UNIVERSITY OF
ILLINOIS LIBRARY
AT URBANA-CHAMPAIGN
BIOLOGY
FLORA OF GUATEMALA
DOROTHY L. NASH
and
LOUIS O. WILLIAMS
Genera contributed by:
KENNETH M. BECKER
JOHN J. FAY
JEROLD L. GRASHOFF
THOMAS E. MELCHERT
DAN H. NICOLSON
BIOLOGY
FIELDIANA: BOTANY
Volume 24, Part XII
Published by
Field Museum of Natural History
May 15, 1976
FLORA OF GUATEMALA
PART XII
IN MEMORIUM
Paul Carpenter Standley
1884-1963
Who Initiated the "Flora of Guatemala'
FLORA OF GUATEMALA
Vernonieae, Astereae, Inuleae, Heliantheae,
Anthemideae, Cynareae, Mutisieae, Cichorieae
DOROTHY L. NASH
Research Specialist, Department of Botany
Field Museum of Natural History
Eupatorieae, Helenieae, Senecioneae
LOUIS O. WILLIAMS
Curator Emeritus, Department of Botany
Field Museum of Natural History
Genera contributed by:
KENNETH M. BECKER
JOHN J. FAY
JEROLD L. GRASHOFF
THOMAS E. MELCHERT
DAN H. NICOLSON
FIELDIANA: BOTANY
Volume 24, Part XII
Published by
Field Museum of Natural History
May 15, 1976
LAURA M. SCHLIVEK
Associate 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
Family in Part XII
PAGE
Compositae 1
I <
V- 24 '. ! 2
LIST OF ILLUSTRATIONS
The tribes in systematic order
Page
TRIBE I, Vernonieae
1. Elephantopus angustifolius 455
2. Elephantopus mottis 456
3. Elephantopus spicatus 457
4. Harleya oxylepsis 458
5. Lepidonia paleata 459
6. Liabum bourgeaui 460
7. Pacourina edulis 461
8. Piptocarpha chontalensis 462
9. Spiracantha cornifolia 463
10. Struchium sparganophorum 464
11. Vernonia standleyi 465
TRIBE II, Eupatorieae
12. Adenostemma hirtiflorum 466
13. Ageratum cordatum 467
14. Ageratum rugosum 468
14a. Brickellia paniculata 469
15. Carminatia tenuiflora 470
16. Eupatorium collinum 471
17. Eupatorium lanicaule 472
18. Eupatorium monticola 473
19. Eupatorium muelleri 474
20. Eupatorium nubivagum 475
21. Eupatorium sodali 476
22. Isocarpha oppositifolia 477
23. Macvaughiella standleyi 478
24. Mikania vitifolia 479
25. Oxylobus glanduliferus 480
26. Piqueria trinervis 481
27. Stevia polycephala 482
TRIBE III, Astereae
28. Achaetogeron guatemalense 483
29. Archibaccharis asperifolia 484
30. Aster spinosus 485
31. Astranthium purpurascens 486
32. Baccharis vaccinioides 487
VII
VIII
33. Conyza bonariensis 488
34. Egletes liebmannii var. yucatana 489
35. Erigeron karvinskianus 490
36. Gymnosperma glutinosa 491
37. Haplopappus stoloniferus 492
38. Heterotheca graminifolia 493
39. Lagenophora cuchumatanica 494
40. Solidago stricta 495
TRIBE IV, Inuleae
41. Achyrocline deflexa 496
42. Adenocaulon lyratum 497
43. Epaltes mexicana 498
44. Gnaphalium americanum 499
45. Gnaphalium brachyphyllum 500
46. Pluchea odorata 501
47. Pseudoconyza viscosa 502
TRIBE V, Heliantheae
48. Acanthospermum hispidum 503
49. Aldama dentata 504
50. Alepidocline annua 505
51. Ambrosia cumanensis 506
52. Aphanactis standleyi 507
53. Baltimora recta 508
54. Bidens squarrosa 509
55. Borrichia arborescens 510
56. Calea skutchii 511
57. Calyptocarpus wendlandii 512
58. Chrysanthellum americanum 513
59. Clibadium arboreum 514
60. Coreopsis mutica var. microcephala 515
61. Cosmos caudatus 516
62. Cuchumatanea steyermarkii 517
63. Dahlia coccinea 518
64. Delilea berterii 519
65. Desmanthodium guatemalense 520
66. Eclipta alba 521
67. Eleutheranthera ruderalis 522
68. Galinsoga urticaefolia 523
69. Garcilassa rivularis 524
70. Goldmanella sarmentosa 525
71. Heliopsis buphthalmoides 526
72. Heterosperma pinnatum 527
73. Hidalgoa ternata 528
74. Hymenostephium guatemalense 529
75. lostephane trilobata 530
76. Jaegeria hirta 531
77. Lagascea helianthifolia 532
78. Lasianthaea fruticosa 533
IX
79. Melampodium divaricatum 534
80. Melanthera nivea 535
81. Milleria quinqueflora 536
82. Montanoa guatemalensis 537
83. Neurolaena macrophylla 538
84. Notoptera scabridula 539
85. Oteiza ruacophila 540
86. Otopappus gla bratus 541
87. Parthenium hysterophorus 542
88. Perymenium grande , 543
89. Philactis liebmannii 544
90. Podachaenium eminens 545
91. Polymnia maculata 546
92. Rensonia salvadorica 547
93. Rojasianthe superba 548
94. Rumfordia standleyi 549
95. Sabazia pinetorum 550
96. Salmea scandens 551
97. Sanvitalia procumbens 552
98. Schistocarpha platyphylla 553
99. Sclerocarpus divaricatus 554
100. Selloa obtusata 555
101. Sigesbeckia agrestis 556
102. Simsia amplexicaulis 557
103. Spilanthes ocymifolia 558
104. Synedrella nodiflora 559
105. Tithonia longiradiata 560
106. Tragoceras schiedcanum 561
107. Trichospora verticillata 562
108. Tridax procumbens 563
109. Trigonospermum annuum 564
110. Verbesina fraseri 565
111. Viguiera dentata 566
112. Wedelia acapulcensis 567
113. Wedelia trilobata 568
114. Zexmenia phyllocephala 569
115. Zinnia peruviana 570
TRIBE VI, Helenieae
116. Bahia depauperata 571
117. Dyssodia montana 572
118. Espe/oa mexicana 573
119. Florestina latifolia 574
120. Galeana pratensis 575
121. Heleniwn integrifolium 576
122. Pectis elongata 577
123. Porophyllum punctatum 578
124. Schkurhia virgata 579
125. Tagetes nelsonii 580
TRIBE VII, Anthemideae
l26.Achillea millefolium 581
127. Artemisia mexicana 582
128. Chrysanthemum parthenium 583
129. Matricaria courrantiana 584
TRIBE VIII, Senecioneae
130. Emilia sonchifolia 585
130. Erechites valerianae folia 585
131. Psacalium pinetorum 586
132. Senecio conbanensis 587
133. Senecio nubivagus 588
134. Werneria nubigena 589
TRIBE IX, Cynareae
135. Cirsium mexicanum 590
TRIBE X, Mustisieae
136. Acourtia carpholepis 591
137. Chaptalia nutans 592
138. Jungia guatemalensis 593
139. Lycoseris crocata 594
140. Onoseris onoseroides 595
141. Perezia nudicaulis 596
142. Trixisinula 597
TRIBE XI, Cichoreae
143. Hieracium abscissum 598
144. Hypochoeris glabra 599
145. Lactuca graminifolia 600
146. Pinaropappus spathulatus 601
147. Sonchus oleraceus 602
148. Taraxacum officinale 603
Flora of Guatemala — Part XII
COMPOSITAE
Sunflower Family
Reference: S. F. Blake in Standl. Contr. U. S. Nat. Herb. 23:
1401-1641. 1926.
Annual or perennial herbs, shrubs, or trees, sometimes scandent; leaves opposite
or alternate, sometimes verticillate or all basal, entire to dissected, never truly
compound but often appearing so; flowers collected in a head (this rarely 1-flowered)
on a receptacle, surrounded by an involucre of phyllaries (involucral bracts); corolla
gamopetalous, regular, tubular, and 5-dentate (rarely 2-4-dentate), bilabiate, or
ligulate (flattened, strap-shaped, and usually 2-5-dentate), rarely wanting in the
pistillate flowers; stamens (in the hermaphrodite or staminate flowers) almost always
5, united by the anthers or rarely free, inserted on the corolla; ovary inferior, 1 -celled,
with a single erect anatropous ovule; style usually 2-branched, the branches
stigmatiferous inside, often bearing sterile appendages at the apex; fruit an achene,
with a single, erect seed, this without endosperm, the achene often bearing a pappus
of bristles, awns, or scales.
The corollas are of four chief kinds, ligulate or strap-shaped,
bilabiate, tubular, and filiform. Heads composed of one kind of
flowers only are called homogamous, those composed of two or
more kinds heterogamous; when heterogamous, the central flowers
(disc) are always hermaphrodite, the peripheral flowers pistillate or
neutral (lacking a style). Homogamous heads in which all the
flowers are hermaphrodite and have ligulate (in this case always 5-
dentate) corollas are called ligulate. Homogamous heads in which
all the corollas are tubular and hermaphrodite or staminate, or
filiform and pistillate, are called discoid. Heterogamous heads in
which the peripheral corollas are ligulate are called radiate', those
in which the peripheral flowers are pistillate, with tubular, filiform,
or abortive corollas are called disciform. The receptacle may be
naked, bristly, or paleaceous (bearing pales or chaff). The generic
characters are drawn to a considerable extent from the character of
the pappus, which may be of bristles, awns, scales, or teeth, or
reduced to a crown or cup, or entirely wanting.
2 FIELDIANA: BOTANY, VOLUME 24
One of the largest families of plants, with about 900 genera and
perhaps as many as 20,000 species, found in almost parts of the
earth, most numerous in temperate regions. A very few genera
besides those listed here are found in southern Central America.
The family is perhaps more abundantly represented in North
America than in any other continent. It is very well represented in
Guatemala, particularly in the mountains, the number of species
being astonishingly large for an area of this size. In the wet lowland
tropics the number of Compositae is greatly reduced, and consists
largely of weedy plants.
The Compositae, a large and complicated family in Guatemala,
has been prepared in systematic order so far as the tribes are
concerned. We have prepared a tribal key which follows these notes.
A key to the genera in each of the tribes will appear in the
appropriate place. The plates will appear at the end of the family
treatment grouped systematically by tribe, but arranged alphabet-
ically within the tribes. This should simplify the determination of
an unrecognized genus since the plates of related genera will be
grouped. Most genera are illustrated, the larger ones often with
more than one plate.
The Compositae have attracted many students in recent years
so that several useful monographs and revisions have made our
work easier. These are cited under the appropriate genera.
The great size and complexity of the Compositae has meant
that no one botanist has had a "grasp" and understanding of the
neotropical genera. Dr. S. F. Blake, friend and colleague, was the
last botanist with a broad understanding of the American
Compositae. We have used his account of the family and key to the
tribes (see reference above) as a basis for ours, with modifications
that are necessary because of the broader scope of our work. —
Louis O. Williams, August, 1975.
KEY TO THE TRIBES OF COMPOSITAE THAT OCCUR IN GUATEMALA
Corollas all ligulate or all bilabiate.
Corollas all ligulate; anthers sagittate at the base; sap milky XI. CICHORIEAE.
Corollas all bilabiate (in ours); anthers caudate at the base; sap not milky.
X. MUTISIEAE.
Corollas not all ligulate nor all bilabiate, at least the central flowers with regular 4-5-
cleft corollas; sometimes no ligulate corollas present.
Anthers caudate at the base.
Anther apices long-appendaged; receptacle setiferous (in ours); leaves more or
less spinose or prickly IX. CYNAREAE.
WILLIAMS: FLORA OF GUATEMALA 3
Anther apices not conspicuously appendaged; receptacle naked; leaves not
spinose nor prickly IV. INULEAE.
Anthers often sagittate at the base but not caudate.
Tips of the style branches truncate or subtruncate.
Phyllaries multiseriate, unequal; pappus a low crown or wanting; plants
usually strong-scented VII. ANTHEMIDEAE.
Phyllaries uniseriate and equal or nearly so, or if bi- or triseriate, the outer
series composed of shorter scales; pappus commonly composed of soft,
slender hairs but sometimes reduced to a crown; plants seldom if ever
strong-scented VIII. SENECIONEAE.
Tips of the style branches (in ours) usually acute, only rarely subtruncate.
Leaves usually opposite, or at least the lower ones usually so.
Heads homogamous, discoid.
Receptacle paleaceous; corollas usually yellow, rarely white; style
branches various, usually acute.
Pappus of 2-3 or more rigid awns or scales V. HELIANTHEAE.
Pappus of numerous, slender bristles II. EUPATORIEAE.
Receptacle not paleaceous; corollas yellow, pink, purplish, or white; style
branches usually elongated and obtuse or subacute; pappus of 1-2
series or none.
Corollas pink, purplish, or white; pappus of 1-2 series of slender bristles,
these rarely plumose, or pappus none II. EUPATORIEAE.
Corollas yellow; pappus biseriate, the inner bristles long, the outer ones
very short (Liabum) I. VERNONIEAE.
Heads heterogamous, usually radiate.
Pappus of 2-3 or more rigid awns and/or scales or sometimes absent.
Receptacle paleaceous; achenes usually thickened, hard, sometimes
flattened and winged; pappus of 2-3 or more rigid awns and/or
scales or absent V. HELIANTHEAE.
Receptacle naked (except fimbrillate in Gaillardia); achenes usually
long and narrow, rarely flattened or winged; pappus squamellae
entire, dissected into setae, or lacking VI.HELENIEAE.
Pappus biseriate, the inner bristles elongated, the outer squamellae very
short; receptacle naked, alveolate or fimbrillate (Liabum).
I. VERNONIEAE.
Leaves usually alternate.
Heads discoid.
Flowers yellow III. ASTEREAE.
Flowers white or purplish I. VERNONIEAE.
Heads radiate.
Ray flowers always yellow or red.
Plants vines; pappus of soft, slender, barbellate hairs.
VIII. SENECIONEAE.
Plants not vines; pappus squamellae entire, dissected into setae, or
lacking VI.HELENIEAE.
Ray flowers neither yellow nor red III. ASTEREAE.
4 FIELDIANA: BOTANY, VOLUME 24
TRIBE I. VERNONIEAE
By DOROTHY L. NASH
Annual or perennial herbs, or sometimes shrubs or small trees, rarely
subscandent; leaves usually alternate (in ours except Liabum); inflorescences spicate,
paniculate, often in scorioid cymes or corymbose; involucres cylindric to camp-
anulate, the phyllaries in pairs or more usually in 3 or more imbricate series;
receptacles flat or somewhat convex, epalaeceous; heads homogamous, consisting of
1-several-many disc flowers, these usually white or cyanic, never yellow (except
Liabum); anthers sagittate at the base, often with obtuse auricles; style branches
subulate or at least acute; achenes usually costate or striate; pappus composed of
scales or bristles.
The tribe in Guatemala consists of nine genera and a few more
than 40 species, of which two-thirds belong to the enormous and
complex genus Vernonia. Most species of the tribe occur from mid-
elevation down to the lowlands. None are conspicuous in the flora
and most are fairly wide ranging, as is to be expected from the
ecological situations in which they occur. A few are abundant as
weeds in disturbed situations and especially around old corrals
( Elephantopus ) .
Heads aggregate, forming glomerules.
Bracts subtending the glomerules complicate, each tipped with a short spine; style
branches short, almost obsolete Spiracantha.
Bracts subtending the glomerules neither complicate nor spinose; style deeply
bilobate Elephantopus.
Heads separate, not aggregate in glomerules.
Pappus bristles none, the achenes bearing only a persistent, cartilaginous, shallowly
lobate or entire crown.
Leaves sparsely strigillose beneath or glabrate; involucres hemispheric to
depressed-globose; heads about 50-flowered; corollas greenish or creamy
white Struchium.
Leaves white-tomentose beneath; involucres oblong-turbinate; heads 8-9-
flowered; corollas purple Harleya.
Pappus bristles and /or scales present.
Receptacle paleaceous Lepidonia.
Receptacle naked.
Plants aquatic; achenes about 10 mm. long Pacourina.
Plants terrestrial; achenes commonly 1.5-3 mm. long.
Lower leaf surfaces stellate-tomentose Piptocarpha.
Lower leaf surfaces not stellate-tomentose.
Leaves alternate; heads homogamous Vernonia.
Leaves opposite; heads often heterogamous Liabum.
ELEPHANTOPUS Linnaeus
References: Baker, C. F., A revision of the Elephantopeae,
Trans. Acad. Sci. St. Louis 12: 43-56. 1902; Gleason, H. A.,
Elephantopus L. in Vernonieae, N. Amer. Fl. 33: 106-109. 1922.
NASH: FLORA OF GUATEMALA 5
Erect perennial herbs, simple or sparsely branched, usually more or less sericeous
or villous; leaves alternate or often chiefly basal, sessile or the basal ones sometimes
short-petiolate, the margins entire, crenate, or dentate; inflorescences spicate,
corymbose, or sometimes paniculate, the heads homogamous, 1-5-flowered, without
rays, in dense, ovoid or globose glomerules, the clusters leafy-bracteate, mostly long-
pedunculate and openly corymbose; involucre of 2-4 decussate pairs of phyllaries, the
outer phyllaries shorter than the inner ones, the alternate pairs conduplicate;
receptacle flat or nearly so, naked; disc corollas white or purple, the tube slender, the
limb unequally 5-cleft; stamens 5, the anthers sagittate at the base, the auricles
obtuse; style branches subulate, minutely hispidulous or glabrate; achenes 10-costate
or 10-striate; pappus mostly of 5-30 bristles, these rigid, flat, and scalelike or
prolonged into terminal setae.
About 15 species, in tropical and temperate regions of both
hemispheres. Only three are in Guatemala, but four are treated
here, as one occurring in Chiapas, Mexico, may be expected in
Guatemala. Many species are much alike in general appearance, and
several reductions could probably be made without doing great
violence to taxonomic proprieties.
Pappus bristles dissimilar, composed of 10-15 bristles, 2 lateral ones longer than the
others and plicate at the apex, 2 straight ones almost as long, and several short,
scarious scales E. spicatus.
Pappus bristles all alike, straight.
Inflorescences appearing spicate, the glomerules of heads sessile, subtended by
triangular to lanceolate bracts; pappus bristles numerous (about 30).
E. angustifolius.
Inflorescences not spicate, the glomerules of heads pedunculate, subtended by
broad, ovate to cordiform bracts; pappus bristles 5-8.
Pappus bristles about 4 mm. long E. mollis.
Pappus bristles 6-8 mm. long E. tomentosus.
Elephantopus angustifolius Swartz, Prodr. Veg. Ind. Occ.
115. 1788. Elephantosis angustifolia DC. Prodr. 5: 87. 1836.
Orthopappus angustifolius Gleason, Bull. N. Y. Bot. Gard. 4: 238.
1906. Figure 1.
Open places, mostly in savannas, sometimes in pine forest, sea
level to 1,300 m.; Huehuetenango; Peten. Southern Mexico; British
Honduras; Honduras; El Salvador; Panama; West Indies; tropical
South America.
Erect plants arising from hard, woody roots, the stems simple, to about 1 m.
high, strigose with white hairs; basal leaves short-petiolate, the blades narrowly
oblanceolate, mostly 10-25 cm. long, 1.5-4 cm. wide, obtuse or subacute, long-
attenuate to the base, thinly strigose on both surfaces, the margins shallowly and
irregularly crenate, the cauline leaves linear to oblong; inflorescences spicate or with
a few short branches below, the clusters of heads crowded or remote, each head 4-
flowered; involucres about 1 cm. long; phyllaries 8, in 2 decussate pairs, acuminate,
6 FIELDIANA: BOTANY, VOLUME 24
glabrous or sparsely strigose, 7-11 mm. long; corollas white, 6-8 mm. long; achenes
brown, pubescent, about 2 mm. long; pappus of about 20-40 more or less uniform
bristles, pale brown or silver, 7-8 mm. long, attenuate from base to apex.
Elephantopus mollis HBK. Nov. Gen. & Sp. 4: 26. 1818. E.
hypomalacus Blake, Contr. Gray Herb. 52: 20. 1917. Ore/a de cache
(Jalapa; Quezaltenango). Figure 2.
