweiz. Zeitschr. Gartenb. 8: 143. 1850. Tupa laxiflora Planch. & Oerst. Vidensk. Meddel. 1857: 154. 1857. T. costaricana var. stricta Planch. & Oerst. torn. cit. 155. T. costaricana var. patula Planch. & Oerst. I.e. L. persicaefolia var. mollis Vatke, Linnaea 38: 722. 1874. L. persicaefolia var. war- scewiczii Vatke, torn. cit. 723. L. nelsonii Fernald, Proc. Am. Acad.
NASH: FLORA OF GUATEMALA 421
36: 503. 1901. L. rensonii Wimmer in Fedde, Repert. Sp. Nov. 38: 85. 1935. L. laxiflora var. nelsonii McVaugh, Ann. Mo. Bot. Gard. 27: 349. 1940. L. laxiflora var. stricta (Planch. & Oerst.) McVaugh, No. Am. Fl. 32A(1): 96. 1943. L. laxiflora f. lutea Standl. & Steyerm. Field Mus. Bot. 23: 98. 1944. L. laxiflora var. patula (Planch. & Oerst.) Wimmer, Pflanzenr. IV. 276b(2): 683. 1953. L. laxiflora var. patula f. concolor (Mart. & Gal.) Wimmer, torn. cit. 684. L. laxiflora var. patula f. flava Wimmer, I.e. Chilillo and quiebra-machete (Guatemala, fide Aguilar); cohetillo and srukatzunum (Alta Vera- paz).
Wet to dry thickets, often on roadside banks and brushy slopes or in open, mixed or pine-oak forest, 600-2,900 m. (rarely 3,500 m.), most common at the lower levels; Alta Verapaz; Baja Verapaz; Chimaltenango; Chiquimula; Escuintla; Guatemala; Huehue- tenango; Jalapa; Jutiapa; El progreso; Quezaltenango; El Quiche; Sacatepequez; San Marcos; Santa Rosa; Solola; Suchitepequez; Totonicapan; Zacapa. Arizona and Mexico; Honduras and El Salvador to Panama; West Indies; Colombia and Peru.
Erect or ascending, usually more or less woody or suffrutescent perennials, simple or sparsely branched, usually more than 1 m. tall and sometimes as much as 2.5 m., the stems pubescent or glabrous; leaves numerous, short-petiolate or sessile, the blades commonly lanceolate to oblong-lanceolate, sometimes linear-lanceolate or narrowly elliptic, acuminate, attenuate to the base or acute to nearly rounded at the base, mostly 3-15 (-25) cm. long (the uppermost leaves in the area of inflorescence sometimes smaller and bractlike), densely pubescent to glabrate, the margins mostly serrate or serrulate (rarely subentire); flowers axillary, few or numerous, the pedicels erect or spreading, 1-8 (-10) cm. long, glabrous or villous, usually inconspicuously bracteolate at or below the middle; hypanthium glabrous or pubescent, in anthesis campanulate to cupular, becoming broadly hemispheric in fruit, to about 9 mm. in diameter; calyx lobes triangular, acute, entire, varying from 1-6.5 mm. in length but averaging 3-4 mm.; corolla 2-3 cm. long, glabrous or pubescent, bright to pale red or orange-red, often with yellow lobes, rarely the entire corolla yellow, the tube fenestrate, the lower lip 10-18 mm. long; anther tube 5-9 mm. long; capsule more than half inferior, 5-10 mm. long; seeds ellipsoid or ovoid, inconspicuously striate longitudinally, about 0.6 mm. long.
This variable and weedy species is probably one of the most abundant and widely distributed of Guatemalan plants, and may be found almost anywhere in the mountains of much of Central America, except at the highest elevations. McVaugh recognizes four varieties, two of them recorded from Guatemala, said to differ in amount of indument, in leaf shape, and whether the pedicels are stiff or flexuous, but the differences are not too well marked and there are no definite ranges. This is also true of the six varieties and numerous forms recognized by Wimmer. At one extreme are plants
422 FIELDIANA: BOTANY, VOLUME 24
with very narrowly elliptic leaves attenuate to each end, and with very short, triangular calyx lobes 1-3 mm. long; at the other extreme are plants with broadly lanceolate leaves, acuminate at the apex but acute to nearly rounded at the base, and narrow calyx lobes 3.5-6.5 mm. long. However, the most common plants are the intergrading forms with mostly lanceolate to oblong-lanceolate leaves and calyx lobes 3-4 mm. long. Plants with erect pedicels more or less appressed near the base and plants with spreading pedicels may be found in all groups; plants with mostly sessile leaves and plants with mostly petiolate leaves exist in both extremes. Glabrous and densely pubescent forms also occur throughout the spectrum.
