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.
286 FIELDIANA: BOTANY, VOLUME 24
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.
288 FIELDIANA: BOTANY, VOLUME 24
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.
296 FIELDIANA: BOTANY, VOLUME 24
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.
308 FIELDIANA: BOTANY, VOLUME 24
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.
310 FIELDIANA: BOTANY, VOLUME 24
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.
312 FIELDIANA: BOTANY, VOLUME 24
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.
313
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.
316 FIELDIANA: BOTANY, VOLUME 24
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.
317
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.
319
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.
323
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.
325
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.
335
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.
339
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.
341
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;
344
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.
357
358 FIELDIANA: BOTANY, VOLUME 24
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.
359
360 FIELDIANA: BOTANY, VOLUME 24
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.
362
DIETERLE: FLORA OF GUATEMALA 363
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.
364
DIETERLE: FLORA OF GUATEMALA 365
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.
366
DIETERLE: FLORA OF GUATEMALA 367
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.
368
FIELDIANA: BOTANY, VOLUME 24
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.
DIETERLE: FLORA OF GUATEMALA 369
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.
370
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.
373
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.
379
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.
381
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.
404
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.
406
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.
410
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.
413
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.
415
416 FIELDIANA: BOTANY, VOLUME 24
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
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