Damp or dry thickets and fields, rocky hillsides, or often in pine
or pine-oak forest, sea level to 1,800 m.; Alta Verapaz; Baja
Verapaz; Chiquimula; Escuintla; Guatemala; Huehuetenango;
Izabal; Jalapa; Jutiapa; Peten; Quezaltenango; Retalhuleu; San
Marcos; Santa Rosa; Suchitepequez. Mexico; Honduras and El
Salvador to Panama; Cuba; tropical South America.
Erect perennials, the stems stout, mostly 30-70 cm. tall, simple below, sometimes
branched above, pilose or hirsute, sometimes naked and scapelike but usually sparsely
leafy below; leaves basal and cauline, or the cauline ones much reduced and
bractlike, sessile, clasping, the blades oblong to obovate or spathulate, rarely elliptic,
mostly 8-15 cm. long and 3-6 cm. wide, obtuse or acute, attenuate to the base, the
margins crenate or subentire, thinly pilose above and often scabrous, resin-dotted
beneath and often densely velutinous-pilose; glomerules of heads 1-2 cm. in diameter,
the peduncles short or elongated, the glomerules subtended by several broad, green,
sessile bracts, these mostly broadly ovate to cordiform, densely pubescent to almost
glabrous; phyllaries 8, in 4 decussate pairs, about 7 mm. long, paleaceous, with stiff
acuminate tips, strigose or almost glabrous; corollas white, pink, or purplish white;
achenes minutely pilose, 2.5-3.2 mm. long; pappus of 5-8 rigid bristles about 4 mm.
long, dilated below into a narrowly or broadly triangular base.
Widely distributed in Central America and very common in
some regions. This species differs only slightly from the East Indian
E. scaber L., which has larger achenes, mostly 3-4 mm. long. The
key characters usually employed in separating species are not
constant. I find that some specimens of E. scaber from Africa and
Asia have stem hairs no more appressed than ours; in fact, a few
have totally erect hairs, others merely ascending. Some East Indian
specimens have the undersurface of leaves softly tomentose (as in
the American E. tomentosus L.). Some with few hairs are a little
scabrous. Phyllaries of Asiatic specimens of E. scaber tend to be
more villous, but there are variations from glabrous to puberulent
to densely villous and long-ciliate. Pollen studies also show no
significant differences. It would appear, therefore, that there may be
biological and nomenclatural problems that cannot be solved by
studying only the New World specimens, as our E. mollis, E.
tomentosus, and E. carolinianus Willd. probably represent only a
part of the E. scaber complex.
NASH: FLORA OF GUATEMALA 7
Elephantopus spicatus Juss. ex Aubl. PL Guian. 808. 1775.
Pseudelephantopus spicatus Rohr, Skr. Nat. Selsk. Kjoebenhavn 2:
213. 1792. Distreptus spicatus Cass. Diet. Sci. Nat. 13: 367. 1819.
Matamoria spicata La Llav. et Lex. Nov. Veg. Descr. 1: 8. 1824. E.
spicatus var. roseum Klatt, Bull. Soc. Bot, Belg. 35: 279. 1896. Oreja
de cache (Jutiapa, Quezaltenango, Retalhuleu, Santa Rosa); ore/a
de cone/o (Guatemala, Suchitepequez). Figure 3.
Damp or wet thickets or open fields, often a weed in waste or
cultivated ground or on sandbars along streams, sea level to 1,600
m.; Alta Verapaz; Baja Verapaz; Chimaltenango; Chiquimula;
Escuintla; Guatemala; Huehuetenango; Jutiapa; Peten; Quezal-
tenango; El Quiche; Retalhuleu; Sacatepequez; Santa Rosa;
Suchitepequez; probably in all the lowland departments with the
possible exception of Totonicapan. Mexico; British Honduras to El
Salvador and Panama; West Indies; tropical South America;
naturalized in the Old World tropics.
Erect, stout plants, usually less than 1 m. tall, the stems appressed-hirsute or
glabrate, striate; leaves cauline, mostly obovate or oblong-obovate to spathulate, 5-15
cm. long, the uppermost chiefly lanceolate to linear and small, acute or obtuse, long-
attenuate to the base, the margins crenulate or entire, thinly appressed-pilose or
glabrate, the lowermost ones usually broadly spathulate, obtuse; spikes of heads few
or numerous, often forming large, open panicles; bracts when present linear or
narrowly oblong, slightly longer than the heads; involucres about 1 cm. high,
narrowly campanulate or cylindrical; phyllaries appressed, acutely acuminate, green,
pale-marginate, glabrous or sparsely pilose, often resinous-dotted; corollas white or
pale purple; achenes densely hirsute on the costae, resin-dotted in the intervals, 7-8
mm. long; pappus fulvous, of 10-15 bristles, 4-6.5 mm. long, with 2 long, stout lateral
ones plicate at the apex, exserted from the involucre, 2 straight ones almost as long,
and several short, scarious scales, all of them gradually dilated and fimbriate-ciliate
at the base.
Elephantopus tomentosus L. Sp. PL 1814. 1753. E. tomen-
tosus f. rotundatus Fern. Rhodora 38: 445. 1936.
Not reported from Guatemala, but may be expected there as it
occurs in Chiapas, Mexico, as well as in the United States.
Erect perennials, mostly 30-60 cm. tall, the stems more or less pilose; leaves
essentially basal, the cauline ones reduced and bractlike, the basal blades mostly 10-
30 cm. long, 3-10 cm. wide, narrowly or broadly obovate, or rarely elliptic, acute or
obtuse, long-attenuate to the base or narrowly cuneate, the margins commonly
crenate, pilose above, tomentose beneath; inflorescences corymbose, becoming
paniculate; glomerules of heads 1-2.5 cm. in diameter, subtended by broad, green,
sessile bracts, these mostly cordate, acute or obtuse, resin dotted and pilose;
phyllaries 8-11 mm: long, sharply acuminate, pubescent to pilose; corollas white or
purplish, about 8 mm. long; achenes pubescent, 4-5 mm. long, pappus of 5 bristles 6-8
mm. long, gradually dilated below into a triangular base.
8 FIELDIANA: BOTANY, VOLUME 24
HARLEYA Blake
Erect perennials, stoloniferous, the stems simple or branched; leaves alternate,
petiolate, the blades broad, penninerved, repand-dentate, glabrous above, covered
beneath with very dense, appressed, white tomentum; heads homogamous, eradiate,
8-9-flowered, subsessile in small, dense cymules, these aggregate at the ends of the
branches or in the axils of the upper leaves; involucres oblong-turbinate; phyllaries
multiseriate, graduated, erect, with only the apex becoming more or less reflexed,
cuspidate-acuminate; receptacle small, naked, flat, somewhat alveolate; corollas
purple, regular, equal, the limb 5-parted; anthers deeply sagittate at the base, the
auricles obtuse, not caudate; style branches subulate, hirtellous; achenes turbinate, 4-
5-angulate and often with 1-5 less distinct, intermediate costulae, glandular-papillose
between the costae; pappus coroniform, cartilaginous, pale, obscurely crenate.
The genus, composed of a single species, was named for one of
its collectors, Harley H. Bartlett.
Harleya oxylepis (Benth.) Blake, Journ. Wash. Acad. Sci. 22:
381, f. 1. 1932. Oliganthes oxylepis Benth. in Benth. & Hook. Gen.
PI. 2: 233. 1873. Figure 4.
Open, wet banks or fields, sometimes on shale slopes, at or a
little above sea level; Alta Verapaz; Peten. British Honduras;
Mexico (Tabasco).
Herbaceous or perhaps suffrutescent plants, to about 1 m. tall, the stems white-
tomentose above the middle, glabrate below; leaves short-petiolate, the blades
rhombic-ovate to lance elliptic, mostly 8-12 cm. long, 2.5-5 cm. wide, acute, acute or
attenuate to the base and decurrent on the petiole, glabrous above, white-tomentose
below; heads sessile or short-pedicellate, densely clustered; corolla purple or lavender,
the lobes about one-third as long as the tube, becoming revolute; involucres 9-10 mm.
high; phyllaries about 6-seriate, lanceolate, glabrous, erect, but the cuspidate apices
sometimes becoming more or less reflexed; achenes pale brown, scarcely 2 mm. long.
LEPIDONIA Blake
Erect herbs or suffrutescent plants; leaves alternate, petiolate, the blades
membranaceous, penninerved, densely whitish-tomentose beneath, the margins
serrulate; heads homogamous, discoid, rather large, few at the ends of the stems or
branches, the peduncles shorter than the leaves, each bearing 1-2 heads; involucres
hemispheric; phyllaries about 8-seriate, graduated, the outer ones small, ovate,
indurate below, unicostate, each bearing a short, herbaceous, lanceolate or ovate,
more or less squarrose appendage, the inner ones linear-oblong, thick-chartaceous, the
appendages ovate, subscarious, apiculate; receptacle broad, rounded, paleaceous, the
pales linear, stramineous, persistent; flowers all hermaphrodite and fertile, the
corollas funnelform, the limb 5-dentate; anthers sagittate at the base, the auricles
obtuse, the apical appendages ovate; style branches linear-subulate, hispidulous
dorsally; achenes obovoid, 5-6-angulate, glabrous, truncate; pappus bristles capillary,
deciduous, numerous, multiseriate, unequal, hispidulous-barbellate.
The genus consists of a single species.
NASH: FLORA OF GUATEMALA 9
Lepidonia paleata Blake, Journ. Wash. Acad. Sci. 26: 454. f. 1.
1936. Vernonia salvinae var. canescens Coult. Bot. Gaz. 16: 95.
1891. Figure 5.
Known only from the type, in dense, wet forest, 1,450 m., near
Coban, Alta Verapaz, Tuerckheim 583.
A shrub about 2 m. high, the stems densely lanate-pilose with brownish hairs;
leaves on petioles 1-2 cm. long, the blades obovate, 16-21 cm. long, 6-8 cm. wide,
acuminate, attenuate-cuneate at the base, the margins serrulate, thinly short-pilose
above, pale beneath, very densely and closely tomentose, long-pilose along the veins
with brownish (when dried) hairs, the lateral veins about 10 pairs; peduncles 4-6 cm.
long, ebracteate; involucres 7-8 mm. high, 13-15 mm. broad; outer phyllaries
pilosulous on the margins, the appendages densely pilosulous, the inner phyllaries
glandular and ciliate on the appendages; corollas 10.5-12 mm. long, bright violet;
pales linear-lanceolate, acuminate, obscurely ciliolate and sparsely strigillose, 5-7.5
mm. long; immature achenes olivaceous, glabrous, 1.8 mm. long; pappus stramineous,
1.5-3 mm. long.
LIABUM Adanson
References: P. A. Rydberg, Liabum and Sinclairia, N. Amer.
Fl. 34: 289-301. 1927; H. Robinson and R. D. Brettell, Tribal
revisions in the Asteraceae, III. A new tribe, Liabeae, Phytologia 25:
405-407. 1973.
Mostly shrubs or small trees, terrestrial or epiphytic, rarely herbaceous, simple or
branched, the stems often with arachnoid tomentum; leaves opposite, commonly
more or less triangular-ovate, rarely lobate, the margins entire, denticulate, or
serrate, the lower surfaces usually closely white arachnoid-tomentose; inflorescences
cymose or cymose-paniculate (in ours); heads usually pedicellate, heterogamous and
radiate or homogamous and discoid; involucres campanulate or hemispheric;
phyllaries multiseriate, imbricate, graduated; receptacle flat or nearly so, naked,
alveolate or fimbrillate; ray flowers 1-2-seriate, pistillate, fertile, the ligules yellow,
narrow, entire or tridentate; disc flowers hermaphrodite, fertile, the corollas regular,
yellow, the tube slender, the limb cylindric, usually deeply 5-cleft; anthers sagittate
at the base, the apical appendages broad and flat; style of the disc flowers hirtellous
on the outer surface to below the base of the linear-subulate branches; achenes
glabrous or pubescent, oblong-turbinate, sometimes slightly incurved; pappus usually
biseriate (in ours), the inner bristles scabrous or barbellate, the outer ones very short
or minute.
Liabum has most commonly been placed in the tribe
Senecioneae, following Bentham's disposition of it about a century
ago. Cassini, a specialist in the Compositae, had placed the genus in
the Vernonieae, establishing for it the section Liabeae, which
appears to us the most satisfactory disposition. Robinson and
Brettell have recently established the tribe Liabeae to contain the
genus.
10 FIELDIANA: BOTANY, VOLUME 24
Perhaps 100 species, in tropical America, with 12 in Guatemala.
Petioles broadly winged, greatly dilated and clasping at the base; plants herbaceous.
L. bourgeaui.
Petioles naked, at least below, not dilated at the base; plants shrubs or small trees.
Involucres 10-20 mm. high; phyllaries acuminate; achenes 4-5 mm. long.
Heads discoid L. tajumulcense.
Heads conspicuously radiate L. andrieuxii.
Involucres 4-8 mm. high; phyllaries obtuse or acute; achenes 1.5-3 mm. long.
Lower leaf surfaces essentially glabrous, minutely glandular-puberulent, or
sparsely tomentulose.
Heads discoid; inner phyllaries oblong-linear, rounded at the apex; achenes
glabrous or nearly so L. glabrum.
Heads radiate; inner phyllaries lanceolate, acute; achenes conspicuously
pubescent L. hypochlorwn.
Lower leaf surfaces densely and closely white-tomentose.
Heads discoid.
Involucres 4-5 mm. high.
Heads with about 6 flowers; achenes about 2.5 mm. long L. deamii.
Heads with 10-12 flowers; achenes about 1.3 mm. long L. dimidium.
Involucres 6-8 mm. high.
Achenes glabrous L. sublobatum.
Achenes hispidulous L. brachypus.
Heads radiate.
Ray flowers inconspicuous, the ligules 2-3 mm. long; disc corollas about 7
mm. long L. polyanthum.
Ray flowers conspicuous, the ligules (4-) 5-8 mm. long; disc corollas about 9
mm. long.
Phyllaries more or less brownish-tomentulose, the inner ones acute, or
obtuse and apiculate; achenes hispidulous L. vagans.
Phyllaries usually glabrate, rarely somewhat tomentulose, the inner ones
rounded at the apex; achenes glabrous L. discolor.
Liabum andrieuxii (DC.) Benth. & Hook. Gen. PL 2: 437.
1876. Vernonia andrieuxii DC. Prodr. 6: 16. 1836.
Rocky cliffs, 1,400 m., Huehuetenango. Mexico.
Weak shrubs, 1-2 m. high, the stems terete, more or less arachnoid-tomentose
and glandular-pilose; leaves opposite, the uppermost ones subsessile or short-
petiolate, the blades elliptic-lanceolate to angular-ovate, the remaining ones on
petioles 1-4 cm. long, the blades broadly angular-ovate or rhombic-ovate to cordiform
or sometimes almost reniform, sometimes sublobate, mostly 6-20 cm. long, 4-18 cm.
wide, acute or acuminate, the base rounded or almost truncate or subcordate and
then abruptly cuneate, the margins remotely dentate-serrate or denticulate, minutely
scabrous above, densely white-tomentose beneath; heads 2-several at the ends of the
branches, pedunculate, radiate, large, often 3 cm. high and 5 cm. across; involucres
broadly campanulate, 1-2 cm. high; phyllaries several-seriate, linear, long-acuminate,
often bent or twisted, more or less arachnoid-tomentose; ray flowers numerous, the
ligules orange, narrow, 20-30 mm. long; disc flowers numerous, the corollas orange,
NASH: FLORA OF GUATEMALA 11
11-15 mm. long, the tube very slender, 6-8 mm. long, the lobes 3-4 mm. long; achenes
a ppressed- pubescent, 4-5 mm. long; pappus bristles biseriate, the outer ones less than
1 mm. long, the inner ones 7-10 mm. long.
Liabum bourgeaui Hieron. in Ule, Verb. Bot. Ver. Brandenb.
48: 208. 1907. Papelillo (Huehuetenango). Figure 6.
Dense, wet, mixed forest or thickets or on open, damp or wet
banks, frequently along the borders of mountain streams, 300-2,000
m. Alta Verapaz; Escuintla; Huehuetenango; Quezaltenango; El
Quiche; Retalhuleu; San Marcos; Santa Rosa; Solola; Suchitepe-
quez; Zacapa. Southern Mexico; Honduras; Costa Rica; Panama.
Erect perennial herbs (perhaps sometimes suffrutescent at the base but
essentially herbaceous), mostly 0.5-1 m. high, rarely to 2 m., the stems usually simple,
stout, densely and closely white-tomentose; leaves opposite, on petioles to 5 cm. long,
the petioles broadly winged, much dilated at the base and amplexicaul, the leaf
blades ovate or broadly elliptic, mostly 10-20 cm. long and 4-13 cm. wide, acute or
acuminate, cuneate or abruptly contracted at the base and then decurrent on the
petiole, the margins finely and remotely denticulate, glabrous above, very densely
white-tomentose beneath, conspicuously triplinerved, the lateral nerves prolonged
almost to the apex of the blade; inflorescences cymose-paniculate, large, the heads
very numerous, on short, stout pedicels; involucres 8-9 mm. high; phyllaries 6-7-
seriate, puberulent and floccose-tomentose, narrowly lanceolate, acute or acuminate;
ray flowers inconspicuous, yellow, the tube very narrow, about 4 mm. long, the ligule
1-3 mm. long; disc flowers numerous, the corollas yellow to orange-yellow; achenes
little more than 1 mm. long, hispidulous; inner pappus bristles about 5 mm. long,
scabrous, the outer ones few and minute, or sometimes none.
This species has been reported from Guatemala as L.
asclepiadeum Sch. Bip. In Huehuetenango, the cottonlike pubes-
cence of the stems and leaves is sometimes employed as tinder in
starting a fire with flint and steel, a practice still continued in
remote regions.
Liabum brachypus (Rydb.) Blake, Journ. Wash. Acad. Sci. 22:
386. 1932. Sinclairia brachypus Rydb. N. Amer. Fl. 34: 299. 1927.
Known only from the type collection, dense, damp or wet
forest, 1,700 m.; Chiquimula, Volcan de Ipala, Pittier 1886.
Shrubs about 3 m. high, the branches stout, floccose-tomentose when young,
glabrate in age; leaves on slender, naked petioles 3-5 cm. long, the blades rhombic-
ovate, 9-17 cm. long, acuminate or long-acuminate, cuneate at the base and short-
decurrent on the petiole, glabrous above in age, very densely and closely white-
tomentose beneath, remotely and inconspicuously denticulate, triplinerved;
inflorescence dense, cymose-paniculate; heads discoid, 10-14-flowered, on stout or
slender, arachnoid-tomentulose pedicels; involucres 7-8 mm. high; phyllaries 4-5-
seriate, the outer ones short, ovate, the inner broadly linear, about 1 mm. wide,
obtuse, more or less arachnoid-tomentose on the outermost phyllaries, glabrate in
12 FIELDIANA: BOTANY, VOLUME 24
age; corollas yellow, about 6 mm. long; achenes sparsely hispidulous, 1-1.5 mm. long;
inner pappus bristles brownish, about 5 mm. long, the outer ones minute.
Liabum deamii Robinson & Bartlett, Proc. Amer. Acad. 43:
60. 1907. L. subglandulare Blake, Contr. U. S. Nat. Herb. 24: 31.
1922. Sinclairia subglandularis Rydb. N. Amer. Fl. 34: 298. 1927. S.
deamii Rydb. torn. cit. 299.
Damp or wet thickets and along rocky stream banks, near sea
level to 600 m.; Alta Verapaz; Peten; Zacapa (type from Gualan,
along the Rio Motagua, Deam 194). British Honduras; Honduras;
El Salvador.
Erect or sprawling shrubs, woody vines, or weak trees to about 6 m. high, the
branches terete, more or less arachnoid-tomentose and short-hirsute with brownish
hairs, sometimes glabrate in age; leaves (often absent at anthesis) on petioles 1.5-4
cm. long, the blades lance-ovate to broadly rhombic, mostly 5-10 cm. long, 2.5-11.5
cm. wide, acuminate or abruptly short-acuminate, broadly cuneate at the base or
sometimes rather abruptly short-decurrent on the petiole, the margins entire or
remotely and inconspicuously denticulate, the upper surface hispidulous or glabrate,
the lower one densely white-tomentose, triplinerved; inflorescences broadly thyrsoid
to paniculate, mostly 10-20 cm. high, 5-15 cm. broad; heads discoid, numerous, on
short pedicels, these more or less glandular, pubescent with short, multiseptate hairs,
and with arachnoid tomentum; involucres about 4 mm. high; phyllaries 3-4 -seriate,
more or less tomentulose or pubescent, the outer ones short, broadly ovate, the inner
ones oblong, obtuse or rounded at the apex; flowers about 6, the corollas yellow, 6-8
mm. long; achenes about 2.5 mm. long, hispidulous; pappus biseriate, the inner
bristles 6-7 mm. long, stramineous or reddish, the outer squamellae less than 1 mm.
long.