Sometimes used as a remedy for mange of horses and dogs.
Lobelia longicaulis Brandg. Univ. Calif. Publ. Bot. 6: 73. 1914. L. neglecta Vatke, Linnaea 38: 720. 1874 (not L. neglecta Roem. & Schult. 1819). Dortmannia neglecta Kuntze, Rev. Gen. 2: 973. 1891. L. plebeia Wimmer in Fedde, Repert. Sp. Nov. 22: 194. 1926. L. urticifolia Wimmer, torn. cit. 195. L. poasensis Wimmer, Ann. Naturh. Mus. Wien 46: 239. 1933. L. longicaulis var. plebeia Wimmer, Pflanzenr. IV. 276b(2): 507. 1953.
In damp or wet thickets or in mixed or Cupressus or pine forest, sometimes on sandbars along streams, rarely in cornfields or oak forest, 1,300-2,700 m.; Alta Verapaz; Chimaltenango; Huehue- tenango; Jalapa; Jutiapa; Quezaltenango; Sacatepequez; Solola; Totonicapan. Central and southern Mexico; Honduras and El Salvador; Costa Rica; Panama.
Perennial herbs, commonly with several stems arising from a single root, the stems suberect, decumbent, or prostrate, to 50 cm. long, simple or with a few weak branches, sparsely furfuraceous-pubescent or almost glabrous, angled and often narrowly winged; cauline leaves rather distant, evenly distributed, sessile, subsessile, or on petioles to 1 cm. long, the blades ovate or broadly lanceolate, mostly 2-6 (-7) cm. long, acuminate or long-acuminate, acute at the base or contracted and decurrent on the petiole, sparsely furfuraceous-pubescent, especially on the veins, the margins coarsely and irregularly serrate, the bractlike leaves of the area of inflorescence lanceolate or linear; pedicels mostly 8-30 mm. long, sparsely pubescent; hypanthium in anthesis campanulate, pubescent or almost glabrous, the calyx lobes linear-subulate, entire, more or less ciliate, 3-4 mm. long; corolla pale blue, pink, purple, or white, 6-7 mm. long, dorsally cleft; filament tube 3-5 mm. long; anther tube 1-1.6 mm. long; capsule one-half to two-thirds inferior, 5-10 mm. long, ellipsoid to oblong; seeds ellipsoid, smooth, lustrous, about 0.5 mm. long.
Lobelia mexicana Wimmer, Ann. Nat. Mus. Wien. 56: 358. 1948.
NASH: FLORA OF GUATEMALA 423
Damp forest and thickets, 250-1,000 m.; Alta Verapaz; Baja Verapaz. Chiapas, Mexico.
Perennial herbs, the stems simple, solitary or numerous, erect or ascending, mostly 10-30 cm. tall, more or less puberulent, usually conspicuously so in the area of inflorescence; leaves cauline or sometimes crowded near the base of the stem, numerous, sessile or short-petiolate, the blades narrowly elliptic to narrowly oblong, mostly 2-7 cm. long, 0.3-0.7 cm. wide, acute or obtuse, frequently minutely mucronulate, attenuate to the base, glabrous or rarely more or less puberulent beneath, the margins subentire or callose-denticulate; area of inflorescence appearing pedunculate, mostly 3-15 cm. long, the pedicels filiform, puberulent, mostly 1.5-4 cm. long, solitary in the axils of the much reduced, upper bractlike leaves, these linear- filiform, 2-10 mm. long; hypanthium puberulent, in anthesis short-campanulate, campanulate in fruit, the calyx lobes linear-subulate, puberulent, entire, 2-3.5 mm. long; corolla blue, blue and white, or lavender, 7-12 mm. long, commonly puberulent, the tube 3-5 mm. long, corsally cleft, the lower lobes spreading; anther tube 1-1.5 mm. long; capsule ovoid, one-half to three-fourths inferior, 3.5-5 mm. long; seeds ellipsoid, light brown, shining, 0.4-0.5 mm. long.
Lobelia nana HBK. Nov. Gen. & Sp. 3: 317. t. 272. 1819. Rapuntium nanum Presl, Prodr. Monogr. Lobel. 22. 1836. Dort- mannia nana Kuntze, Rev. Gen. 2: 973. 1891. L. bellis Wimmer in Fedde, Repert. Sp. Nov. 22: 193. 1926.
Alpine meadows, 3,300-3,500 m.; Huehuetenango (Sierra de los Cuchumatanes). Mexico; Argentina; Bolivia; Ecuador; Peru.
Perennial herbs, glabrous, the stems very slender, arising from a stout caudex and first appearing caespitose, later creeping and rooting at the nodes, mostly 2-15 cm. long; leaves numerous, the blades spathulate to oblanceolate, mostly 0.6-1.5 cm. long, rarely to 3 cm., usually obtuse, sometimes acute, narrowed at the base into a short, marginate petiole, sometimes ciliate near the base, the margins shallowly and obscurely crenate; flowers 1-4, solitary in the upper leaf axils, the pedicels commonly 1.5-4 cm. long (rarely to 15 cm.); hypanthium in anthesis cupular or short- campanulate, in fruit campanulate, 2.5-4 mm. broad, the calyx lobes narrowly linear- lanceolate, 2.5-4.5 mm. long, entire, acute or obtuse; corolla blue, lavender, or white, 6-10 mm. long, the tube 2-4 mm. long, dorsally cleft, not fenestrate; anther tube 1.2-2 mm. long; capsule one-third to one-half inferior, 5-6.5 mm. long; seeds ellipsoid, glabrous smooth or obscurely striate, about 0.5 mm. long.
Lobelia nubicola McVaugh, N. Am. Fl. 32A: 94. 1942.
Known only from the type, collected in Liquidambar forest, middle slopes of Montana Norte to El Jutal, Cerro Brujo, southeast of Concepcion de las Minas, Chiquimula, 1,700-2,000 m., Steyermark 31048.
A slender shrub, 1.5 m. tall, glabrous; leaves short-petiolate, the blades narrowly lanceolate or linear-lanceolate, 5-12 cm. long, 1-1.5 cm. wide, narrowly long- attenuate, acute at the base, minutely and remotely crenate; flowers few, solitary in the axils of the uppermost leaves, the pedicels 3-3.5 cm. long; hypanthium in anthesis
FIG. 54. (Left) Lobelia nana. A, habit, X 1; B, flower dissected, x 3; C, anther column and stigma, X 4. (Right) Lobelia laxiflora. D, habit, X W, E, flower partially dissected, x 3; F, calyx and style, x 3; G, base of corolla with attached filaments, x 3; H, anther column with protruding style, x 3; I, seeds, much enlarged.
424
NASH: FLORA OF GUATEMALA 425
turbinate or cupular, 7-8 mm. long, the calyx lobes narrowly triangular, acute, entire or obscurely denticulate, 4-5 mm. long; corolla orchid-purple (fide Steyermark), about 4 cm. long, glabrous outside, cleft dorsally to a point about 3.5 mm. from the base, the tube narrowest about 5 mm. above the base, the 2 upper lobes about 15 mm. long, the 3 lobes of the lower lip about 13 mm. long; anther tube about 7 mm. long; fruit capsular (mature fruit not seen); seeds unknown.
Lobelia sartorii Vatke, Linnaea 38: 721. 1874. L. novella Robins. Proc. Am. Acad. 26: 167. 1891. Dortmannia sartorii Kuntze, Rev. Gen. 2: 973. 1891. L. piscinula Wimmer in Fedde, Repert. Sp. Nov. 38: 84. 1935. L. sartorii var. piscinula Wimmer, Pflanzenr. IV. 276b (2): 517. 1953. Lachpzc (Quecchi, Alta Verapaz).
Grassy banks or in wet forest, often in pine forest, sometimes on limestone, 350-3,500 m.; Alta Verapaz; Baja Verapaz; Huehue- tenango; El Progreso; Peten; El Quiche. Mexico.