The alleged difference between L. deamii and L. subglandulare
is that the former has rather dense arachnoid indument while the
latter has multiseptate hairs; this appears to be a difference in
specimens rather than species. Some specimens have fewer
multiseptate hairs and more arachnoid hairs; in others the
multiseptate hairs are dominent but the arachnoid indument is also
present.
Liabum dimidium Blake, Journ. Wash. Acad. Sci. 22: 385.
1932.
Known only from the type, Tikal, Peten, Bartlett 12602.
A subscandent shrub as much as 6 m. long, the branches more or less arachnoid-
tomentose, glabrate in age, those of the inflorescence sordid-pilosulous with short,
spreading hairs and with arachnoid tomentum; leaves on petioles 1.5-3 cm. long, the
blades ovate to triangular-ovate, mostly 9-12 cm. long, 6-9 cm. wide, acute to
acuminate, broadly cuneate at the base or rounded and abruptly cuneate, the
margins serrate-denticulate, glabrate above, white-tomentose beneath; panicles
NASH: FLORA OF GUATEMALA 13
pyramidal, as much as 28 cm. broad, 20-30 cm. long; heads partly sessile, partly
pedicellate, discoid; involucres about 5 mm. high; phyllaries 4-5-seriate, the outer
ones sordid-pilosulous, ovate, short, acute, ciliate, the inner ones linear-oblong, acute
or subacute; flowers 10-12, the corollas yellow, 7-8 mm. long; achenes about 1.3 mm.
long, densely hispidulous; pappus yellowish white, the inner bristles about 7 mm.
long, the outer squamellae about 1.5 mm. long.
Liabum discolor (Hook. & Arn.) Benth. & Hook, ex Hemsl.
Biol. Centr. Amer. Bot. 2: 232. 1881. Sinclairia discolor Hook. &
Arn. Bot. Beechey Voy. 433. 1841.
Damp or wet, dense or open forest or thickets, 300-2,500 m.;
Alta Verapaz; Guatemala; El Progreso; Quezaltenango; San
Marcos; Sacatepequez. Mexico.
Shrubs of 1-4 m., usually terrestrial, rarely epiphytic, often sprawling or
subscandent, the branches stout, floccose-tomentose at first but soon glabrate; leaves
on slender, naked petioles 2-4 cm. long, the blades broadly rhombic-ovate to lance-
ovate, mostly 8-15 cm. long, acute or acuminate, cuneate or broadly cuneate at the
base, not or scarcely decurrent on the petioles, the margins denticulate or subentire,
glabrous above or nearly so, covered beneath with dense, white tomentum, trinerved
or triplinerved; inflorescences large, cymose-paniculate; heads radiate, very numer-
ous; involucres about 8 mm. high; phyllaries 4-5-seriate, usually glabrous or glabrate,
rarely more or less arachnoid-tomentose, striate, the outer ones very short, ovate,
subacute, the inner elongated, linear-oblong, rounded at the apex; ray flowers
few, the ligules 4-7 mm. long, yellow, 3-4-dentate at the apex; disc corollas yellow,
about 9 mm. long; achenes about 10-striate, glabrous, 1.5 mm. long; pappus biseriate,
the inner bristles yellowish, about 5 mm. long, the outer squamellae 1-1.5 mm. long.
Liabum glabrum Hemsl. Biol. Centr. Amer. Bot. 2: 232. 1881.
Sinclairia glabra Rydb. N. Amer. Fl. 34: 297. 1927.
Damp or rather dry thickets or roadside hedges, 500-1,800 m.;
Chimaltenango; Escuintla; El Quiche; Santa Rosa; Solola. South-
ern Mexico.
Shrubs or trees, sometime 6 m. high, with rather few, irregular branches, these
thick, glabrate in age; leaves opposite or sometimes ternate, on slender petioles 2-5
cm. long, the blades lanceolate or triangular-ovate, mostly 5-15 cm. long, acuminate
to long-acuminate, cuneate at the base or abruptly contracted and long-decurrent on
the petiole, but not to the base, the margins denticulate, triplinerved, almost glabrous
on both surfaces or sometimes puberulent or sparsely tomentulose beneath;
inflorescences cymose-paniculate, the panicles more or less pyramidal, large and
broad; heads discoid, sessile or short-pedicellate; involucres about 8 mm. high,
glabrous or nearly so; phyllaries 3-5-seriate, striate, the outer ones short, ovate,
subacute or obtuse, the inner ones oblong- linear, elongated, rounded at the apex; disc
corollas bright yellow, 7-8 mm. long; achenes glabrous or essentially so, 10-striate;
pappus biseriate, the inner bristles stramineous, 5-6 mm. long, the outer squamellae
to 1 mm. long.
14 FIELDIANA: BOTANY, VOLUME 24
Liabum hypochlorum Blake, Contr. Gray Herb. 53: 27. 1918.
Sinclairia hypochlora Rydb. N. Amer. Fl. 34: 301. 1927.
Damp or wet, mixed forest, 500-900 m.; Quezaltenango;
Retalhuleu (type from San Felipe, Holway 703)', Suchitepequez.
Mexico (Chiapas).
Small, sometimes epiphytic shrubs about 2 m. high, or woody vines as much as
12 m. long, forming dense tangles over dead trees, the stems sometimes 5 cm. thick,
the young branches stout, densely glandular and sparsely pilose; leaves on petioles 2-
6 cm. long, the blades broadly triangular-ovate to almost cordiform, mostly 10-16 cm.
long, 8-12 cm. wide, abruptly acute or short-acuminate, broadly cuneate or rounded
and abruptly cuneate at the base, the margins remotely denticulate or almost entire,
triplinerved, essentially glabrous on both surfaces but minutely glandular-punctate
beneath and sometimes sparsely pilosulous along the costae and veins; panicles
mostly 12-15 (-20) cm. broad; heads radiate, about 15-flowered, on glandular, more or
less short-tomentose pedicels; involucres 4-5 mm. high; phyllaries 4-seriate, glandular -
puberulent and pubescent to pilose, acute, the outer ones ovate, short, the inner ones
lanceolate; ray flowers 5, the ligules yellow, 3-4 mm. long; disc flowers 8-10, yellow,
about 8 mm. long; achenes 2.5 mm. long, about 5-angulate, pubescent; inner pappus
bristles about 7 mm. long, the outer squamellae subulate, minute.
Liabum polyanthum Klatt, Bull. Soc. Bot. Belg. 31, pt. 1: 209.
1892. Sinclairia polyantha Rydb. N. Amer. Fl. 34: 299. 1927.
Dense, wet forest, at or a little above sea level; Izabal. Mexico;
British Honduras; Costa Rica.
Small, epiphytic shrubs or woody vines as much as 12 m. long and climbing over
trees, the branches stout, terete, when young glandular and somewhat arachnoid-
tomentulose; leaves on petioles 2-4 cm. long, the blades rhombic-ovate, mostly 9-14
cm. long, abruptly acute or acuminate, more or less cuneate at the base, glabrous
above, densely white-tomentose beneath; panicles 15-30 cm. broad, the branches
glandular and puberulent; heads inconspicuously radiate; involucres 5-7 mm. high,
phyllaries about 4 -seriate, the outer ones ovate, short, acute, more or less
tomentulose, the middle ones acute, the inner ones linear-oblong, obtuse, glabrate;
ray flowers inconspicuous, 3-5, the ligules yellow, 2-3 mm. long; disc flowers 6-15,
yellow, about 6 mm. long; achenes 2.5 mm. long, appressed-pubescent; inner pappus
bristles about 6 mm. long, yellowish or brownish white, the outer squamellae minute,
subulate.
The achenes were originally described as glabrous, but those of
the isotype I examined are hispidulous.
Liabum sublobatum Robinson, Proc. Amer. Acad. 51: 539.
1916. L. glabrum var. hypoleucum Greenm. Proc. Amer. Acad. 32:
294. 1897. Sinclairia sublobata Rydb. N. Amer. Fl. 34: 297. 1927. S.
hypoleuca Rydb. I.e. Ivoy (Santa Rosa).
Wet to dry, brushy or grassy slopes, or in oak-pine forest, 240-
2,200 m.; Alta Verapaz; Chimaltenango; Chiquimula; Guatemala;
NASH: FLORA OF GUATEMALA 15
Quezaltenango; Retalhuleu; Sacatepequez; Santa Rosa; Solola
(type from San Lucas Tollman, Holway 179)', Zacapa. Southern
Mexico; El Salvador; Honduras; Nicaragua.
Shrubs or weak trees, 3-8 m. high, the branches usually few, stout, soon glabrate;
leaves on petioles mostly 3-6 cm. long, the blades rather thick, usually more or less
rugose, rhombic-ovate to lanceolate, mostly 8-13 cm. long, acuminate to long-
acuminate, sometimes somewhat hastate-lobate near the base, cuneate at the base,
the margins denticulate to irregularly serrate or almost entire, glabrous above, more
or less densely and closely tomentose beneath, triplinerved; panicles usually large,
often pyramidal, 6-30 cm. long; heads usually numerous, discoid, pedicellate or
subsessile; involucres about 8 mm. high, glabrous or minutely puberulent, or with
some arachnoid tomentum at the base of the outer phyllaries; phyllaries 4-seriate,
the outer ones ovate, subacute, the inner ones oblong, obtuse to rounded at the apex,
sometimes obscurely ciliate; disc flowers 6-8, the corollas yellow, about 8 mm. long;
achenes about 2 mm. long, glabrous; pappus bristles about 6 mm. long, dirty white,
the outer squamellae less than 2 mm. long.
Liabum tajumulcense Standl. & Steyerm. Field Mus. Bot. 23:
27. 1943.
Known only from the type, San Marcos, moist thickets
bordering a stream, barranco southwest of Tajumulco,
northwestern slopes of Volcan de Tajumulco, 2,300-2,500 m.,
Steyermark 36453.
A shrub 5 m. high with very thick, pale branches, the young branches arachnoid-
tomentose; leaves unknown, deciduous; panicles dense, thyrsoid, the branches densely
ochraceous-tomentose; heads large, numerous, discoid, sessile or on short, stout
pedicels; involucres broadly campanulate, 12-13 mm. high, 17-20 mm. broad,
phyllaries 4-seriate, 5-13 mm. long, ovate-lanceolate to linear- lanceolate, acuminate,
ochraceous-lanate; corollas numerous, yellow, 12-13 mm. long; immature achenes 5-
costate, about 4 mm. long, densely hirsutulous: inner pappus bristles 35-40, dirty
white, 10-11 mm. long, the outer scales minute.
This plant is like many other members of the genus in Central
America, in having deciduous leaves. The shrubs or trees are usually
leafless throughout the dry season.
Liabum vagans Blake, Brittonia 2: 354. 1937. Gamusa
(Jalapa).
Dense, wet, mixed forest, sometimes in pine forest, 900-2,700 m.;
Chimaltenango; Guatemala; Jalapa; Quezaltenango; El Quiche
(type collected along trail between Nebaj and Aguacatan, Skutch
1913); San Marcos.
Erect or subscandent shrubs to about 4 m. high, the branches stout, thick,
sordid-tomentulose and brown-pilose; leaves on slender petioles 3-7 cm. long, the
blades broadly ovate to broadly rhombic-ovate or sometimes almost cordiform,
16 FIELDIANA: BOTANY, VOLUME 24
mostly 10-15 cm. long, 8-14 cm. wide, acute or rather abruptly short-acuminate,
subtruncate to broadly cuneate at the base and short-decurrent on the petiole, the
margins denticulate or almost entire, 3-nerved or triplinerved, at first thinly
arachnoid-tomentose above, soon green and glabrate, covered beneath with a very
dense, close, white tomentum; panicles large, convex, 12-20 cm. wide, sordid-
tomentose and pilose with multiseptate, brownish hairs, the pedicels 2-6 mm. long;
involucres 6-8 mm. high; phyllaries about 4-seriate, brownish-tomentulose, at least
the inner ones about 1.5 mm. wide and acute, or if obtuse, usually minutely apiculate
at the apex; ray flowers 5-7, the ligules bright yellow, 7-8 mm. long; disc flowers 4-6,
the corollas about 9 mm. long; achenes oblong, hispidulous, 2-2.5 mm. long; pappus
bristles yellowish white, 5.5-7 mm. long, the outer scales 1-1.5 mm. long.
Skutch reports that in this, and in some other species, the sap
is milky.
PACOURINA Aublet
Erect, aquatic herbs, essentially glabrous; leaves alternate, amplexicaul or
decurrent on the petiole, the blades penninerved, spinose-dentate; heads
homogamous, large, sessile, solitary in the axils of the upper leaves or opposite the
leaves; involucres hemispheric to depressed-globose; phyllaries multiseriate, imbricate,
the apices spinulose and spreading, the margins scarious, the outer phyllaries
gradually shorter; receptacle flat, naked; corollas regular, the tube slender, dilated at
the base, the limb narrow, the 5 lobes spreading, reflexed, becoming revolute; anthers
sagittate at the base, the auricles subobtuse; style branches subulate, hirtellous;
achenes linear, 10-costate, crowned by an entire or denticulate, cartilaginous ring;
pappus bristles numerous, short, multiseriate, caducous.
Two species are known, one in Central America.
Pacourina edulis Aubl. PL Guian. 800, pi. 316. 1775. Figure 7.
Growing in water 50-100 cm. deep, in lake margins, 500 m.;
Jutiapa (Lago de Atescatempa, Steyermark 31876). Nicaragua;
Dominican Republic; tropical South America.
Plants coarse and stout, 1-2 m. tall, the stems hollow, mostly 1-3 cm. thick, the
entire plant glabrous or very sparsely pubescent; leaves sessile, the blades narrowly
oblong to broadly elliptic, oblanceolate, or ovate, mostly 15-20 cm. long, 5-8 cm. wide,
deeply and irregularly serrate or runcinate-dentate with spine-tipped teeth, rarely
subentire, acute or acuminate, narrowed below into a broad, petioliform base 2-3 cm.
long; heads mostly in the upper leaf axils or opposite them, about 3 cm. broad, the
flowers roseate or purplish; phyllaries broadly ovate-oblong; achenes linear, about 1
cm. long, glandular.
PIPTOCARPHA R. Brown
Reference: H. A. Gleason, N. Amer. Fl. 33: 103-104. 1922.
Erect or scandent shrubs or rarely, trees, the pubescence stellate or lepidote;
leaves alternate, penninerved, the margins mostly entire; heads small, homogamous,
NASH: FLORA OF GUATEMALA 17
3-20-flowered, aggregate in axillary or terminal corymbs, umbels, or panicles, or
sessile in axillary clusters, the flowers hermaphrodite; involucres ovoid or
campanulate; phyllaries imbricate in several series, the outer ones smaller, the inner
ones often deciduous with the achenes; receptacle convex, naked; corollas regular,
tubular, the limb 5-lobate; anthers caudate or subcaudate at the base; style minutely
bifid (in ours); achenes truncate, 10-costate; pappus bristles biseriate, the inner ones
capillary, the outer ones short and inconspicuous, unequal, sometimes absent.
About 40 species, in tropical America. Only one is found in
continental North America.
Piptocarpha chontalensis Baker in Mart. Fl. Bras. 6 (2): 132.
1873. P. costaricensis Klatt, Bull. Soc. Bot. Belg. 31 (1): 184. 1892.
Figure 8.
Wet thickets or mixed forest, sea level to 350 m.; Alta Verapaz;
Huehuetenango; Izabal. British Honduras to Costa Rica.
Erect or arching shrubs to about 3 m. tall, or often scandent and more than 6 m.
long, or rarely trees 8-9 m. tall, the younger branches densely and closely ochraceous-
tomentose; leaves on petioles 1-1.5 cm. long, the blades subcoriaceous, oblong-ovate
or elliptic to broadly ovate, mostly 10-20 cm. long, 5-12 cm. wide, acute or abruptly
short-acuminate, obliquely rounded at the base, the margins entire or obscurely and
remotely denticulate, in age glabrous and lustrous above but stellate-tomentulose
when young, very densely and closely stellate-tomentose beneath; heads 4-8-flowered,
numerous, crowded in axillary, corymbiform inflorescences little longer than the
petioles; involucres ovoid, 2-4 mm. high; phyllaries closely imbricate, appressed,
ovate, acute to rounded at the apex, the outer ones tomentose, the inner ones
glabrous except at the tomentose apex, deciduous; corolla limb 5-lobate, the lobes
recurving in age; achenes about 3 mm. long, glabrous or sparsely pubescent, 10-
costate; pappus white, the inner bristles 6-7 mm. long, the outer ones 1-2 mm. long.
SPIRACANTHA HBK.
Reference: H. A. Gleason, Spiracantha HBK. in Vernonieae, N.
Amer. Fl. 33: 110. 1922.
Erect annual, or perhaps sometimes more enduring, herbs; leaves alternate,
short-petiolate, broad, penninerved, white-tomentose beneath, the margins denticu-
late or subentire; heads very small, 1-flowered, numerous, aggregate in globose,
headlike glomerules, each head subtended by a coriaceous, spinose bract, each
glomerule subtended by 3-4 foliaceous, complicate bracts; involucres narrow;
phyllaries 2 (5), complicate, mucronate, unequal; receptacle minute; corolla regular,
the limb 4-5-cleft; anthers sagittate at the base, the auricles short, obtuse; style
almost simple, appearing subulate at the apex, the branches almost obsolete,
hirtellous; achenes slightly compressed, obscurely 5-striate, glabrous; pappus of
numerous stout, erect, unequal squamellae.
The genus consists of a single species.
Spiracantha cornifolia HBK. Nov. Gen. & Sp. 4: 29. 1818.
Figure 9.
18 FIELDIANA: BOTANY, VOLUME 24
Brushy hillsides, wooded, rocky banks, or weedy fields, 150-400
m.; Santa Rosa. Yucatan Peninsula of Mexico; British Honduras;
El Salvador; Nicaragua: Costa Rica; Panama; Colombia.
Erect plants to about 50 cm. tall, frequently much branched, the stems often
reddish or purplish, long-strigose; leaves short-petiolate, the blades ovate to elliptic or
oblong-ovate, mostly 2-6.5 cm. long, acute at each end, the margins entire or
minutely denticulate, thinly long-pilose or almost glabrous above, closely and densely
white-tomentose beneath; glomerules of heads long-pedunculate, subtended at the
base by several broad, leaflike bracts, these complicate, arachnoid-tomentose
beneath, tipped with a short spine; bracts subtending the heads oblong, rounded at
the apex and bearing a stout, squarrose spine; heads 3-4 mm. high, almost concealed
by the bracts; phyllaries linear, scarious, acuminate, 1-costate, about 3 mm. long,
pilose at the base, otherwise glabrous; corollas purple; achenes about 2 mm. long.
These plants are rather frequent on the low hills of the Pacific
slope in Santa Rosa, but have not been observed elsewhere in
Guatemala.
STRUCHIUM P. Browne
Reference: H. A. Gleason, N. Amer. Fl. 33: 48. 1922.
Erect annuals of wet soil, simple or branched, with short internodes, almost
glabrous; leaves alternate, petiolate, the blades penninerved, the margins coarsely
serrate or almost entire; heads homogamous, solitary or glomerate in the leaf axils;
involucres hemispheric to depressed- globose; phyllaries numerous, multiseriate,
imbricate, graduated, scarious or scarious-marginate; receptacle slightly convex,
naked; corollas tubular, greenish white to creamy white, commonly 3-4-lobate;
anthers sagittate at the base, the auricles attenuate, not caudate; style branches
subulate, hirtellous; achenes 3-4-angulate and costate, glabrous or minutely
glandular, truncate at the apex, bearing a persistent, cartilaginous, shallowly lobate
or entire crown.
The genus consists of a single species.
Struchium sparganophorum (L.) Kuntze, Rev. Gen. 366.