Slender herbs, simple or branching, the stems erect, mostly 15-30 cm. tall, glabrous above, more or less pubescent below; cauline leaves usually few and below the middle of the stem, sessile or short-petiolate, the uppermost ones reduced and usually narrower than the others, basal leaves on petioles 1-3 cm. long, the blades lanceolate to broadly ovate, mostly 2-5 cm. long, acute or obtuse, acute to cordate at the base, the margins subentire to denticulate or coarsely serrate, at least the lower leaves usually villous-hirsute with multiseptate hairs, sometimes glabrous or glabrate; inflorescences racemose, often appearing pedunculate, the few to many flowers on bracteolate pedicels 1-3 cm. long; bracts linear-subulate, 2-6 mm. long; hypanthium glabrous, in anthesis short-campanulate, broadly obconic in fruit and about 3.5 mm. broad, the calyx lobes linear-subulate or narrowly triangular, entire, glabrous or rarely pubescent, 2-4 mm. long; corolla blue, violet, or white with blue or lavender markings, 7-12 mm. long, glabrous outside, cleft dorsally, the tube 4-6 mm. long, sometimes also fenestrate laterally, the 2 upper lobes erect, the lower ones spreading; anther tube 1-2 mm. long; capsule one-half to two-thirds inferior, 3.5-6 mm. long; seeds ellipsoid, brown, somewhat lustrous, faintly longitudinally striate, about 0.6 mm. long.
Lobelia standleyi McVaugh, N. Am. Fl. 32A: 47. 1942.
Damp or wet, alpine or subalpine meadows, 3,000-3,700 m.; Huehuetenango (type from Chimal, Sierra de los Cuchumatanes, Standley 81108}; Solola; Totonicapan.
Small, repent plants, glabrous throughout or the stems, pedicels, and hypanthium minutely hispidulous, the stems commonly 3-6 cm. long, rooting at most of the nodes; leaves numerous and close together, somewhat imbricate and sometimes concealing the stems, subsessile, the blades orbicular to rounded-ovate, mostly 3-5 mm. long and wide (rarely to 7 mm.), rounded at each end, abruptly and shortly narrowed to the base, the margins coarsely and sharply dentate with uniform teeth; flowers solitary in the upper leaf axils, the pedicels to 4 mm. long; hypanthium campanulate to shallowly cupular in anthesis, hispidulous or glabrous, about 2 mm. long, the calyx lobes linear, acute, entire, 4-6 mm. long; corolla pale blue to white,
426 FIELDIANA: BOTANY, VOLUME 24
glabrous or minutely hispidulous, 8-10 mm. long, the tube about 3 mm. long, dorsally cleft about halfway to the base, not fenestrate; anther tube 1.7-2 mm. long; capsule about half inferior, about 6 mm. long.
Lobelia stenodonta (Fern.) McVaugh, Bull. Torrey Bot. Club 67: 144. 1940. Heterotoma stenodonta Fern. Proc. Am. Acad. 36: 504. 1901.
Not in Guatemala but may be expected, as it is frequent in the mountains of southern Chiapas, Mexico.
Erect perennials from slender rootstocks, simple or sparsely branched, the stems glabrous, to about 50 cm. tall; leaves cauline, sessile to short-petiolate, the blades thin, ovate to lanceolate or lance-elliptic, mostly 6-15 cm. long, 1.5-5 cm. wide, long- acuminate, the base rather abruptly cuneate, the margins conspicuously and sharply serrate to doubly serrate, the principle teeth commonly at least 2 mm. long; inflorescences terminal and axillary, appearing pedunculate and racemose, the pedicels solitary in the axils of reduced, bractlike leaves, ebracteolate, mostly 1.5-3 cm. long, hispidulous to sparsely papillose, the area of inflorescence 5-15 cm. long; hypanthium in anthesis campanulate, glabrous, the calyx lobes linear-lanceolate, acute, obscurely denticulate, 3-5 mm. long; corolla pale bluish, 5-9 mm. long, the tube fenestrate; anther tube 1.5-2 mm. long; capsule ellipsoid, about half inferior, narrowly beaked; seeds light brown, ellipsoid, smooth, about 0.6 mm. long.
Lobelia stolonifera Donn.-Sm. Bot. Gaz. 27: 338. 1899.
Commonly in damp or wet, alpine meadows, 3,200-3,500 m.; Huehuetenango (type collected between Todos Santos and Chiantla, Sierra de los Cuchumatanes, Seler 3017); Totonicapan.
Perennial herbs, the rootstocks sometimes sending out filiform, leafy stolons, the stems simple, the flowering stems erect or suberect, solitary, mostly 7-15 (-20) cm. tall; cauline leaves 1-4 below the middle of the stem, the upper ones sessile, the lower subpetiolate, the blades mostly spathulate or oblanceolate, sometimes elliptic, 1-3 cm. long, acute to rounded at the apex, attenuate to the base, glabrous, the margins appearing crenate but actually obscurely callose-denticulate, the basal leaves and those of the stolons broadly spathulate, very obtuse, decurrent on the petiole; inflorescence bracteate, appearing pedunculate, the 3-4 flowers terminating a long, naked stem, the bracteolate pedicels suberect, 4-8 mm. long, glabrous; hypanthium in anthesis campanulate, pubescent, in fruit about 3 mm. broad; calyx lobes narrowly triangular, glabrous, acute, entire, purple, 3-5 mm. long; corolla white, blue, or white marked with purple, 10-12 mm. long, glabrous outside, the tube 3-4 mm. long, the lobes spreading, rounded or mucronulate, those of the lower lip 7-8 mm. long; anther tube about 2 mm. long; capsule ellipsoid-fusiform, more than half inferior, about 6 mm. long; seeds ellipsoid, brown, obscurely longitudinally striate, lustrous, about 0.5 mm. long.
Lobelia umbellifera McVaugh, Bull. Torrey Bot. Club 67: 144. 1940. L. fasciculata Donn.-Sm. Bot. Gaz. 27: 338. 1899 (not L. fasciculata Kuntze, 1891). Chancol, violeta (Huehuetenango).
NASH: FLORA OF GUATEMALA 427
Wet or sometimes rather dry meadows, chiefly in alpine regions, sometimes in pine forest, 2,100-4,000 m.; Chimaltenango; Huehuetenango (type from Todos Santos, Seler 3022); Quezal- tenango; San Marcos; Solola; Totonicapan. Mexico (Chiapas).
Perennial herbs from taproots, sometimes forming large mats, the stems decumbent or prostrate, slender, 5-35 cm. long, simple or sparsely branched, rather densely and evenly leafy, glabrous; leaves sessile, the blades elliptic or narrowly obovate to oblong, 0.5-2 cm. long, subacute to rounded at the apex and sometimes very minutely mucronulate, cuneate at the base, the margins crenate, sometimes strigose or scaberulous above, glabrous beneath; inflorescences subumbellate, pedunculate, the filiform, pubescent pedicels in a terminal cluster of 1-6, subtended by linear bracts mostly 5-7 mm. long; hypanthium in anthesis short- campanula te to shallowly cupular, more or less inflated in fruit and 3-7 mm. wide, glabrous or puberulent to somewhat strigose at the base, the calyx lobes narrowly linear- lanceolate, glabrous, minutely callose-denticulate, 3-5 mm. long; corolla blue or lavender, 11-13 mm. long, the tube about 3 mm. long, fenestrate laterally, the limb spreading, the emarginate lobes usually conspicuously apiculate; anther tube 1-1.5 mm. long; capsule about two-thirds inferior; seeds ellipsoid, brown, lustrous, about 0.7 mm. long.
Lobelia xalapensis HBK. Nov. Gen. & Sp. 3: 315. 1819. L. monticola HBK. torn. cit. 316. L. cliffortiana Roem. & Schult. Syst. V: 53. 1819 (not L. cliffortiana L. Sp. PL ed. 3, 2: 1320. 1753). L. palmaris Willd. ex Roem. & Schult. torn. cit. 56. L. mollis Graham, Edinb. New Phil. Journ. 8: 185. 1829. Rapuntium affine, xalapense, monticolum Presl, Prodr. Monogr. Lobel. 25. 1836. R. molle Presl, torn. cit. 30. L. ocimoides Kuntze, Linnaea 24: 178. 1851. Dortmannia mollis and monticola Kuntze, Rev. Gen. 2: 972. 1891. D. ocimoides and xalapensis Kuntze, torn. cit. 973. D. cliffortiana var. xalapensis Kuntze, op. cit. 3: 187. 1898.
Damp or wet thickets and wet, open pine, pine-oak, or pine- juniper forest, often on steep banks or slopes, frequently on limestone, rarely on dry hills, mostly 60-1,400 m., rarely to 3,400 m.; Alta Verapaz; Baja Verapaz; Huehuetenango; Izabal; Peten; Quezaltenango; Retalhuleu; San Marcos; Santa Rosa. Southern Mexico; British Honduras; Honduras; Costa Rica; Nicaragua; Panama; Lesser Antilles; Venezuela and Colombia southward to Argentina; Galapagos Islands.