1891. Ethulia sparganophora L. Sp. PI. ed. 2. 1171. 1763.
Sparganophorus vaillantii Crantz, Inst. Herb. 1: 261. 1766. Figure
10.
Wet thickets or forest, often a weed in banana plantations, at
or little above sea level; Izabal; Peten. Southern Mexico; British
Honduras, along the Atlantic coast to Panama; West Indies;
tropical South America; naturalized in Africa.
Erect herbs, commonly 40-60 cm. tall, the stems simple or much branched, stout,
terete, sparsely short-pilose or almost glabrous; leaves short-petiolate, the blades thin,
mostly oblanceolate or elliptic, 5-12 cm. long, acuminate, attenuate to the base,
conspicuously or obscurely and coarsely serrate, very sparsely and inconspicuously
strigillose or glabrate, punctate on both surfaces; heads about 50-flowered; involucres
NASH: FLORA OF GUATEMALA 19
4-5 mm. high; phyllaries appressed, with broad, pale margins, abruptly contracted
into a short, spinulose tip; achenes oblong, 1.5-2 mm. long, the crown whitish, about
half as long as the achene.
VERNONIA Schreb.
References: H. A. Gleason, A revision of the North American
Vernonieae, Bull. N. Y. Bot. Gard. 4: 144-243. 1906; Vernonia and
Eremosis, N. Amer. Fl. 33: 52-101. 1922. S. B. Jones, Jr., Revision of
Vernonia sect. Eremosis (Compositae) in North America, Brittonia
25: 86-115. 1973.
Annual or perennial herbs, shrubs, or trees, rarely subscandent; leaves alternate,
the blades narrow or broad, the margins serrate, dentate, crenate, or entire; stems
usually leafy and branched, at least above; heads homogamous, 1-many-flowered,
often disposed in scorpioid cymes, these terminal or arising in the upper leaf axils, the
inflorescence sometimes paniculate or corymbiform, or rarely composed of solitary
heads; flowers perfect; involucres campanulate to subcylindrical; phyllaries laxly or
closely imbricate in few-many series, the inner ones longer; receptacle flat or
somewhat convex; corolla regular, white, pink, or purple, the limb 5-cleft; style
branches elongated, acute; anthers sagittate at the base; achenes more or less
cylindrical, usually costate; pappus biseriate, the outer series very short, of scales or
bristles, the inner bristles capillary, terete or slightly flattened, as long as, or longer
than, the involucre.
Veronia is one of the largest genera of Compositae, with more
than 600 reported species, sometimes freely hybridizing, chiefly
tropical and most numerous in South America, a few extending into
temperate North America, with 20 in Guatemala, one in Chiapas,
Mexico, and two in British Honduras, making a total of 23 species
included here. The genus also occurs in Asia and Africa. One Old
World species, V. cinerea (L.) Lessing, is treated here, as it has
become well established in many parts of the American tropics, has
been collected frequently in British Honduras, and may be expected
in Guatemala.
Heads few, usually large, the involucres 1.5-2 cm. high.
Phyllaries red-purple, almost glabrous or somewhat puberulent, the margins entire
or only obscurely denticulate; corollas 20-28 mm. long; pappus 5-8 (10) mm.
long V. salvinae.
Phyllaries greenish, densely appressed-scaberulous, the margins ciliate, more or less
fimbriate and denticulate; corollas 15-17 mm. long; pappus 2.5-4 mm. long.
V. corae.
Heads often very numerous, relatively small, the involucres not more than, and
usually less than, 1 cm. high.
Cymes of inflorescences scorpioid; flowers commonly 15-40 in each head.
Involucres about 10 mm. high.
Inner phyllaries oblong, the apices rounded, obtuse, or emarginate; corollas
white or pinkish V. tortuosa.
20 FIELDIANA: BOTANY, VOLUME 24
Inner phyllaries linear, the apices attenuate to subulate-spinose; corollas
purple V. argyropappa.
Involucres 2-8 mm. high.
Bracts subtending the heads all large and leaflike.
Leaves thinly puberulent and glandular-punctate above, closely pilose or
glabrate beneath and glandular-punctate; involucres thinly puberulent
(British Honduras) V. ctenophora.
Leaves papillose-scabrous above, laxly long-pilose beneath, not glandular-
punctate on either surface; involucres more or less arachnoid-tomentose.
V. acilepis.
Bracts subtending the heads mostly small, commonly linear or subulate, or
only those subtending the lowermost heads of the inflorescence leaflike.
Involucres (6-)7-8 mm. high V. medialis.
Involucres 2-5 (-6) mm. high.
Middle phyllaries with spinose tips.
Leaf blades ovate, obovate, oblong, or elliptic, commonly 2-4 times
longer than wide; involucres 4-5 mm. high.
Upper leaf surfaces scabrous; lower leaf surfaces more or less pilose,
with appressed or lax hairs V. canescens.
Upper leaf surfaces densely hirsute-sericeous; lower leaf surfaces
densely velutinous-tomentose, with matted hairs V. mollis.
Leaf blades mostly narrowly lanceolate or linear-lanceolate, commonly
6-10 times longer than wide; involucres 3-4 mm. high.
V. canescens var. pilata.
Middle phyllaries acute, acuminate, or sometimes obtuse, but never with
spinose tips.
Pappus white; heads crowded on the cymes (British Honduras).
V. scorpioides.
Pappus fulvous; heads not crowded on the cymes.
Inner phyllaries ovate-lanceolate, 3-4 mm. long.
Leaf blades mostly 2.5-7 cm. wide; usually densely tomentose
beneath V. deppeana.
Leaf blades mostly 1.5-3 cm. wide; more or less puberulent or
pubescent beneath but not tomentose V. patens.
Inner phyllaries linear-oblong and often somewhat constricted near
the apex, 5-8 mm. long.
Lower leaf surfaces densely pilose; cymes mostly 15-30 cm. long;
heads 17-25-flowered V. polypleura.
Lower leaf surfaces glabrous or with some pubescence, mostly
confined to costae and veins; cymes mostly 5-10 cm. long;
h°ads about 15-flowered V. tenetta.
Cymes of inflorescences not scorpioid, commonly forming either corymbiform,
hemispheric, or pyramidal panicles; flowers commonly 1-14 in each head
(about 20 only in the introduced V. cinerea).
Flowers 9-14 (-20) in each head.
Plants annual herbs; leaf blades mostly 2-4 cm. long (introduced) V. cinerea.
Plants shrubs; leaf blades mostly 4-12 cm. long.
Heads often 5-10 on a common peduncle; involucres about 8 mm. high, the
phyllaries densely white- or grayish-tomentose; achenes glabrous.
V. mima.
NASH: FLORA OF GUATEMALA 21
Heads usually only 2-3 on a common peduncle; involucres 4-5 mm. high, the
phyllaries glabrous, or if tomentulose not densely so, greenish, with
prominent purplish markings; achenes glandular-pubescent (Mexico:
Chiapas and Oaxaca) V. oaxacana.
Flowers 1-6 (-7) in each head.
Lower leaf surfaces white or grayish, with dense tomentum, the hairs matted.
V. leiocarpa.
Lower leaf surfaces green, glabrate, or if tomentose, not densely so, the hairs
mostly straight, not matted.
Heads 1-flowered V. angusta.
Heads 3-6-flowered.
Involucres 8-13 mm. high V. shannonii.
Involucres 4-6 mm. high.
Heads disposed in globose glomerules V. standleyi.
Heads not disposed in globose glomerules.
Leaf blades mostly 4-6 cm. wide; heads 5-6-flowered; achenes glabrous
or very minutely glandular V. heydeana.
Leaf blades mostly 1.5-4 cm. wide; heads commonly 3-flowered;
achenes pilose, usually densely so V. triflosculosa.
Vernonia acilepis Benth. in Oerst. Vid. Medd. Kjoeb. 1852: 68.
1852.
Damp or dry, open or brushy, often rocky plains and hillsides,
sometimes in pine forest, 600-1,500 m.; Chiquimula; Guatemala;
Jutiapa. Mexico (Chiapas); El Salvador; Nicaragua.
Erect annuals, sometimes as much as 1 m. tall but usually much lower, simple or
sparsely branched, the slender stems densely pilose with short, ascending hairs; leaves
short-petiolate or sessile, the blades ovate to oblong-lanceolate, mostly 3-6 cm. long,
acute or acuminate, the base cuneate, the margins entire or serrate, papillose-
scabrous above, rather densely pilose beneath with long, spreading hairs;
inflorescence simple or of a few loosely spreading, scorpioid cymes, each bearing 4-8
remote, secund, sessile heads; bracts foliaceous, oblong or lanceolate; heads about 15-
flowered; involucres 7-8 mm. high; phyllaries erect or suberect, narrowly lanceolate,
with spinose, subulate tips and scarious, pinkish margins, arachnoid-tomentose;
corollas pink or lavender; achenes about 2 mm. long, densely pilose; pappus almost
white, the inner bristles about 5 mm. long.
Vernonia angusta (Gleason) Standl. Field Mus. Bot. 11: 276.
1936. Eremosis angusta Gleason, N. Amer. Fl. 33: 98. 1922.
From 125-1,100 m.; Jalapa; Zacapa (type from Gualan,
Kellerman 6132).
Shrubs or small trees to 5 m. tall or more, the branches densely tomentulose;
leaves sessile or on petioles 5-10 mm. long, the blades elliptic-lanceolate, to about 7
cm. long and 2 cm. wide, acute at the apex and at the base, the margins entire,
minutely scabrous-puberulent above, thinly tomentulose and glandular-punctate
beneath; inflorescences about 10 cm. broad, the heads numerous, crowded, sessile or
short-pedicellate, 1-flowered; involucres narrowly cylindric, 6-8 mm. high; phyllaries
22 FIELDIANA: BOTANY, VOLUME 24
yellowish, the outer ones thinly tomentulose, ovate to rounded, obtuse or rounded at
the apex, minutely cuspidate, the inner ones elongated, acute, scarious-marginate;
achenes dark, about 2 mm. long, pilose; pappus white, the inner bristles about 6 mm.
long.
Apparently rare, not represented by recent collections.
Vernonia argyropappa Buck, Index DC. Prodr. 2: V. 1840. V.
poeppigiana A. DC. in DC. Prodr. 5: 55. 1836, not DC. Prodr. 5: 20.
1836. V. friedrichsthalii Sch. Bip. ex Ekman, Arkiv Bot. 13 (15): 40,
41. 1914, nomen.
Damp or wet fields or thickets, sometimes in marshes or in pine
savannas, near sea level to 1,400 m.; Alta Verapaz; Izabal; Peten;
Santa Rosa. Southern Mexico; British Honduras; Honduras to
Costa Rica; tropical South America.
Erect annuals, 0.5-1.5 m. tall, simple or sparsely branched, sordid-pilose with
appressed or ascending, often long hairs; leaves sessile or nearly so, the blades mostly
oblong-lanceolate, 6-12 cm. long, 1.5-3 cm. wide, acute or acuminate, cuneate or
obtuse at the base, papillose-scabrous above, appressed-pilose and glandular-punctate
beneath, the margins entire or minutely serrulate; cymes scorpioid, few or numerous,
often forming large panicles, the branches bearing 4-10 remote, secund, sessile heads,
2 1-34 -flowered; bracts similar to the leaves but smaller; involucres campanulate or
hemispheric, 8-10 mm. high; phyllaries linear, erect, sparsely pilose, attenuate to a
subulate, spinose tip; corollas purple or lavender; achenes about 3 mm. long,
hispidulous; pappus white, the inner bristles 6-8 mm. long.
Vernonia canescens HBK. Nov. Gen. & Sp. 4: 35, t. 317. 1820.
V. bullata Benth. in Oerst. Vid. Medd. Kjoeb. 67. 1853. Cacalia
bullata Kuntze, Rev. Gen. 969. 1891. Caratillo (Guatemala, fide
Aguilar).
Dry to wet, brushy slopes, thickets, mixed forest or oak forest,
1,000-2,400 m.; Chimaltenango; Chiquimula; Guatemala; Quezal-
tenango; Sacatepequez; Santa Rosa. Southern Mexico; Honduras
and El Salvador to Panama; tropical South America.
Shrubs or suffrutescent herbs, 1-3 m. tall, sparsely or abundantly branched, the
branches often elongated and arching, very densely pubescent with short, ascending,
whitish or yellowish hairs; leaves on very short petioles, the blades commonly very
rugose and somewhat bullate, mostly ovate or lanceolate to elliptic, 5-8 cm. long,
acute or acuminate, rounded or very obtuse at the base, more or less scabrous above,
glandular-punctate beneath and usually very densely pilose with appressed or lax
hairs; inflorescences often large, lax, and broad, much branched, usually 10-30 cm.
broad; heads about 21-flowered, remote and sessile on the scorpioid branches;
involucres campanulate, 4-5 mm. high; phyllaries rather loosely imbricate, arachnoid-
tomentose, the middle and outer ones triangular-subulate, with spinose tips, the inner
ones lanceolate, long-acuminate; corollas white, pinkish or purplish white; achenes
about 2 mm. long, hispidulous; pappus white, the inner bristles about 4 mm. long.
NASH: FLORA OF GUATEMALA 23
Vernonia canescens var. pilata Blake, Brittonia 2: 331. 1937.
Damp or wet, brushy slopes, thickets, or on open banks in wet
forest, 1,000-2,500 m.; Quezaltenango (type from Colomba, Skutch
1993); San Marcos; Zacapa. Mexico (Chiapas); Costa Rica.
Slender shrubs, commonly 1-2.5 m. tall with weak, often recurved branches, the
stems sometimes 5 m. long and reclining on other shrubs or even subscandent,
puberulent or hispidulous with short, ascending hairs; leaves short-petiolate, the
blades more or less rugose, lanceolate or linear-lanceolate, mostly 6-16 cm. long and
1-2.5 cm. wide, narrowly long-acuminate, acute or subobtuse at the base, thinly
tuberculate-hispidulous or glabrate above and slightly scabrous, the veins often
impressed, densely glandular-punctate beneath and minutely hispid or sometimes
pilose on the veins; cymes scorpioid, very slender, usually elongated, more or less
recurved, often much branched and forming large, broad, open panicles; heads sessile,
only the lower ones leafy-bracteate; involucres about 4 mm. high, campanulate;
phyllaries sparsely ciliate, glabrous or nearly so, appressed, pale, tinged with purple,
lance- linear, the outer and middle ones attenuate into a short, subulate, spinose tip;
corollas pale pink; achenes brownish, 1.5-2 mm. long, appressed-hispidulous; pappus
white, the inner bristles 3-3.5 mm. long.
Vernonia cinerea (L.) Lessing, Linnaea 4: 291. 1829. Conyza
cinerea L. Sp. PL 2: 862. 1753.
Native of Old World; now a pantropical weed, well established
in some parts of the American tropics. Occasional in pine groves,
grassy slopes, gardens; Izabal. British Honduras.
Erect or sometimes procumbent annuals, the stems simple or branching, terete,
striate, to 1.5 m. tall, more or less pilose; leaves alternate, short-petiolate, the blades
mostly 2-4 cm. long, 0.5-3 cm. wide, obovate, broadly ovate, or oblong-spathulate,
obtuse or subacute, rounded and usually abruptly contracted and attenuate to the
base, the margins sinuate to crenate or coarsely dentate, sparsely pilose or pubescent
above or glabrate, more or less pilose beneath; inflorescence divaricate, branching
dichotomously, lax; heads pedicellate, about 20-flowered; involucres more or less
campanulate, about 3 mm. high; phyllaries 3-seriate, linear-lanceolate, acuminate to
mucronate, pubescent, dark-tipped; corollas purplish; achenes oblong, pilose; pappus
white, the inner bristles about 3 mm. long.
Vernonia corae Standl. & Steyerm. Field Mus. Bot. 22: 395.
1940.
Damp or wet thickets or mixed forest, 1,200-2,700 m.; San
Marcos (type from Volcan de Tajumulco, Steyermark 36787).
Shrubs or small trees to about 5 m. tall, the young branches densely tomentose;
leaves short-petiolate, the blades thin, oblong-lanceolate to narrowly obovate, mostly
10-25 cm. long, 2.5-8 cm. wide, acuminate, long-attenuate to the base, the margins
more or less serrate, sparsely appressed-pilose above or glabrate, glandular-punctate
beneath and more or less hispid-pilosulous, especially on the costae and veins; heads
1-4 on stout peduncles 3-8 cm. long, many-flowered; involucres broadly campanulate,
1.5-2 cm. high, 2-4 cm. broad; phyllaries multiseriate, greenish, densely appressed-
24 FIELDIANA: BOTANY, VOLUME 24
scaberulous, imbricate, the outer ones often squarrose, broadly ovate to triangular,
mucronate, the margins scarious, ciliate, commonly fimbriate and denticulate, the
innermost ones linear to linear-oblong, acuminate or acute, usually fimbriate or
pectinate; corollas purple or lilac, 15-17 mm. long; achenes dark, glabrous, about 3
mm. long; inner pappus 2.5-4 mm. long.
Vernonia ctenophora Gleason, Bull. Torr. Bot. Club 46: 243.
1919.
At or little above sea level, British Honduras (Freshwater
Creek Reserve), Campeche; type from Apazote; collected also at
Tuxpena.
Herbs, probably annual, to 1.5 m. tall, the slender branches puberulent and
minutely viscid; leaves sessile or short-petiolate, the blades thin, ovate-lanceolate or
the upper ones lanceolate, mostly 4-6 cm. long (lower leaves not seen, probably much
larger), acuminate, rounded or obtuse at the base, the margins entire or nearly so,
thinly and minutely puberulent above and densely glandular-punctate, closely
grayish-pilose beneath (rarely glabrate) and glandular-punctate; inflorescence of few,
greatly elongated, slender, scorpioid cymes, the bracts similar to the leaves but
smaller; heads remote, secund, sessile, 18-21-flowered; involucres campanulate, 6-7
(-8) mm. high; phyllaries closely imbricate, purplish, linear-lanceolate, thinly
pubescent and glandular-punctate, with subulate, spinose tips; achenes strigose,
about 1.5 mm. long; pappus white, the inner bristles about 4 mm. long.
Vernonia deppeana Less. Linnaea 6: 398. 1831. Suquinay
(Alta Verapaz; Chiquimula; Sacatepequez; Suchitepequez); suqui-
nay hembra (Alta Verapaz); semem (Alta Verapaz, Quecchi).
Wet to dry thickets and forest, sometimes in pine or oak forest,
300-2,000 m.; Alta Verapaz; Baja Verapaz; Chimaltenango; Chi-
quimula; Huehuetenango; Guatemala; Jalapa; Quezaltenango;
Retalhuleu; Sacatepequez; San Marcos; Santa Rosa; Solola;
Suchitepequez. Mexico; El Salvador and Honduras to Costa Rica.
Shrubs or small trees, sometimes to 9 m. tall, with a broad, dense, rounded
crown, the branches stout, densely pilosulous; leaves short-petiolate, the blades thick,
oblong or broadly elliptic, mostly 8-15 cm. long and 2.5-7 cm. wide, obtuse or acute,
acute to rounded at the base, the margins entire or obsoletely serrulate, papillose-
scabrous above, pale beneath and densely tomentose or floccose-tomentose;
inflorescences large, much branched, 10-30 cm. broad, the cymes scorpioid; heads
sessile, very numerous, 18-21-flowered; involucres campanulate, 3-4 mm. high;
phyllaries closely imbricate, pubescent, the outer ones ovate, acute or acuminate, the
inner oblong, acute or subacute, arachnoid-ciliate; corollas white, pinkish, or pale
lavender; achenes acutely costate, about 2.5 mm. long, thinly or densely pubescent;
pappus fulvous, the inner bristles about 4 mm. long.
Gleason, in North American Flora, used for this species the
older name of V. stellaris La Llave and Lex., but the application of
that name is uncertain, and it seems safer to adopt the later V.
NASH: FLORA OF GUATEMALA 25
deppeana, as its application to this species is not in doubt (see
Blake, Contr. U. S. Nat. Herb. 26: 228. 1930).
Vernonia heydeana Coulter, Bot. Gaz. 20: 42. 1895. Eremosis
heydeana Gleason, Bull. N. Y. Bot. Card. 4: 234. 1906.
In Guatemala, known only from the type, San Miguel
Uspantan, El Quiche, 1,800 m., Heyde & Lux 3392. Southern
Mexico.
Shrubs, the branches puberulent or glabrate; leaves on petioles 1-2 cm. long, the
blades elliptic, ovate, or ovate-lanceolate, mostly 7-10 cm. long, 3-6 cm. wide, acute or
abruptly short-acuminate, entire or remotely denticulate, acute at the base, thinly
papillose-puberulent or glabrous above, thinly tomentulose and glandular-punctate
beneath; inflorescences hemispheric or pyramidal panicles; heads crowded on short
pedicels, mostly 5-6-flowered, rarely with only 3 flowers; involucres campanulate,
about 6 mm. high; phyllaries 3-4-seriate, the outer ones short, ovate, obtuse, minutely
tomentose, ciliate, the inner ones elongated, brown with green centers, obtuse or
subacute, glabrous or nearly so, the margins scarious; corollas more or less glandular
near the apex; achenes brown, about 3 mm. long, glabrous or very minutely
glandular; pappus white, the inner bristles 5-6 mm. long, the outer bristles about 1
mm. long.
Vernonia leiocarpa DC. in DC. Prodr. 5: 34. 1836. Eremosis
leiocarpa Gleason, Bull. N. Y. Bot. Gard. 4: 232. 1906. E.
melanocarpa Gleason, I.e. (type from Santa Rosa, Heyde & Lux
3416). V. melanocarpa Blake, Contr. Gray Herb. 52: 18. 1917. Qan
ta'ax, supup (Alta Verapaz, Quecchi); suquinay (Guatemala).
Wet to dry, often rocky slopes, thickets or forest, frequently in
pine-oak forest, 1,000-2,500 m.; Alta Verapaz; Baja Verapaz;
Chimaltenango; Chiquimula; Guatemala; Huehuetenango; Jalapa;
Jutiapa; Quezaltenango; Sacatepequez; San Marcos; Santa Rosa;
Solola; Totonicapan; Zacapa. Southern Mexico; British Honduras
and Honduras to Nicaragua.
Shrubs or trees, rarely to 12 m. tall, with dense, usually rounded crowns, the
young branches densely tomentose; leaves on stout petioles 1-3 cm. long, the blades
ovate to lanceolate, mostly 7-14 (-20) cm. long and 2-6 cm. wide, acute or acuminate,
rarely obtuse, broadly cuneate or rounded at the base, the margins entire or dentate,
puberulent or glabrate above, white or grayish beneath with very dense tomentum;
inflorescences large and broad, leafy, 15-20 cm. long; heads very numerous, aggregate,
sessile or short-pedicellate, mostly 3-flowered, sometimes 4-7-flowered; involucres
cylindric or narrowly campanulate, 4-5 mm. high; phyllaries commonly purplish at
the apex, the outer ones broadly triangular-ovate, obtuse, densely tomentose, the
inner ones oblong; corollas pinkish to lavender; achenes pale brown to almost black,
glabrous, 2.5-3 mm. long; pappus white, the inner bristles 5-7 mm. long, the outer
ones 1-2 mm. long.
26 FIELDIANA: BOTANY, VOLUME 24
Called "acerillo" and "hoja blanca" in Honduras, "palo de
asma" in El Salvador, this species is known commonly in
Guatemala as "suquinay." There is a village called El Suquinay in
the Department of Guatemala.
This species is one of the most abundant shrubs on the open,
cleared hills of the Pacific coastal plain, where it forms extensive,
almost pure stands over wide areas. It hybridizes with V. standleyi
Blake, to form what has been called V. X calderonii Blake. In El
Salvador the plant is said to be a home remedy for asthma.
Vernonia medialis Standl. & Steyerm. Field Mus. Bot. 23:
148. 1944.
Damp or dry thickets, plains and slopes, 125-300 m.; Chimal-
tenango; Escuintla; Retalhuleu (type from Rio Coyote, 4 km. west
of Retalhuleu, Standley 87473).
Herbaceous or suffrutescent plants 1-1.5 m. tall, laxly branched, the branches
multistriate, densely and minutely puberulent; leaves on short, stout petioles to 0.5
cm. long, the blades membranaceous, lance-oblong or elliptic-oblong, sometimes
oblanceolate-oblong, mostly 7-15 cm. long, 2-6.5 cm. wide, acuminate to long-
acuminate, the base acute to almost rounded, the margins entire or nearly so,
sparsely and shortly tuberculate-pilosulous on the upper surface, in age scaberulous
or glabrate, densely glandular-punctate beneath, minutely hispidulous or pilosulous,
especially on the veins, in age almost glabrous; inflorescence composed of few or
numerous scorpioid cymes, elongated, simple or laxly branched and sometimes
forming large panicles, foliaceous-bracteate near the base; heads usually remote, the
upper ones usually naked (if bracteate, the bracts small, linear to oblong, often
inconspicuous), all sessile; involucres campanulate, 6-8 mm. high; phyllaries
numerous, laxly arachnoid-tomentose, the outermost short, somewhat spreading,
spinose-subulate, the costa thick, excurrent, the middle ones subappressed, linear-
lanceolate, attenuate and spinose-subulate, the innermost phyllaries broadly linear,
pale-marginate, subacute to acuminate; flowers 15-27 per head; corollas white or
pinkish, glabrous; achenes pale, 2-3 mm. long, densely antrorse-hispidulous; pappus
white, the inner bristles about 6 mm. long.
Vernonia mima Standl. & Steyerm. Field Mus. Bot. 23: 264.
1947.
Damp or rather dry thickets, 1,500-2,000 m.; Huehuetenango
(type collected near crossing of Rio San Juan Ixtan, east of San
Rafael Petzal, Standley 82871); Quezaltenango.
Erect shrubs 2-3 m. tall, the branches brownish, when young densely
subtomentose or pilose with sordid hairs; leaves on petioles 1.5-3 cm. long, the blades
broadly ovate or elliptic-ovate, mostly 7-12.5 cm. long, 5-8 cm. wide, acute, rounded
or obtuse at the base, sometimes abruptly contracted and short-decurrent on the
petiole, the margins subentire, sparsely puberulent or glabrate above, scarcely paler
beneath, minutely puncticulate, loosely subtomentose or short-pilosulous, the lateral
NASH: FLORA OF GUATEMALA 27
veins 6-8 pairs, prominent beneath; heads turbinate-campanulate, sessile or subsessile
in dense cymules, very numerous, the cymules forming a large, ovoid or rounded,
leafy panicle 20-30 cm. long, the branches densely tomentulose or sordid-pilosulous;
involucres about 8 mm. high; phyllaries 4-5-seriate, pale and greenish, commonly
glabrous or glabrate, sometimes tomentulose, closely appressed, the outer ones
triangular-ovate, acute, the innermost oblong- linear, obtuse, about equalling the
pappus; flowers about 10 in each head; immature achenes turbinate, brown, glabrous,
1-2 mm. long; pappus white, the inner bristles 4-5 mm. long.
Vernonia mollis HBK. Nov. Gen. & Sp. 4: 36. 1820. Chicuta
(Jutiapa). Semem, sukinay (Quecchi, Alta Verapaz).
Dry, brushy, often rocky plains and hillsides, 600-1,500 m.; Alta
Verapaz; Chimaltenango; Chiquimula; Jalapa; Jutiapa. Honduras;
tropical South America (fide Standley).
Erect, stiff shrubs or sometimes suffrutescent herbs, to 1.5 m. tall, the stems
densely tomentose; leaves sessile or nearly so, the blades thick and usually rugose,
ovate to oblong-ovate or rarely oblong, mostly 4-8 cm. long and 1.5-3 cm. wide, acute
or acuminate, obtuse or rounded at the base, the margins entire or rarely more or less
denticulate, usually densely hirsute-sericeous on the upper surface, densely
velutinous-tomentose beneath, the long, interlaced hairs concealing the numerous
glands; cymes scorpioid, few or numerous, short or enlongated, usually recurved,
sometimes forming rather large panicles; heads secund, sessile, about 21-flowered,
only the lower heads leafy-bracteate; involucres broadly campanulate, 4-5 mm. high;
phyllaries numerous, closely imbricate, triangular-linear, villous, with subulate,
spinose tips, the costa indurate and conspicuous, the outer phyllaries somewhat
spreading, the middle and inner ones appressed; corollas pink or white; achenes
hispidulous; pappus white, the inner bristles about 4.5 mm. long, the outer scales
broad, less than 1 mm. long.
Perhaps only a variety of V. canescens HBK., distinguished
from it by the abundant tomentum, but the material referred here
is uniform and readily separated from the typical form. Moreover,
in Guatemala, V. mollis is limited in distribution, having been found
so far only in the dry, eastern departments.
Vernonia oaxacana Sch. Bip. ex Klatt, Leopoldina 20: 74.
1884.
Not reported from Guatemala, but may be expected there as it
has been collected in Chiapas, as well as in Oaxaca, Mexico, in open,
sunny locations around 900 m.
Shrubs 1-3 m. tall, the stems terete, multistriate, densely tomentose when young,
much less so in age, sometimes short floccose-tomentose; leaves short-petiolate, the
blades elliptic-oblong to ovate-oblong, mostly 4-12 cm. long, 2-6 cm. wide, acute,
short-acuminate or sometimes obtuse, broadly cuneate or rounded at the base, often
scabrous above, tomentose or short floccose-tomentose beneath, the margins entire or
28 FIELDIANA: BOTANY, VOLUME 24
serrate; inflorescence a large, more or less corymbiform panicle, the primary
peduncles 1.5-3 cm. long, the secondary ones 0.5-1.5 cm. long, with usually only 2-3
heads sessile or short -pedicellate on a common peduncle; heads commonly with 12-14
flowers each; involucres campanulate, 4-5 mm. high; phyllaries ovate-lanceolate to
narrowly oblong, acute, sometimes minutely cuspidate, glabrous or somewhat
floccose-tomentulose, but always appearing greenish with prominent purple mark-
ings; corollas pinkish to purple or rarely white; achenes brown, glandular- pubescent,
about 3 mm. long; pappus white, the inner bristles about 6 mm. long
Vernonia patens HBK. Nov. Gen. & Sp. 4: 41. 1820. V.
lanceolaris A. DC. in DC. Prodr. 5: 37. 1836. V. aschenborniana
Schauer in Nees & Schauer, Linnaea 19: 714. 1847. V. salamana
Gleason, Bull. Torr. Bot. Club 46: 242. 1919. Suquinay.
Wet or dry thickets or forest, sea level to 1,700 m.; Alta
Verapaz; Chiquimula; Escuintla; Guatemala; Izabal; Jalapa; Peten;
El Progreso; Retalhuleu; Sacatepequez; San Marcos; Santa Rosa;
Suchitepequez. Southern Mexico; British Honduras; Honduras and
El Salvador to Panama; tropical South America.
Erect shrubs of 2-3 m. or sometimes small trees to 6 m. tall, the branches
tomentulose or glabrate; leaves short-petiolate, the blades narrowly oblong to
lanceolate or lance-oblong, mostly 6-15 cm. long, 1.5-3 cm. wide, acute or acuminate,
obtuse or acute at the base, the margins entire or sometimes denticulate, nearly
glabrous above or sparsely puberulent, more or less puberulent beneath, especially on
costae and veins, or somewhat scabrous; inflorescence usually much branched,
commonly 20-30 cm. broad, the cymes scorpioid; heads sessile, remote, ebracteate;
involucres broadly campanulate, 3-5 mm. high; phyllaries closely imbricate, glabrous
or puberulent, ciliate, pale green with brown centers or tips, acute or subacute,
sometimes obtuse and inconspicuously mucronulate; florets 11-21-per head; corollas
pinkish or white; achenes costate, hispidulous, about 2 mm. long; pappus fulvous, the
inner bristles about 5 mm. long.
This is probably the species reported from Baja Verapaz by
Loesener as V. bangii Rusby, a South American species.
Vernonia polypleura Blake, Journ. Wash. Acad. Sci. 28: 478.
1938.
Wet, mountain forest, about 1,800 m., San Marcos. Mexico
(Chiapas).
Shrubs or trees, sometimes as much as 19 m. tall, the branches sulcate-angled,
densely pilosulous; leaves on pubescent petioles mostly 1-3 cm. long, the blades
elliptic-oblong to lance-oblong, 10-18 cm. long, 2.5-5 cm. wide, acuminate, attenuate
to the base, the margins entire or nearly so, somewhat shiny above and essentially
glabrous or glabrate, except along the costae, densely pilose beneath; inflorescences
branching, usually large, the scorpioid cymes sometimes as much as 30 cm. long;
heads sessile, 17-25-flowered, often subtended by much reduced, bractlike leaves;
involucres campanulate; phyllaries multiseriate, closely imbricate, the outer ones
NASH: FLORA OF GUATEMALA 29
short, lanceolate, sharply acute or acuminate, the middle ones oblong-ovate, the
innermost ones commonly 5-7 mm. long, linear-oblong, often somewhat constricted
below the subtriangular, acute apex; corollas pink or purplish; achenes about 2 mm.
long, 10-costate, densely pilose; pappus fulvous, the inner bristles 4-5 mm. long.
Vernonia salvinae Hemsl. Biol. Centr. Amer. Bot. 2: 73. 1881.
Cacalia salvinae Kuntze, Rev. Gen. 970. 1891. Leiboldia salvinae
Gleason, Bull. N. Y. Bot. Card. 4: 162. 1906. Araha (Quezal-
tenango).
Damp or wet thickets or dense, mixed forest, 1,200-2,700 m.;
Quezaltenango (type from Volcan de Zunil, Salvin s.n.); El Quiche;
San Marcos. Southern Mexico.
Sparsely branched shrubs or small trees, 1.5-6 m. tall, the young branches
densely tomentose, sometimes glabrate in age; leaves short-petiolate, the blades thin,
oblong-oblanceolate to narrowly obovate, mostly 10-25 cm. long, 2.5-8 cm. wide,
acuminate, long-attenuate to the base, the margins commonly serrate, sometimes
entire, essentially glabrous above or sparsely appressed-pilose, glandular-punctate
beneath and thinly appressed-pilose, especially on the veins; heads 1-4 on stout
peduncles 2-8 cm. long, at the ends of the branches, the heads many-flowered;
involucres broadly campanulate or hemispheric, about 2 cm. high and 3-4 cm. broad;
phyllaries multiseriate, imbricate, more or less puberulent or almost glabrous, broadly
ovate to broadly triangular, mucronate, the margins scarious, almost entire or
obscurely denticulate, the outer ones commonly more or less squarrose, bright purple
or maroon; corollas pale purple, mostly 20-27 mm. long; achenes black, glabrous,
about 3 mm. long; inner pappus bristles 5-8 (-10) mm. long.
Vernonia scorpioides (Lam.) Pers. Syn. PI. 2: 404. 1807.
Conyza scorpioides Lam. Encycl. Meth. 2: 88. 1786.
Open places, at or near sea level; British Honduras (Stann
Creek). Yucatan; Honduras; Nicaragua; tropical South America.
Erect, sparsely branched, herbaceous or suffrutescent plants, the stems densely
short-pilose with ascending hairs; leaves short-petiolate, the blades ovate to
lanceolate, mostly 5-12 cm. long, 2-7 cm. wide, acute or acuminate, abruptly cuneate
at the base and decurrent on the petiole, the margins entire, softly papillose-
pilosulous or glabrate above, thinly pilose to sericeous or almost tomentose beneath,
not glandular; cymes few, spreading, scorpioid, crowded at the ends of the branches,
short, forming a depressed panicle; heads crowded, sessile, ebracteate, about 21-
flowered; involucres campanulate, about 4 mm. high; phyllaries few-seriate, pale,
loosely imbricate, lanceolate, acuminate, villous; corollas lavender or pinkish;
achenes pubescent; pappus white, the inner bristles about 5 mm. long.
Vernonia shannonii Coulter, Bot. Gaz. 20: 42. 1895. Eremosis
shannoni Gleason, Bull. N. Y. Bot. Gard. 4: 234. 1906.
Damp, mixed forest or in oak or pine-oak forest, 2,500-3,800 m.;
30 FIELDIANA: BOTANY, VOLUME 24
Huehuetenango; Quezaltenango; San Marcos (type from San
Lorenzo, Shannon 605); Totonicapan.
Shrubs or trees, sometimes 10 m. tall, the branches obscurely tomentulose or
almost glabrous; leaves on petioles 0.5-2 cm. long, the blades thin, ovate-lanceolate,
mostly 10-17 cm. long, 4-7 cm. wide, acuminate to obtuse, acute or rather abruptly
attenuate to the base and often somewhat decurrent on the petiole, the margins
entire or nearly so, when young floccose-tomentose, but in age essentially glabrous,
except along the costae; inflorescences broad, much branched, dense; heads on
pedicels sometimes more than 1 cm. long, usually 2-5 on a common, slender peduncle,
each head 5-6-flowered; involucres narrowly campanulate, 8-11 mm. high, dark
yellowish to light brown; outer phyllaries broadly ovate, obtuse or subacute,
glabrous, often somewhat arachnoid-ciliate, the inner ones elongated, linear-oblong,
deciduous, rounded at the apex, ciliate, glabrous or tomentulose at or near the apex;
corollas lilac; achenes about 5 mm. long, glabrous, 10-costate, more or less glandular;
pappus white, the inner bristles 7-8 mm. long.
Vernonia standleyi Blake, Journ. Wash. Acad. Sci. 13: 143.
1923. Figure 11.
Dry, brushy, often rocky plains or hillsides, sometimes in pine-
oak forest, 850-1,400 m.; Chimaltenango; Jalapa; Jutiapa. El
Salvador; Honduras; Nicaragua.
Erect shrubs, 1-3 m. tall, rather densely branched, the branches brown, glandular
and puberulent or in age glabrate; leaves subsessile to short-petiolate, the blades
lance-oblong to elliptic, mostly 6-9 cm. long and 1.5-3 cm. wide, acuminate, cuneate
at the base, the margins entire or serrulate, glandular-punctate above and
puberulent, in age glabrate, somewhat paler beneath, glandular-punctate, short-pilose
or finally glabrate; heads (4-) 5- flowered, very numerous, sessile or subsessile and
forming dense globose glomerules 1-3 cm. thick; involucres about 4.5 mm. high;
phyllaries 5-6-seriate, pale, the outermost ones ovate, the middle ones ovate-oblong,
the innermost linear and deciduous, obtuse or subacute, glandular and somewhat
villous, arachnoid-ciliate; corollas white, glandular, 4-6 mm. long; achenes about 2.5
mm. long, pale, densely antrorse-pilosulous; pappus white, the inner bristles about 4
mm. long.
Vernonia tenella D. Nash, Fieldiana: Botany 36 (9): 74. 1974.
Wet mountain forest, 1,800-2,400 m., San Marcos (type from
slopes of Tajumulco Volcano, Sierra Madre Mountains, 2,300 m., L.
O. Williams et al. 26876). Also collected near Aldea Fraternidad,
between San Rafael Pie de la Cuesta and Palo Gordo, Williams,
Molina, & Williams 26094 and 26264.
Erect shrubs to 3 m. tall, or sometimes sprawling or somewhat scandent, the
stems terete, striate, puberulent or thinly appressed-pubescent to glabrate; leaves on
petioles 0.5-1.5 cm. long, the blades narrowly lanceolate to lance-oblong or elliptic, 5-
11 cm. long, 1.5-3 cm. broad, acuminate, attenuate to the base and decurrent on the
petiole, essentially glabrous above or puberulent along the costae, glabrous beneath
or with some pubescence usually confined to the costae and veins, the margins entire;
NASH: FLORA OF GUATEMALA 31
inflorescences composed of branching scorpioid cymes, the branches very slender,
mostly 5-10 cm. long, about 0.5 mm. in diameter, minutely tomentose; heads sessile,
secund, remote, the flowers about 15 to each head; involucres 6-8 mm. high, more or
less tomentose; outer and middle phyllaries acute, the inner longer ones linear-
oblong, obtuse to subacute or subtriangular at the apex; corollas lavender or purple;
achenes 1.5-2 mm. long, pilose; pappus pale fulvous, the inner bristles about 6 mm.
long.
Vernonia tortuosa (L.) Blake, Proc. Biol. Soc. Wash. 39: 144.
1926. Conyza tortuosa L. Sp. PI. 862. 1753. V. schiedeana Less.
Linnaea 6: 399. 1831. Rash k^am (Quecchi; Alta Verapaz).
Wet to dry thickets or open forest, often in pine or pine-oak
forest, 100-1,500 m.; Alta Verapaz; Chiquimula; Escuintla; Guate-
mala; Izabal; El Progreso; Peten; Quezaltenango; Retalhuleu;
Sacatepequez; San Marcos; Santa Rosa; Suchitepequez. Southern
Mexico; British Honduras; Honduras; El Salvador; Costa Rica.