Erect annuals, the stems simple or with few ascending branches, usually sparsely pubescent or puberulent, at least below; leaves cauline, on short or long, slender petioles, the blades somewhat pubescent on the veins beneath or almost wholly glabrous, broadly ovate to lance-ovate, mostly 1-4 cm. long, acute to very obtuse, rounded or cordate at the base and abruptly contracted into the narrowly winged petiole, the margins sinuate or irregularly and coarsely dentate; racemes axillary and terminal, appearing pedunculate, short or elongating and many -flowered, the pedicels
428 FIELDIANA: BOTANY, VOLUME 24
slender, spreading, 0.5-1.5 cm. long, minutely puberulent, subtended by minute, linear or subulate bracts; hypanthium in anthesis flat or shallowly cupular, puberulent, in fruit cupular and often glabrate, the calyx lobes linear-subulate, entire, 2-4 mm. long, ciliate or not; corolla commonly blue or purplish, sometimes white or pinkish, about 4 mm. long, the tube glabrous outside, cleft dorsally to a point less than 1 mm. from the base, the lower lip 1.5-2 mm. long; anther tube 0.5-1 mm. long; capusle one- third inferior or less, ellipsoid, mostly 3.5-5 mm. long; seeds ellipsoid, brown, lustrous, to 0.5 mm. long.
Common and somewhat weedy plants in many parts of Guatemala. They have been reported as L. cliffortiana L., a species not known to occur in Central America.
Lobelia yucatana Wimmer in Fedde, Repert. Sp. Nov. 38: 83. 1935.
Thickets in dry arroyo, Peten (Uaxactun, Bartlett 12221). Yucatan Peninsula of Mexico, the type from Pocoboh, Gaumer 1327.
Slender annuals, the stems apparently decumbent or procumbent, 15-30 cm. long, branching, glabrous above, short-pubescent below; leaves cauline, few, the lower ones largest, slender-petiolate, the blades ovate, obtuse, mostly 2-4 cm. long, gradually attenuate to the base or abruptly contracted and decurrent on the petiole, strigose above or essentially glabrous, the margins coarsely crenulate, the upper leaves short- petiolate, the blades ovate or elliptic, obtuse; inflorescence very lax and few-flowered, not distinctly pedunculate, the pedicels 8-15 mm. long, pubescent, the bracts linear; hypanthium glabrous, narrowly obconic in anthesis, ellipsoid or oblong in fruit, the calyx lobes narrow, subulate, entire, glabrous, 1-1.5 mm. long; corolla white, about 3 mm. long; anther tube about 0.5 mm. long; capsule three-fourths to four-fifths inferior, 4-5 mm. long; seeds smooth, lustrous, to 0.5 mm. long.
PRATIA Gaudichaud
Herbaceous, suffrutescent, or shrubby plants, the stems erect, ascending, or decumbent; leaves alternate, petiolate or sessile, the margins of the blades subentire or crenate to dentate or serrate-denticulate; flowers pedicellate, solitary in the axils of the uppermost leaves or in the axils of bracts, the inflorescence than appearing terminal and racemose or subcorymbose; calyx 5-lobate, the tube adnate to the inferior ovary except for a small portion exceeding the ovary and appearing as a rim above it, the segments equal or subequal, persistent; corolla red, purple, or white, tubular, the tube usually broadest at the base, somewhat curved, cleft dorsally to the base or nearly so, or rarely cleft for only about one-third its length, the bilabiate limb 5-lobate, the 2 upper lobes linear, longer than the 3 lobes of the lower lip; filaments distinct at the base, free from the corolla or nearly so, the anther tube somewhat narrowed at the orifice or closed by the incurved tips of the 3 longer anthers, the 2 shorter ones with a tuft of hairs at the apex; style geniculate and separating with the corolla and stamens from the fruit; fruit somewhat fleshy, indehiscent; seeds minute, broadly ellipsoid or almost globose, foveolate.