Stout shrubs, commonly 1-3 m. tall, the branches often elongating and recurving
or reclining on other plants, sordid-pubescent; leaves on short, stout petioles mostly
0.3-1.5 cm. long, the blades usually thick and firm, mostly 8-15 cm. long and 2-5 (-8)
cm. wide, commonly lustrous, elliptic-oblong to oblong-lanceolate or oblanceolate-
oblong, abruptly short-acuminate, obtuse, or rounded and apiculate, rounded or
cuneate at the base, the margins entire or nearly so, the upper surface sometimes
bullate, more or less papillose-scabrous, papillose-strigose or puberulent beneath,
sometimes tomentose, the venation prominent (commonly 7-10 pairs of lateral veins);
inflorescences usually large and freely branched, the scorpioid cymes elongated, often
recurved; heads sessile, (18-) 30-40-flowered, the lowermost heads leafy-bracteate;
involucres broadly campanulate, 9-10 mm. high and about as broad; outer and middle
phyllaries lanceolate, acute or acuminate, glabrate, with a thick, elevated costa,
cuspidate, the inner phyllaries oblong, ciliate, broadly rounded or emarginate at the
apex, the apices incurved in bud, recurved in anthesis; corollas white or pinkish;
achenes hispidulous, 1.5-2.5 mm. long; pappus pale yellowish or almost pure white,
the inner bristles 7-8 mm. long.
A decoction of these plants is a common household remedy in
Alta Verapaz, used mostly to treat disturbances of the digestive
system.
Vernonia triflosculosa HBK. Nov. Gen. & Sp. 4: 40. 1820.
Cacalia triflosculosa Kuntze, Rev. Gen. 2: 971. 1891. V. luxensis
Coult. Bot. Gaz. 20: 41. 1895 (type from Santa Rosa, Heyde & Lux
3421). V. dumeta Klatt, Bot. Beibl. Leopoldina 1895, 1. 1895.
Eremosis triflosculosa Gleason, Bull. N. Y. Bot. Gard. 4: 233. 1906.
V. chaclana Blake, Contr. Gray Herb. 52: 19. 1917. Pie de paloma
(San Marcos).
Wet to dry thickets on plains or rocky hillsides, 120-1,250 m.;
Escuintla; Guatemala; Jalapa; Quezaltenango; Retalhuleu; Sacate-
32 FIELDIANA: BOTANY, VOLUME 24
pequez; San Marcos; Santa Rosa; Suchitepequez. Southern Mexico;
Honduras and El Salvador to Panama.
Shrubs or small trees, sometimes to 8 m. tall, with dense, rounded crowns, the
branches thinly pubescent or glabrate; leaves short-petiolate, the blades thin,
oblanceolate to elliptic, mostly 6-13 cm. long and 1.5-3 cm. wide, acute to long-
acuminate, attenuate to the base and more or less decurrent on the petiole, the
margins entire to coarsely serrate or dentate, almost glabrous on both surfaces,
glandular-punctate beneath; inflorescences large, conic or hemispheric, commonly
dense and leafy; heads commonly 3-flowered, sessile or very short-pedicellate, usually
2-5 on a common peduncle; involucres cylindric, pale brown or greenish, 4-5 mm.
high; phyllaries laxly imbricate, the outer ones rounded or ovate, subacute, the inner
ones oblong, acute or subacuminate, glabrous or nearly so, ciliate; achenes about 3
mm. long, pubescent; pappus white, the inner bristles 5-6 mm. long.
TRIBE II. EUPATORIEAE
By Louis O. WILLIAMS
Reference: B. L. Robinson, A generic key to the Compositae-
Eupatorieae, Proc. Amer. Acad. 49: 429-437. 1913.
Herbs, shrubs or weak tree; leaves opposite or rarely the uppermost alternate;
inflorescences mostly corymbose but sometimes otherwise or even reduced to a single
head; involucres may have a somewhat definite number of phyllaries (usually 4 in
Mikania and 5-6 in Stevia) or an indefinite number and often very many (Ageratum,
Eupatorium, etc.), these appear to be in 1, 2, 3 or sometimes more series, usually
reduced in length and broader outward, often accompanied by reduced bracts or
scales at the base; phyllaries often or usually stiff and bi- or tricostulate with
thickened costulae (Eupatorium, Brickellia, Ageratum, etc.) or sometimes relatively
thin with more nerves and these not so prominent; receptacles sometimes conical but
more often nearly flat, without pales or rarely pales present; florets in a head may be
subdefinite in number (in ours), as few as three or four (Stevia, Mikania) or contain a
great number (Eupatorium, Brickellia); ray florets are unknown in the tribe; achenes
from 4-12-costate but usually with 5 or 10 costae or rarely apparently not costate,
small in size, often shining, glabrous or not; the pappus in the tribe exhibits most of
the forms found in the Compositae, in Guatemala the common form is smooth or
barbellate, capillary hairs or bristles, simply because this is the form found in
Eupatorium, which contains well more than half of the species in the tribe, pappus
scales joined at the base or not and bristle-tipped or not, clavate and sometimes
gland-tipped, a plumose capillary pappus or bristles occur as do awn-like or setiform
pappuses, or the pappus may be reduced to a disc, or crown, or ring, or appear to be
completely lacking, a pappus may be present or lacking in the same species (often in
Ageratum); corollas mostly small, often minute, but because of their abundance
often attractive, colors vary from white through blues and purples, while pale green is
rare and yellows are not known in ours, color is usually constant within a species,
anthers (4-)5, small or minute, unappendaged at the base, a prominent apical anther
appendage, usually thin, is almost always present but may be lacking (t*iqueria) or
sometimes difficult to distinguish (some Eupatorium); styles usually exserted and
relatively long in comparison to the corolla, usually clavellate.
WILLIAMS: FLORA OF GUATEMALA 33
Members of the tribe are to be found in most ecological
situations in Guatemala, but mostly are plants of the middle to
highest elevations, where a definite wet-dry season regimen occurs.
Pappus none or a very short and obscure crown.
Heads 3-5-f lowered; herbs, shrubs or subshrubs.
Heads 3-flowered; herbs; anthers not appendaged at the apex Piqueria.
Heads 4-5-flowered; shrubs or subshrubs; anthers obscurely appendaged at the
apex Eupatorium sodali.
Heads more than 20-flowered; herbs; anthers appendaged at the apex Isocarpha.
Pappus present, better developed.
Achenes obovoid or prismatic and 8-10-costate.
Achenes obovoid; plants herbaceous Adenostemma.
Achenes prismatic, 8-10-costate, shrubs or herbs Brickellia.
Achenes mostly 4-5-costate, usually prismatic.
Pappus, at least in part, of squamellae or awns.
Phyllaries only 5-8, uniseriate, subequal Stevia.
Phyllaries more numerous, usually imbricated.
Corollas with short tubes and scarcely distinct throats Ageratum.
Corollas with slender tubes, abruptly dilated into the throat Oxylobus.
Pappus wholly of soft bristles.
Phyllaries 4; heads 4-flowered; plants scandent Mikania.
Phyllaries usually more numerous; heads with usually numerous flowers;
plants rarely scandent.
Bristles of the pappus plumose; annuals Carminatia.
Bristles of the pappus not plumose; mostly perennials.
Achenes strongly compressed; pappus bristles 2 Macvaughiella.
Achenes not compressed; pappus bristles usually numerous; receptacle
epaleaceous or rarely paleaceous Eupatorium.
ADENOSTEMMA Forster
Small or rather large herbs, glandular-pubescent or glabrate, perennial or
annual; leaves opposite, petiolate, usually triplinerved, dentate; heads small or
medium-sized, white, discoid; inflorescence corymbose; involucre broadly camp-
anulate or hemispheric; phyllaries herbaceous, in ours connate to the middle and 1-2-
seriate; receptacle almost flat, naked; corollas subequal, regular, the tube short, the
limb campanulate, 5-dentate; anthers subtruncate at the apex, not appendaged,
obtuse and subtruncate at the base, the connective bearing a small and sometimes
apiculate gland; achenes obtuse, 5-costate, glandular-tuberculate; pappus of a few
short, rigid, often clavate, gland-like bristles.
Species 10 or more, in the tropics of both hemispheres. Only
one is found in Central America.
Adenostemma hirtiflorum Benth. PI. Hartweg. 75. 1841.
Figure 12.
Wet mixed mountain forest, or in openings in forest, 1,200-1,500
m.; endemic; Quezaltenango (type from "Rancho Santa Rita,"
34 FIELDIANA: BOTANY, VOLUME 24
Hartweg 531; collected also at Palmar and Finca Pirineos).
Plants decumbent and rooting at the base, probably perennial, the stems erect,
rather stout, mostly 50 cm. high or less, simple or sparsely branched, glabrate; leaves
few, mostly 3-4 pairs, slender-petiolate, ovate or deltoid-ovate, thin, 6-11 cm. long,
acute or acuminate, rather abruptly cuneate or broadly cuneate at the base and
decurrent, coarsely and unevenly serrate-dentate, glabrous above or nearly so,
somewhat puberulent beneath, especially on the nerves; heads few, in open corymbs,
on long slender peduncles, the peduncles densely puberulent; involucre in anthesis 5
mm. long and broad, in fruit 8 mm. broad; phyllaries green, connate to the middle,
oblong or obovate-oblong, rounded at the apex, glabrate, densely and conspicuously
ciliate; corollas hirtellous; achenes obovoid, angulate, 3 mm. long, almost smooth;
pappus bristles usually 5.
The relationship of this species to those of South America and
the West Indies remains to be determined. As now understood, the
present species is severely limited in its distribution.
AGERATUM Linnaeus
References: B. L. Robinson, Revisions of Alomia, Ageratum,
and Oxylobus, Proc. Am. Acad. 49: 438-492. 1913. Miles F. Johnson,
A monograph of the genus Ageratum L., Ann. Mo. Bot. Gard. 58: 6-
88. 1971. R. M. King & H. Robinson, Additions to the genus,
Ageratum, Phytologia 24: 112-117. 1972. R. M. King & H. Robinson,
A new genus, Blakeanthus, I.e. 118-119.
Annual or perennial herbs, sometimes fruticose; leaves all or chiefly opposite,
mostly ovate or lanceolate, crenate, serrate, or rarely entire, sessile or petiolate, often
glandular-punctate; inflorescence generally terminal, corymbose or cymose, usually
compound; heads homogamous, discoid, the involucre usually campanulate;
phyllaries narrow, subequal, imbricate in 2-3 or rarely more series, generally lance-
linear, acute or attenuate, mostly 2-costate; receptacle flat, convex, or conic, naked
or paleaceous; corolla 5-lobate, commonly blue or purple, sometimes white or pink;
anthers oblong or linear, rounded at the base, with an apical, membranaceous, ovate
or oblong appendage; achenes prismatic, 5-angulate; pappus scales setiferous or
muticous, distinct or connate at the base, sometimes forming an entire or dentate,
crateriform corona, or none.
Species about 45, one A. conyzoides, pantropical as a weed.
The genus Ageratum in this flora contains all of the Ageratum-
like plants known in the region except Oxylobus. It includes
Ageratum, Alomia of our region, without saying whether or not the
genus should be maintained in a reduced sense elsewhere, and
Blakeanthus.
Pappus absent or represented by a low undivided crown on the apex of the achene.
Heads large, as broad as long; leaves attenuate to the base A. platylepis.
WILLIAMS: FLORA OF GUATEMALA 35
Heads smaller and longer than broad; leaves truncate or cordate at the base.
A. cordatum.
Pappus present, of membranaceous scales, these sometimes setiferous or coroniform
or cupuliform.
Pappus of 5-6 setiferous scales free at the base.
Plants erect; leaves eldiptic or linear.
Heads single; pappus scales blunt, lacerate or minute and apparently
coroniform A., gaumeri.
Heads several: pappus scales constricted above to entire or lacerate setae, or
truncate and lacerate to entire.
Plants usually pubescent; leaves elliptic; coastal E. ellipticum.
Plants usually glabrous; leaves linear; pine savannas E. peckii.
Plants erect or decumbent or repent; leaves lanceolate, oblong, ovate or deltoid.
Phyllaries triseriate, firm, glabrous; lacustrine A. radicans.
Phyllaries biseriate, pilose to stipitate-glandular; not lacustrine.
Phyllaries eglandular, linear-lanceolate; leaves ovate, obtuse at the base but
not cordate not truncate A. conyzoides.
Phyllaries stipitate-glandular, linear-lanceolate, acuminate; leaves ovate to
deltoid, truncate to cordate at the base A. houstonianum.
Pappus coroniform, or if setiferous the setae joined at the base.
Receptacles paleaceous A. isocarphoides.
Receptacles epaleaceous or the pales few and scattered.
Plants perennial, decumbent or repent and of the seashore; pappus coroniform
or entire A. littorale.
Plants shrubby or suffrutescent, not of the seashore.
Leaves entire, lanceolate to ovate, white tomentose below A. chortianum.
Leaves dentate or crenate, elliptic to deltoid.
Lower leaf surface tomentose; phyllaries conspicuously pilose.
A. rugosum.
Lower leaf surface pilose or scabrous; phyllaries rarely densely pilose.
A. corymbosum.
Ageratum chortianum Standl. & Steyerm. Field Mus. Bot.
23: 98. 1944.
Brushy places, sometimes along bluffs, 300-1,500 m.; Jalapa;
Chiquimula (type from Montana Castilla, southeast of Quezalte-
peque, Steyermark 31269); Guatemala. Honduras.
Plants suffrutescent, branched, 1 m. high, the stems terete, at first densely
grayish-velutinous with short antrorse hairs; leaves short -petiolate, thick, oblong-
lanceolate, 3.5-7 cm. long, 10-30 mm. broad, obtuse or acute, broadly cuneate at the
base, green and somewhat lustrous above, thinly puberulent, densley grayish-
tomentose beneath; heads about 10, corymbose, pale lilac, long-pedunculate, the
pedicels 4-5 mm. long; involucre campanulate, 4-5 mm. high, 5-7 mm. broad;
phyllaries about 3-seriate, graduate, narrowly lanceolate or linear-lanceolate,
attenuate, 2-3-costate, appressed-pilosulous; receptacle conic, obtuse; corollas 2.2-2.5
mm. long, sparsely pilose and gland-dotted; achenes brownish, glabrous, 2 mm. long,
5-angulate; pappus a slightly 5-lobate crown 0.2 mm. high.
36 FIELDIANA: BOTANY, VOLUME 24
Ageratum conyzoides L. Sp. PI. 839. 1753. Coelestina
microcarpa Benth. ex Oersted, Vidensk. Meddel. 1852: 72. 1852.
Ageratum microcarpum Hemsl. Biol. Cent. Am. Bot. 2: 82. 1881.
Alomia microcarpa Rob. Proc. Am. Acad. 49: 452. 1913. Mejorana;
mejorana chaparro; flor noble.
Moist or wet thickets, often in waste ground, a frequent weed
in gardens and grain fields, occasionally in wet meadows or in pine
forest, frequent on sandbars along streams, 2,000 m. or less, most
common at 900 m. or lower; Alta Verapaz; Izabal; Zacapa;
Chiquimula; Jalapa; Jutiapa; Santa Rosa; Escuintla; Solola;
Suchitepequez; Retalhuleu; Quezaltenango; San Marcos; Huehue-
tenango. Mexico; El Salvador and Honduras to Panama; West
Indies; South America; Old World tropics, introduced.
A rather coarse annual, erect or sometimes decumbent, branched, the stems
puberulent and villous; leaves thin, on long slender petioles, ovate to broadly deltoid-
ovate, 2-8 cm. long, obtuse, rounded to subcordate at the base, crenate-serrate, thinly
villous on both surfaces, sometimes glandular-punctate beneath; corymbs often
several at the ends of the branches, of few or numerous heads, short-pedunculate, the
pedicels slender, 3-7 mm. long, glandular or hispidulous; heads usually lavender or
pale blue, about 50-flowered, 6 mm. broad; involucre campanulate; phyllaries oblong,
green, mostly 2-costate, rather abruptly acuminate, scarious-margined, erose-
denticulate and long-ciliate near the apex, sparsely villous or glabrate dorsally;
receptacle naked; corollas glabrate or glandular- puberulent; achenes black, lustrous,
often hispidulous on the angles; pappus scales 5, lanceolate, fimbriolate, setiferous at
the apex, about equaling the corolla.
A. conyzoides is sometimes called "hierba de perro" and "hierba
de chucho" in El Salvador and is one of the common lowland weeds
in many regions of Central America. In the Coban region and in
some parts of eastern Guatemala this is one of the common weeds
of cornfields, often covering them with solid sheets of lavender or
pale blue and affording a beautiful show of color, just like that
displayed in Costa Rica by Santa Lucia, the epappose form of this
species, which usually has been called Alomia microcarpa. These
two forms, exactly alike except that the northern form bears a
pappus while the common southern form is without a pappus, come
together in Honduras and beautify the same old cornfields in the
beginning of the dry season. The epappose form of the species has
not been found in Guatemala but is to be expected.
A form with white flowers is occasionally found and is of no
taxonomic importance.
Ageratum cordatum (Blake) L. Wms. Fieldiana, Bot. 36: 78.
1975. Alomia cordata Blake, Proc. Biol. Soc. Wash. 60: 41. 1947.
WILLIAMS: FLORA OF GUATEMALA 37
Blakeanthus cordatus King & Robinson, Phytologia 24: 119. 1972.
Figure 13.
Dry oak or oak-pine woods, 650-1,200 m. or perhaps higher;
Zacapa (type, above Santa Rosalia, Steyermark 42712); Honduras.
Low, erect or spreading, suffrutescent herbs or shrubs, the stems terete, with
short, dense and crisped pubescence, internodes 2-6 cm. long; leaves petiolate, broadly
ovate-lanceolate, triplinerved from the base, acute or acuminate, subcordate or
cordate at the base, rarely somewhat acute, densely short crisped pubescent
interspersed with stipitate glandular pubescence below, pubescence thinner above, the
blades 3.5-9 cm. long and 2-7 cm. broad, the petioles mostly 1-2 cm. long;
inflorescence corymbose, terminal, pedunculate with peduncles about 6 cm. long, the
separate corymbs with 10-40 heads or perhaps more, dense, subglobose; heads 6-8
mm. high and 3-4 mm. in diameter, each with 15-35 flowers; involucre campanulate,
biseriate; phyllaries 5-6 mm. long and 0.8-1 mm. broad, linear-elliptic, acuminate,
obscurely 3-nerved, sparsely crisped puberulent, the inner phyllaries less pubescent;
corolla about 3 mm. long, tubular but slightly expanded toward the apex; styles
clavate, exserted about 2 mm. at anthesis; achenes about 2 mm. long, glabrous or
nearly so, with an obscure cartilagineous crown; pappus none.
Known from but two collections in Guatemala, but known
from many collections in central Honduras.
Ageratum corymbosum Zuccagni ex Pers. Syn. PL 2: 402.
1807. Coelestina ageratoides HBK. var. latifolia DC. Prodr. 5: 108.
1836. Ageratum corymbosum var. latifolium Rob. Proc. Am. Acad.
49: 476. 1913. A. elachycarpum Rob. I.e. 477 (type from Santa Rosa,
Heyde & Lux 4228). A. guatemalense M. F. Johnson, Ann. Mo. Bot.
Gard. 58: 64, fig. 1971 (type probably from Solola, Ownbey &
Muggli 3972). Mejorana; mota; monillos; polinegra (fide Aguilar).
Moist or dry, brushy or grassy, often rocky slopes, common in
pine and oak forest, rarely in denser forest, 2,500 m. or lower,
mostly at 1,000 m. or higher, except in the north coast where it
descends to sea level; Peten; Alta Verapaz; Baja Verapaz; Izabal;
Zacapa; Chiquimula; Jalapa; Jutiapa; Santa Rosa; Escuintla;
Guatemala; Sacatepequez; Chimaltenango; Solola; Suchitepequez;
El Quiche; Huehuetenango; Quezaltenango; San Marcos. Mexico.
Plants perennial, herbaceous or somewhat woody, commonly about 1 m. high but
sometimes as much as 2.5 m., the stems simple or branched, sparsely or densely
tomentose or grayish-pubescent; leaves opposite or the uppermost alternate, rather
thick and firm, ovate, rhombic-ovate or rhombic- lanceolate, mostly 4-10 cm. long,
acute or subacute, rounded or obtuse at the base, sometimes cuneate, coarsely
crenate or dentate, usually 3-nerved from the base, often reticulate- veined, scabrous
and puberulent above, green or grayish, densely crisp-puberulent or more often
tomentose with white to grayish hairs beneath, the petioles mostly 1 cm. long or less;
heads lavender, 7 mm. broad; inflorescence corymbose, the corymbs terminal on the
38 FIELDIANA: BOTANY, VOLUME 24
branches, simple or compound, many-headed, commonly dense, the pedicels stout, 3-
12 mm. long, glandular-tomentulose; involucre campanulate or hemispheric;
phyllaries lance-linear, attenuate, hispidulous, 2-costate, often colored at the apex;
receptacle naked; corollas sparsely glandular-atomiferous and often pubescent;
achenes about 2.2 mm. long, dark brown or blackish, glabrous; pappus crateriform,
0.3 mm. long, the margin entire or shallowly and irregularly dentate.
A white flowered form, A. corymbosum f. album Rob. (Proc.
Am. Acad. 49: 475. 1913) is represented in Guatemala but is of little
systematic importance. The flowers of Ageratum often fade to
white or near white when past maturity.
A form with phyllaries and achenes a bit shorter than the other
forms of the species, Ageratum corymbosum f. elachycarpum (Rob.)
M. F. Johnson (Ann. Mo. Bot. Gard. 58: 56. 1971) is found in
Mexico, Guatemala (from whence the type) and Honduras.
The type of Ageratum guatemalense is an attenuated and
somewhat immature specimen of A. corymbosum, probably from the
department of Solola rather than Totonicapan.
Ageratum ellipticum Rob. Contr. Gray Herb. 90: 5. 1930.
Chiefly in pine ridges, at or little above sea level; Peten
(Molina 15591). British Honduras (type from Honey Camp, Lundell
512; collected also at Tower Hill and All Pines).
Annual, stiffly erect, 75 cm. high or less, the stems slender, sparsely branched,
remotely leafy, inconspicuously setose-villous and scabrous-puberulent; leaves
opposite, on petioles 1 cm. long or shorter, elliptic-oblong to oblong-lanceolate, 2-5
cm. long, rather thick, obtuse or acute, cuneate at the base, inconspicuously
appressed-crenate to subentire, 3-nerved, almost glabrous but sparsely appressed-
pilose on the nerves; corymbs small, long-pedunculate, dense; heads few, about 40-
flowered, 7 mm. broad, 5 mm. high, lavender; phyllaries lanceolate, attenuate, green
obscurely ciliate, glabrous, 2-nerved, 4 mm. long; achenes black, 1.2 mm. long,
scaberulous on the angles; pappus scales 5, scarious, lacerate at the apex, attenuate
into a short bristle, about equaling the corolla.
Recently discovered near Poptun, Peten, previously known
only from British Honduras.
Ageratum gaumeri Rob. Proc. Am. Acad. 47: 191. 1911.
Known in Guatemala only from Peten. Mexico.
Small annual herbs to 0.5 m. tall, branched above, the stems sparsely pubescent
or glabrous; leaves ovate to deltoid, acute, the base obtuse, crenate, sparsely
pubescent, the upper surface sometimes glabrous, the blade 3-8 cm. long and 1.5-4 cm.
broad, the petiole 1-3 cm. long, pilose or glabrous; inflorescence long pedunculate;
heads campanulate, about 5 mm. long; phyllaries 2-seriate, lanceolate, acute, 3 mm.
long or less, bicostulate, glabrous or nearly so, erose or entire; achenes about 1.5 mm.
WILLIAMS: FLORA OF GUATEMALA 39
long, hispidulous on the angles; pappus of 5 white membranaceous scales; corolla
campanula te, 1.5-2.5 mm. long, lilac or lavender.
Ageratum houstonianum Mill. Gard. Diet. ed. 8. no. 2. 1768.
Mejorana; lokab or docaj (Coban, Quecchi).
Moist thickets or open places, sometimes a weed in waste or
cultivated ground, 1,200-1,500 m.; Alta Verapaz. Mexico; Nicaragua;
Costa Rica; West Indies; South America.
Annual, usually erect and often much branched, the stems leafy throughout,
pilose or villous with spreading hairs; leaves opposite, on rather long, slender petioles,
very thin, broadly deltoid-ovate, mostly 4-8 cm. long, obtuse or acute, generally
shallowing cordate at the base, coarsely crenate, thinly villous-hirsute, not punctate
beneath; corymbs terminal on the branches, often compound, glandular-hirtellous
and villous with several-celled hairs; heads numerous lavender or pale blue, 8 mm.
broad, about 75-flowered; phyllaries narrowly lanceolate or linear, entire, herbaceous,
long-attenuate, ciliolate, usually purplish toward the apex, 2-costate, glandular-
puberulent and hirsutulous; corollas 3 mm. long, the tube sparsely glandular-
puberulent; achenes black, lustrous, commonly hispidulous, at least on the angles, 1.2
mm. long; pappus scales 5, lanceolate, fimbriate-margined, long-setiferous, about
equaling the corolla.
In general appearance this species is exactly like A. conyzoides,
but, although separated from it by rather inconsequential charac-
ters, is fairly distinct.
This species has been found in Guatemala only in the vicinity
of Coban and San Cristobal Verapaz, Alta Verapaz, where it may be
an escape from cultivation. It is to be expected elsewhere in the
country.
Ageratum isocarphoides (DC.) L. Wms. Fieldiana, Bot. 36:
78. 1975. Coelestina isocarphoides DC. 5: 107. 1836. Alomia
isocarphoides Rob. Proc. Am. Acad. 49: 449. 1913.
Known only from the Department of Huehuetenango in the
area of mixed oak-pine forests, 900-1,600 m. Mexico.
Herbs to 1 m. tall, sparingly branched to fastigiately branched above, the stems
terete, nodose, densely pilose-puberulent to almost glabrous; leaves narrowly
lanceolate to ovate-lanceolate, acute or acuminate, triplinerved above the base,
densely tomentulose-puberulent and glandular dotted below, puberulent above, blade
to 14 cm. long and 5.5 cm. broad, reduced upward, petioles mostly less than 3 cm.
long; inflorescences long-pedunculate, corymbose clusters of heads, the pedicels short
and mostly less than 5 mm. long; heads paleaceous, about 5-7 mm. high,
campanulate, often with nearly 100 flowers; involucres 2-3 seriate; phyllaries linear-
lanceolate, acute, the apex indurated, puberulent dorsally and dotted with small,
purple glands, about 3-3.5 mm. long; corolla cylindric, somewhat expanded above,
about 2.5-3 mm. long, purple; achenes black, glabrous, about 1.5-2 mm. long; pappus
coroniform, 0.5 mm. high or less, undivided.
40 FIELDIANA: BOTANY, VOLUME 24
The description is from Guatemalan specimens. Guatemalan
specimens seen by Johnson were annoted as Ageratum echioides
(Less.) Hemsl, which they do not seem to be.
Ageratum littorale Gray, Proc. Am. Acad. 16: 78. 1880. A.
littorale var. hondurense Rob. Proc. Am. Acad. 49: 468. 1913
(described from Roatan Island, Honduras, and Mujeres Island,
Yucatan). A. littorale f. setigerum Rob. I.e. (type from Mujeres
Island).
British Honduras, on sandy sea beaches and limestone rocks;
Turneffe Island, F. E. Egler 42-8; Florida Keys; Mexico.
Plants decumbent or suberect, branched from the base, 30-40 cm. high, almost
glabrous, probably perennial, the stems pale, simple or branched, very leafy below,
terete; leaves opposite, on slender petioles 1-2.5 cm. long, rhombic, mostly 2-3.5 cm.
long, acute, acute or attenuate at the base, coarsely crenate to subentire, glabrous or
nearly so, rather fleshy, apparently pale green; corymbs small, dense, of 4-13 heads,
the peduncles long and slender, the pedicels short, bearing small subulate bractlets;
heads 5 mm. broad, purple or lavender; phyllaries narrowly lanceolate, attenuate, 2-
costate, glabrous; corollas 2.7 mm. long, minutely puberulent; achenes black,
glabrous, lustrous, 1.3 mm. long; pappus none or of 2-3 minute teeth, or the pappus
sometimes of 5 attenuate, setiferous scales.
Two forms of this species are reported from British Honduras
by Johnson, f. littorale and f. setigerum Rob., differing in that the
pappus scales of the latter are well developed and setiferous.
Ageratum peckii Rob. Proc. Am. Acad. 47: 191. 1911; I.e. 49:
458. 1913.
Pine savannas or ridges or in forest, 500 m. or less. Peten (Tun
951, 1609); British Honduras (type from Manatee Lagoon, Peck 80).
Perennial and suffrutescent or possibly flowering as annuals, the stems nodose
below, fastigiately branched or unbranched, glabrous, 1 m. tall or less; leaves linear-
oblong to narrowly elliptic, entire or serrulate, obscurely trinerved, acute or obtuse,
attenuate to a subpetiolar base, glabrous, 2.5-8 cm. long and 0.3-1.5 cm. broad;
inflorescence a sparse corymb or few-headed panicle; heads campanulate, 4.5-6 mm.
long and about as broad, with about 20 flowers; phyllaries bi- or triseriate, linear-
lanceolale, the apices indurated, bicostulate, glabrous or obscurely ciliolate, slightly
shorter than the capitulum; corolla cylindric, about 2 mm. long, the lobes triangular,
0.2 mm. long, blue or lavender; style at anthesis exserted about 1 mm., clavate,
purple; achenes glabrous, about 1.5 mm. long; pappus scales 5, free to the base, the
setiferous apex about as long as the corollas.
Described by Dr. Robinson as an annual but undoubtedly
usually a woody perennial. It is new to Guatemala.
Ageratum platylepis (Rob.) King & Robinson, Phytologia 24:
113. 1972; L. Wms. Fieldiana, Bot. 36: 81. 1975. Alomia platylepis
WILLIAMS: FLORA OF GUATEMALA 41
Rob. Proc. Am. Acad. 49: 448. 1913. Alomia guatemalensis Rob. I.e.
Ageratum benjamin- lincolnii King & Robinson, I.e.
Dry to wet, brushy or forested hillsides, often or usually in oak
forest, 900-1,800 m.; endemic; Guatemala; Chimaltenango; Huehue-
tenango (type of A. platylepis collected near Nenton, E. W. Nelson
3528); Santa Rosa (type of A. guatemalensis, Heyde & Lux 6153).
Stems terete, purplish, at first puberulent, glabrate in age; leaves opposite, short-
petiolate, linear-lanceolate to ovate-lanceolate, 7-10 cm. long, 0.8-3.8 cm. broad,
caudate-attenuate, rounded or acute at the base, rather remotely and obscurely
serrate, minutely glandular-punctate beneath, 3-nerved above the base; corymbs
terminal on the branches, dense, of 5-8 heads, the pedicels 3-5 mm. long; heads 7-9
mm. broad; involucres turbinate-campanulate; phyllaries oblong-lanceolate or linear-
lanceolate, acute, erose, costate, pale, subequal; receptacle paleaceous throughout,
the pales at anthesis shorter than the corollas, subulate at the apex, pale, indurate;
corollas 2.2-3 mm. long, glabrous, apparently blue-purple; achenes black, lustrous,
glabrous, 5-angulate, without pappus.
Ageratum radicans Rob. Proc. Am. Acad. 47: 192. 1911.
Known only from British Honduras, type M. E. Peck 99..
Herbaceous, glabrous, prostrate, and repent; leaves opposite, lanceolate or
narrowly elliptic-oblong, 4-8 cm. long, 5-15 mm. broad, narrowed to each end,
subobtuse, 3-nerved, subentire, not punctate beneath; cymes terminal, of only 2-3
heads, the peduncles elongate, naked; heads short- pedicellate, 8-10 mm. broad;
phyllaries lanceolate or linear, attenuate from almost the base to the narrow apex, 2-
costate, glabrous; receptacle small, convex, naked; corollas glabrous, probably
lavender; achenes acutely 5-angulate, glabrous, 1.2 mm. long; pappus scales 5, ovate,
setiferous at the apex, equaling the corolla.
Ageratum rugosum Coulter, Bot. Gaz. 20: 42. 1895. Figure 14.
Moist or wet thickets or open places, 900-1,400 m.; Alta
Verapaz; Santa Rosa (type from Santa Rosa, Heyde & Lux 4243);
Guatemala; Sacatepequez; Chimaltenango; Solola; Huehuetenango;
Saf Marcos. British Honduras and El Salvador to Costa Rica.
Plants perennial, erect, 1-2 m. high, branched, the stems terete, sordid-
villosulous, leafy throughout; leaves opposite, on petioles 1.5 cm. long or less, ovate,
rather thick and firm, 5-8 cm. long, acute, rounded at the base, serrate, mostly 3-
nerved, scaberulous and rugose above, somewhat paler beneath, densely pubescent or
tomentulose; corymbs terminal, rather large; heads numerous, 7 mm. broad, about
65-flowered; involucre turbinate-campanulate; phyllaries subequal, lance-linear,
attenuate, uncinate, 2-costate, hirtellous; receptacle conic, usually naked; corollas
pale blue, 2.5 mm. long, glandular-granulose; achenes 5-angulate, black, glabrous,
lustrous; pappus scales 5, deltoid, short, acute or obtuse, denticulate, connate for one-
third their length, or pappus reduced to a minute corona 0.1 mm. high.
In general appearance the plant is very like A. corymbosum,
and how it may differ is somewhat of a question.
42 FIELDIANA: BOTANY, VOLUME 24
EXCLUDED SPECIES
AGERATUM ECHIOIDES (Less.) Hemsl. Biol. Cent. Am. Hot. 2:
81. 1881; M. F. Johnson, Ann. Mo. Bot. Card. 58: 40. 1971.
Isocarpha echioides Less. Linnaea 5: 141, t. 2. 1830.
Excluded from the "Flora of Guatemala," since the specimens
cited by Johnson seem rather to be Ageratum isocarphoides, a
species not included in his monograph, since he thought it to be an
Alomia.
AGERATUM MARITIMUM HBK. Nov. Gen. Sp. 4: 150. 1820. A.
maritimum f. calvum Rob. Proc. Am. Acad. 49: 467. 1913.
Reported from the coastal keys of British Honduras, where it
may be expected. Johnson does not cite specimens except from
Cuba, Hispaniola, and Mexico. The species is much like Ageratum
littorale Gray.
BRICKELLIA Elliott
Reference: B. L. Robinson, A monograph of the genus
Brickellia, Mem. Gray Herb. 1: 1-151. 1917.
Annual or perennial herbs, or more often shrubs; leaves opposite or alternate,
sessile or petiolate, mostly dentate; inflorescence usually paniculate, sometimes
racemose, cymose, or corymbose, the corollas generally white, sometimes tinged with
pink or purple; heads rather small or medium-sized, homogamous, 3-many-flowered;
involucre cylindric to campanulate; phyllaries numerous, mostly thin and striate,
imbricate in several series, the innermost linear, the middle ones narrowly oblong,
graduate; receptacle naked, commonly flat, glabrous or short-pubescent; corollas
tubular, the limb 5-dentate; anthers appendaged at the apex, rounded at the base;
style branches clavellate, long-exserted; achenes prismatic, 10-costate, mostly
hispidulous, at least on the costae, sometimes sericeous or glandular-puberulent;
pappus bristles 10-80, capillary, equal or unequal, smooth or barbellate, mostly white,
sometimes fulvescent or tinged with purple, about equaling the corolla.
Species about 90, all American and chiefly in the warmer
regions, most abundant in Mexico. No other Central American
species are known. The generic name Brickellia Elliott is conserved.
Plants annual, glabrous or nearly so; leaves broadly ovate B. diffusa.
Plants perennial, herbaceous or usually woody, abundantly pubescent.
Leaves linear B. scoparia.
Leaves broadly ovate to elliptic-oblong.
Leaves sessile or on very short, thick petioles, oblong or elliptic, cuneate at the
base, thick, conspicuously reticulate-veined B. kellermanii.
Leaves slender-petiolate, all or mostly broadly ovate, chiefly rounded or
shallowly cordate at the base, comparatively thin, usually not conspicuously
reticulate-veined.
WILLIAMS: FLORA OF GUATEMALA 43
Outer phyllaries mostly half as long as the inner ones or longer... B. pacayensis.
Outer phyllaries very short, less than half as long as the inner ones.
B. paniculata.
Brickellia diffusa (Vahl) Gray, PI. Wright. 1: 86. 1852.
Eupatorium diffusion Vahl, Symb. Bot. 3: 94. 1794. Sierra picuda;
sabanera (fide Aguilar); visquita (Peten, fide Lundell); botoncillo
(fide Aguilar); arito (Jutiapa).
Moist or dry thickets, often a weed in waste ground, 1,600 m. or
less; reported from Peten; Alta Verapaz; Zacapa; Jutiapa; Santa
Rosa; Chiquimula; Escuintla; Guatemala; Sacatepequez; Chimal-
tenango; Huehuetenango; El Quiche. Mexico; Honduras and El
Salvador to Panama; Greater Antilles; tropical South America.
An erect annual, usually 1 m. high or less, sometimes as much as 2 m.,
glabrous almost throughout or somewhat puberulent, usually much branched, the
branches brittle, stramineous or brownish, slender; leaves slender-petiolate, thin,
bright green, rounded-ovate or deltoid-ovate, mostly 4-6 cm. long and 2.5-4 cm.
broad, acuminate, rounded or broadly cuneate at the base, sometimes gland-dotted,
serrate or crenate-dentate; heads very numerous, forming large panicles, the filiform
pedicels 8-18 mm. long; outer phyllaries minute, subulate, the middle and inner ones
lance-linear, 2-3-nerved, subscarious, pale green; heads about 8-flowered, 8 mm. long,
greenish white; corollas 4-4.3 mm. long; achenes 1.7-2 mm. long, fuscous-grayish,
villous above; pappus bristles about 24, white.
Called "culantrillo" in Chiapas. A common weed in many parts
of the Central American lowlands.
Brickellia kellermanii Greenm. Field Mus. Bot. 2: 265. 1907.
Dry or moist, rocky hillsides, mostly in pine-oak forest,
sometimes on serpentine, 160-1,950 m.; Alta Verapaz; Baja Verapaz
(type from Sierra de las Minas, W. A. Kellerman 6127); Zacapa;
Jalapa; Chimaltenango; Huehuetenango. Mountains of Honduras.
A stiff shrub about 1 m. high, grayish tomentulose almost throughout, the stems
branched, the branches terete, suberect; leaves almost sessile, very thick, oblong to
oblong-elliptic, mostly 3-6 cm. long and 1-2 cm. broad, obtuse or acute, obtuse or
cuneate at the base, crenate-dentate or entire, pale green and hirtellous above,
whitish tomentulose beneath, the veins very prominent and closely reticulate;
inflorescence leafy, subcorymbose-paniculate, the branches often elongate and
racemiform; heads whitish or pale lilac, 12-14 mm. long., mostly 12-flowered, sessile
or nearly so; phyllaries pubescent, graduate, ciliate, the outer ones ovate-oblong,
mucronate, the inner ones lance-linear, acute; pappus bristles white, 6 mm. long.
Forma podocephala Rob. (Contr. Gray Herb. 68: 43. 1923; type
collected in "Guatemala" by Friedrichsthal) is a form in which the
heads are pedicellate.
44 FIELDIANA: BOTANY, VOLUME 24
Lundell has reported from Peten (La Libertad) Brickellia
oliganthes (Less.) Gray, said to grow there in grasslands. That
Mexican species is very similar to B. kellermanii, if not the same.
Brickellia pacayensis Coulter, Bot. Gaz. 16: 98. 1891; L.
Wms. Fieldiana, Bot. 36: 81. 1975. Coleosanthus pacayensis Coulter,
Bot. Gaz. 20: 46. 1895.
Dry or moist thickets, 750-1,800 m.; Zacapa; Santa Rosa;
Guatemala (type from Pacaya, J. D. Smith 2389); Sacatepequez;
Huehuetenango; Solola. Southern Mexico; Honduras; El Salvador;
Nicaragua.