NASH: FLORA OF GUATEMALA 429
It is interesting that Moeliono & Tuyn, in their treatment of the Campanulaceae for the Flora Malesiana (Ser. I, 6: 122, 133-135. 1960), have reduced certain species of Pratia to Lobelia, and state that they have found both capsular and baccate fruits of L. angulata Forst.
Twenty-nine species in tropical America and Asia, Australia, and New Zealand. Two are in Guatemala and one other is known in Central America, from Nicaragua.
Leaf margins finely serrate; calyx lobes conspicuously longer than the tube (about 3 times as long), 5-12 mm. wide at the base; corolla tube cleft to a point 4-6 mm. from its base P. calochlamys.
Leaf margins crenate; calyx lobes almost equalling the tube in length or sometimes only about half as long, 3-5 mm. wide at the base; corolla tube cleft to a point 15-20 mm. from its base P. guatemalensis.
Pratia calochlamys (Donn.-Sm.) Wimmer in Fedde, Repert. Sp. Nov. 29: 50. 1931. Centropogon calochlamys Donn.-Sm. Bot. Gaz. 46: 112. 1908.
Damp or wet forest, 900-2,400 m.; Alta Verapaz (type collected near Coban, Tuerckheim II. 1893); Huehuetenango.
Erect, glabrous herbs, 30-60 cm. tall, the stems simple; leaves on stout petioles 1- 3.5 cm. long, the blades elliptic to elliptic-oblong or lance-oblong, 5-15 cm. long, abruptly or gradually acuminate, cuneate or attenuate to the base and decurrent on the petiole, the margins evenly and finely serrate, the uppermost leaves often much reduced and crowded; pedicels solitary in the axils of the upper leaves, mostly 2-6 cm. long, bracteolate; calyx lobes triangular or lance-ovate, about 3 times as long as the tube, acute or subacute, dentate, often purplish; corolla rose-purple or bright red tinged with purple, 3.5-4.5 cm. long, glabrous outside, the tube slightly curved, broadest at the base, slightly contracted near the apex, cleft to a point 4-6 mm. from its base, the lobes long-attenuate, the 2 upper ones erect, 15-20 mm. long, the 3 lobes of the lower lip slightly recurved, 7-10 mm. long; filament tube 2.5-3 cm. long, the anther tube 7-8 mm. long; fruit hemispheric, not inflated, about 8 mm. broad; seeds light brown, ellipsoid or oblong, flattened, shallowly pitted-reticulate, 0.5-1 mm. long.
Pratia guatemalensis (Robins.) Wimmer in Fedde, Repert. Sp. Nov. 29: 50. 1931. Centropogon guatemalensis Robins, ex Donn.- Sm. Bot. Gaz. 20: 4. 1895.
Wet, mixed forest, 1,000-1,500 m.; Alta Verapaz (type from Pansamala, Tuerckheim 728); Baja Verapaz.
Erect herbs, essentially glabrous throughout except in the axils of the bracts, the stems mostly simple, to about 40 cm. tall; leaves on petioles mostly 2-8 cm. long, the blades broadly ovate, obovate, or broadly elliptic, mostly 10-20 cm. long, abruptly short-acuminate, acute or cuneate at the base and decurrent on the petiole, the margins crenate; inflorescence appearing racemose or subcorymbose, commonly with
FIG. 55. Pratia calochlamys. A, habit, fruiting plant, x Vr, B, habit, flowering plant, x '/2j C, flower partially dissected, X IVz; D, dissection showing attachment of filaments and ovary, X 2; E, anther column, X 3Vr, F, anther column with stigma dissected out, X 5; G, calyx dissected to show ovary, X 1; H, seeds, X 5.
430
NASH: FLORA OF GUATEMALA 431
10-25 flowers on bracteate, often purplish pedicels mostly 2-4 cm. long; hypanthium in anthesis short, open-campanulate, often purplish, the calyx lobes triangular, about equalling the tube or sometimes only half as long, subacute, obscurely denticulate; corolla purplish red when dry, 4.5-6 cm. long, glabrous outside, cleft to a point 15-20 mm. from its base, subfalcate, the limb bilabiate, the 2 upper lobes erect, narrowly subulate, 15-25 mm. long, the 3 lobes of the lower lip linear or narrowly elliptic, acute, 8-18 mm. long; filament tube 30-40 mm. long, the anther tube 7-8.5 mm. long; mature fruit not seen.
Publication 1238
UNIVERSITY OF ILLINOIS-URBAN*
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