A stout shrub, 1.5-2.5 m. high, often densely branched, the branches stramineous
or ochraceous, when young grayish pubescent or tomentulose, glandular above; leaves
on petioles 1-2 cm. long, broadly ovate or deltoid-ovate, 4-7 cm. long, 3-4 cm. broad,
3-nerved, acute or acutish, rounded or subcordate at the base, crenate, thick,
scabrous-puberulent above, grayish tomentose and somewhat reticulate-veined
beneath; heads 11-13 mm. high, in lax or dense, leafy panicles, 22-25-flowered, pale
yellow or whitish, the pedicels 1-2.5 cm. long, capitate-glandular; phyllaries lance-
oblong, mostly acute, the outer ones almost equaling the inner ones, densely
glandular-puberulent or glandular-hirtellous; receptacle hirtellous; corollas 5.5-7 mm.
long; achenes 2.3-2.9 mm. long, villous above, often glandular; pappus bristles 40-45,
white, minutely barbellate.
Distinguished from B. paniculata principally by the longer
outer phyllaries. Intergression undoubtedly occurs.
Brickellia paniculata (Miller) Rob. Proc. Am. Acad. 42: 48.
1906; L. Wms. Fieldiana, Bot. 36: 81. 1975. Eupatorium panic-
ulatum Miller, Gard. Diet. ed. 8, No. 15. 1768. E. rigidum Benth. PI.
Hartweg. 88. 1841, not Sw. (type from valley of Guatemala,
Hartweg 598). Brickellia hartwegii Gray, PL Wright. 1: 85. 1852.
Coleosanthus rigidus O. Kuntze, Rev. Gen. pi. 1: 328. 1891.
Brickellia argyrolepis Rob. Mem. Gray Herb. 1: 90, fig. 69. 1917
(type from Costa Rica, collection not specified). B. guatemalensis
Rob. I.e. 94, fig. 71 (type from Alta Verapaz, Tuerckheim II. 2145).
B. adenocarpa Rob. I.e. 94, fig. 73 (type from Quezaltenango,
Holway 466). B. adenocarpa var. glandulipes Rob. I.e. (type from
Quezaltenango, Holway 92). Figure 14a.
Open forests, often in oak-pine association, moist or drying
thickets, open hillsides or plains, 1,500-2,700 m.; Alta Verapaz;
Santa Rosa; Zacapa; Chiquimula; Guatemala; Sacatepequez;
Chimaltenango; Solola; Huehuetenango; Quezaltenango; San
Marcos. Mexico; El Salvador; Honduras; Nicaragua; Costa Rica.
WILLIAMS: FLORA OF GUATEMALA 45
A stout, often much branched, shrub to 2 m. tall, the branches crisped-
puberulent when young and often glandular, glabrate and brownish in age; leaves
ovate to broadly ovate-cordate, acute or acuminate, sparsely to sharply crenate-
serrate, trinerved or subtriplinerved, densely to sparsely puberulent above, usually
densely and softly grayish tomentulose beneath but sometimes sparsely tomentulose
and greenish (B. guatemalensis), blades 3-7 cm. long, petioles 2-4 mm. long or in
Guatemalan material mostly 1.5-3 cm. long; inflorescence sparse to dense and many-
headed, corymbose panicles, the pedicels prominently puberulent to densely stipitate-
glandular pubescent; heads about 15-25-flowered, 12-18 mm. high; phyllaries 3-5-
seriate, about 20, 5-striate, the outer mostly ovate-lanceolate, acute, 2 mm. long or
more, sparsely hirsute and often glandular, graduating to linear or linear-lanceolate,
mostly acuminate inner ones, usually 15 mm. long or less; corolla narrowly cylindric,
mostly 7-8 mm. long, the lobes very small; achenes 3.5-5 mm. long, densely hirsute,
glandular or not; pappus white, barbellate, usually slightly shorter than the corollas,
of 50 bristles or fewer.
This is an exceedingly common and variable species at middle
elevations in Guatemala— in fact, throughout its range. There seems
to be no way to segregate the several species described or credited to
Guatemala, although trends are to be noticed in them. Some have
relatively sparse inflorescences with smaller fewer-flowered heads.
The pubescence on most parts of the plants is from very dense to
quite sparse and sometimes glandular. There are several other
species from Mexico involved in this complex, these may include B.
squarrosa (Cav.) Rob. and B. hebecarpa (DC.) Gray.
Brickellia scoparia (DC.) Gray, PL Wright. 1: 84. 1852.
Clavigera scoparia DC. Prodr. 5: 128. 1836.
Open, grassy, often rocky slopes, frequently in pine-oak forest,
sometimes on limestone, 1,000-2,000 m.; Jalapa; Guatemala;
Huehuetenango; Quezaltenango. Mexico.
Plants herbaceous or suffrutescent only at the base, stiffly erect, 1 m. high or
usually lower, the stems often numerous, mostly simple below but with erect narrow
branches above, usually purplish, puberulent or glabrate; leaves crowded, linear,
entire, 2-6 cm. long, 1-5 mm. broad, scabrous above, slightly paler beneath and
punctate, ascending or suberect; inflorescences 20-30 cm. long, narrow and spike-like
or with numerous erect branches; heads 12 mm. long, short-pedicellate, subcylindric;
phyllaries purplish, 3-5-nerved, oblong, rounded at the apex and mucronulate,
glandular near the apex.
This is presumably the plant reported from Guatemala by
Hemsley as B. corymbosa (DC.) Gray.
CARMINATIA Mocino
Erect annuals, weak and rather fragile, more or less pilose; leaves opposite, on
long slender petioles, very thin and delicate, broad, dentate; heads homogamous,
46 FIELDIANA: BOTANY, VOLUME 24
without rays, medium-sized, in more or less secund, elongate racemes, solitary or
usually glomerate, short-pedicellate; involucre cylindric, phyllaries few, imbricate,
thin, narrow, the outer ones much shorter; receptacle flat, naked; corollas pale
greenish or whitish, regular, slender, the limb scarcely broader than the tube, shortly
5-fid, scarcely exceeding the phyllaries and pappus; anthers not appendaged, obtuse
and entire at the base; style branches short-exserted, subacute, glabrous; achenes 5-
angulate, glabrous or sparsely glandular; pappus bristles numerous, 1-seriate, long-
plumose.
The genus consists of a single species and is the only species of
this tribe in our region with plumose pappus bristles.
Carminatia tenuiflora DC. Prodr. 7: 267. 1838. Figure 15.
Moist thickets or open oak or pine forest, 300-1,950 m.;
Chiquimula; Jalapa; Jutiapa; Santa Rosa; Huehuetenango. South-
western United States; Mexico; El Salvador.
An erect annual, sometimes 1 m. high but usually lower, simple or more often
sparsely branched, almost glabrous but with sparse weak white hairs on almost all
parts, the stems very brittle, subterete, pale green; leaves pale green, very thin,
rounded- ovate, 2.5-10 cm. long, obtuse or acute, broadly rounded or subcordate at the
base and abruptly short-decurrent, irregularly and not conspicuously undulate-
dentate, 3-nerved at the base; heads usually numerous or sometimes few, about 1.5
cm. long, the racemes simple or paniculate; phyllaries pale green, erect, linear, very
unequal, scarious-margined, 3-nerved, almost dry; corollas whitish, glabrous; achenes
linear, black or dark olive, 6-8 mm. long, glabrous and smooth or often glandular or
glandular-puberulent, some of them often densely transverse-rugose; pappus long and
soft, white or dirty white, longer than the achenes, the bristles plumose.
This is a frequent plant in moist shaded places in eastern
Guatemala, found only during the wetter months, since it withers
almost as soon as the rains cease. While the plants, throughout
their range, are alike in general appearance, the achenes exhibit a
great deal of variation in size, coloring, pubescence, and sculpturing
of the surface.
EUPATORIUM Linnaeus
Reference: B. L. Robinson in Standley, Contr. U. S. Nat. Herb.
23: 1432-1469. 1926 (account of the woody species of Mexico).
Perennial or rarely annual herbs, in Central America more often shrubs or small
trees, various in habit, rarely epiphytic; leaves mostly opposite and petiolate, filiform
to orbicular, membranaceous to coriaceous, entire or dentate, very rarely compound;
heads usually small, homogamous, generally 5-100-flowered, commonly in corymbose
or thyrsoid panicles; corollas mostly white, pink, purple, or bluish, rarely ochrolecous;
involucre of few to numerous phyllaries; phyllaries narrowly or broadly subequal or
often very unequal and graduate, laxly or closely imbricate; anthers usually
appendaged at the apex, entire at the base; style branches long, at maturity long-
WILLIAMS: FLORA OF GUATEMALA 47
exserted from the corolla, thread-like or clavellate, often colored; achenes columnar
to obovoid, usually 5-costate or 5-angulate; pappus of numerous or few, slender or
rather stiff bristles, rarely epappose.
Eupatorium is a vast genus of probably 600 species or more,
chiefly in tropical regions of America, but a substantial number of
them extend into temperate regions. In Central America the species
are found mostly in the mountains at middle or high
elevations. In Mexico more than 100 species with more or less
woody stems are found. In Central America the group is best
represented in Guatemala, but numerous additional species occur in
the mountains of Costa Rica. A very large number of the
Guatemalan species were described by Dr. B. L. Robinson, who
prepared a monograph of the genus, unfortunately never published.
He was responsible for first bringing order out of the confusion that
had prevailed previously with regard to the species of Mexico and
Central America, as well as those of extra-Brazilian South America.
It should be noted that we have used a somewhat different
terminology than that employed by Robinson in describing the
nervation of the leaves of this genus. He describes as penninerved
leaves which we should consider plinerved, if that term is to be
employed in uniformity with its use in the family Melastomaceae,
where it is best exhibited. On the following pages leaves are
described as plinerved when there are two or more conspicuous
lateral nerves arising in the basal portion of the leaf blade and
conspicuously continued for a long distance above the point of
divergence. Sometimes these more conspicuous nerves arise at a
considerable distance above the base of the blade, and in rare cases
they are somewhat ambiguous, but generally they are much more
conspicuous than the lateral nerves arising above them.
The genus Eupatorium has been subjected to considerable
fragmentation in recent years, principally by Robert M. King and
Harold Robinson. More than 50 eupatorioid genera have been
resurrected or established by these gentlemen. Eighteen of these
segregates have been applied to species which occur in Guatemala.
A list of these genera, considered by the junior author to be
synonyms of Eupatorium, follows: Ageratina, Amolinia, Ayapana,
Bartlettina (Neobartlettia), Chromolaena, Critonia, Critonia-
delphus, Decachaeta, Eupatoriastrum, Fleishmannia, Fleishmann-
iopsis, Heterocondylus, Koanophyllon, Neomirandea, Pach-
ythamnus, Peteravenia, Piptothrix, and Pseudokyrsteniopsis. The
specific synonyms will be found cited in the appropriate places.
48 FIELDIANA: BOTANY, VOLUME 24
Involucre cylindric and 3-5 times as long as broad or rarely only twice as long; the
phyllaries 3-5 or more seriate and closely imbricated and appressed.
SERIES A, p. 48.
Involucre normally campanulate or turbinate, not cylindric, rarely more than twice
as long as thick and often much shorter; the phyllaries closely imbricate or in
age lax, often subequal.
Phyllaries more or less or very unequal, in 3 or more series with the outer ones
gradually shorter.
Leaves penninerved or essentially so, if rarely triplinerved the nerves arising very
far above the base of the blade, usually little below the middle; the blades
mostly obtuse or acute at base, never rounded or cordate.... SERIES B, p. 49.
Leaves 3-5-nerved or 3-5-plinerved, the lower nerves much more conspicuous
than those above SERIES C, p. 50.
Phyllaries subequal in length, in about two scarcely imbricated series, usually 1-3
of the outermost phyllaries shorter than the others SERIES D, p. 52.
SERIES A. The involucre cylindric and mostly 3-5 times longer than wide; the
phyllaries closely imbricated and appressed in 3-5 series.
Heads 4-6-flowered.
Leaf blades acute or long attenuate at the base.
Leaves conspicuously pubescent on both surfaces; stems not conspicuously
angulate, at most striate, pubescent E. macrum.
Leaves glabrous; stems angulate to nearly terete, glabrous.
Stems prominently 6-alate or 6-angulate; leaves penninerved; highland species.
E. sexangulare.
Stems nearly smooth; leaves triplinerved from near the base; lowland species.
E. campechense.
Leaf blades obtuse at the base.
Heads in small dense clusters, sessile E. bartlettii.
Heads corymbose-paniculate, slender pedicellate.
Leaves 3-5-nerved from the very base of the blade E. rojasianum.
Leaves triplinerved E. campechense.
Head 10-40-flowered.
Plants herbaceous; leaf blades attenuate at the base, narrowly lanceolate or linear-
lanceolate; stems abundantly villous E. ivaefolium.
Plants woody; leaf blades usually obtuse to truncate at the base, sometimes acute
but then the stems glabrous.
Leaves penninerved.
Stems below the inflorescence villous-hirsute with spreading brownish hairs;
heads mostly less than twice as long as broad E. lanicaule.
Stems below the inflorescence crisped pubescent with short dark brown hairs
or glabrate; heads more than twice as long as broad.
Leaf bases acute with petioles 4-10 mm. long; heads with 15-20 flowers.
E. quercetorum.
Leaf bases subcordate to acute, sessile or nearly so; heads with about 40
flowers E. glaberrimum.
Leaves 3-nerved or 3-5-plinerved.
WILLIAMS: FLORA OF GUATEMALA 49
Leaf venation 5-plinerved; heads about twice as long as broad E. magistri.
Leaf venation 3-nerved or triplinerved; heads 3 times longer than broad or
more.
Leaves glabrous or essentially so, viscid-resinous E. laevigatum.
Leaves abundantly pubescent, not viscid-resinous E. odoratum.
SERIES B. The involucre normally campanula te or turbinate, not cylindric; the
phyllaries more or less or very unequal, in three or more series with outer ones
gradually shorter; the leaves penninerved or essentially so, if rarely
triplinerved the nerves arising very far above the base of the blade, usually
little below the middle, the blades mostly obtuse or accute at the base, never
rounded or cordate.
Leaves sessile or practically so.
Stems 4-angulate; leaves membranaceous, mostly 15-25 cm. long or larger; heads
white E. quadr angular e.
Stems subterete; leaves coriaceous, mostly 9 cm. long or shorter; heads purple or
pink E. amygdalinum.
Leaves conspicuously petiolate.
Dry leaves, when examined under a strong light, showing pellucid dots or pellucid
veins or both.
Leaves with pellucid dots and sometimes also with pellucid veins.
Veins conspicuously impressed on the upper leaf surface.
Leaf blades very obtuse to broadly rounded at the base E. morifolium.
Leaf blades attenuate to the base E. lanicaule.
Veins not impressed on the upper leaf surface; leaf blades acute or attenuate
at the base.
Branches of the inflorescence glabrous or obscurely puberulent.
E. sexangulare.
Branches of the inflorescence densely pubescent.
Heads pedicellate E. hebebotryum.
Heads sessile.
Leaves brownish tomentose or villosulous beneath, often very sparsely
so, conspicuously and densely pellucid punctate, serrate.
E. daleoides.
Leaves glabrous beneath, obscurely and sparsely punctate, cuspidate-
denticulate E. nubigenum.
Leaves with pellucid veins but without pellucid dots.
Branches of the inflorescence glabrous or essentially so.
Heads purple or lavender; outer phyllaries acute or acuminate.
E. tuerckheimii.
Heads white; outer phyllaries very obtuse E. nubigenum.
Branches of the inflorescence densely pubescent E. luxii.
Dry leaves opaque when examined under a strong light, without translucent dots
or veins.
Veins obscure beneath or obsolete, few and very openly branched; heads
lavender or purple.
Leaves conspicuously serrate E. pinabetense.
Leaves entire E. araliae folium.
Veins conspicuous beneath, numerous, reticulate; heads greenish or white.
Heads 9-11 mm. long E. caeciliae.
50 FIELDIANA: BOTANY, VOLUME 24
Heads 5-6 mm. long.
Lower side of leaves densely and softly pubescent over the whole surface.
Leaves elliptic-oblong or elliptic-ovate; outer phyllaries ovate.
E. montigenum.
Leaves lanceolate or lance-oblong; outer phyllaries lanceolate.
Bases of the leaves cuneately narrowed, the blade elliptic.
E. hypomalacum.
Bases of the leaves acute, the blade lanceolate E. sorensenii.
Lower side of leaves glabrous or glabrate except sometimes on the nerves
and veins.
Phyllaries acute or subacute E. galeottii.
Phyllaries very obtuse and rounded at the apex.
Leaves beneath not prominently reticulate, pubescent at least on the
veins E. luxii.
Leaves beneath prominently reticulate and glabrous.
Leaf margins serrate, with 4-6 pairs of secondary nerves; the
phyllaries much shorter than the pappus E. pittieri.
Leaf margins entire, with 6-10 pairs of secondary nerves, the
phyllaries as long as the pappus E. lucentifolium.
SERIES C. The involucre normally campanulate or turbinate, not cylindric; the
phyllaries more or less or very unequal, in three or more series with the outer
series gradually shorter; the leaves 3-5-nerved or 3-5-plinerved, the lower
nerves much more conspicuous than those above.
Leaf blades acute or attenuate at the base.
Leaves triplinerved, the nerves arising well above the base of the blade.
Heads sessile E. nubigenoid.es.
Heads pedicellate.
Leaves glabrous to evenly and usually sparsely puberulent below;
inflorescences densely headed and the phyllaries whitish.
E. leucocephalum.
Leaves sparsely puberulent, principally along the nerves below; inflorescence
few-headed and the phyllaries greenish E. macrum.
Leaves 3-nerved, the nerves arising at the very base of the blade.
Phyllaries acute or acuminate, puberulent E. imitans.
Phyllaries very obtuse, glabrous or glabrate.
Leaves serrate, thin, acutely attenuate at the apex E. blakei.
Leaves entire or obscurely crenate, thick, obtuse E. crocodilium.
Leaf blades cordate to rounded or very obtuse at the base.
Principal nerves of the leaves arising well above the base of the blade; leaves never
angulate.
Phyllaries pubescent.
Leaves 5-plinerved, with conspicuous, elevated, reticulated veination below.
E. hypodictyon.
Leaves triplinerved.
Phyllaries lance-linear, subequal; leaf veins inconspicuous below; usually
shrubs E. albicaule.
Phyllaries ovate to lanceolate or elliptic, prominently unequal; leaf veins
conspicuous below; vines E. tunii.
WILLIAMS: FLORA OF GUATEMALA 51
Phyllaries glabrous; leaves with the veins obsolete or nearly so, triplinerved.
E. campechense.
Principal nerves of the leaves arising from the very base of the blade or very close
to it; leaves sometimes (rarely) angulate.
Receptacle densely pilose; heads with numerous white flowers; a tall herb.
E. macrophyllum.
Receptacle glabrous.
Phyllaries very closely appressed, even in age, puberulent; heads about 7 mm.
high, a shrub, woody almost throughout E. collinum.
Phyllaries not closely appressed, in age often very lax.
Leaves conspicuously angulate, and acutely dentate along the whole margin,
about as broad as long; plants herbaceous; heads lavender, with linear
or lance-linear phyllaries E. angulifolium.
Leaves not angulate, rarely hastate-lobate at the base.
Heads large, usually many-flowered, mostly 6-10 mm. long or even larger,
often broadly campanulate; shrubs, vines or large coarse herbs.
Base of the petioles thickened and surrounding the stems.
E. perpetiolatum.
Base of the petioles not thickened, if surrounding the stems then thin.
Leaves appearing glabrous beneath, the pubescence mostly confined
to the nerves and of minute, chiefly appressed hairs.
Heads cylindric E. leucocephalum.
Heads broadly campanulate.
Involucre about 7 mm. high E. hastiferum.
Involucre about 10-14 mm. high.
Receptacle with many pales about as long as the flowers.
E. ultraisthmium.
Receptacle not paleaceous.
Leaves at least twice as long as broad, shining; usually vines.
E. vitalbae.
Leaves less than twice as long as broad, surface dull; weak
shrubs E. platyphyllum.
Leaves usually densely pubescent beneath, rarely glabrate, the
pubescence of the nerves of obvious spreading hairs.
Outer phyllaries acute or subacute.
Outermost phyllaries broadly ovate or oval; phyllaries usually
purple or purplish; leaf blades cordate at the base.
E. phoenicolepis.
Outermost phyllaries linear to lanceolate, phyllaries usually green
or whitish, rarely tinged with purple.
Blades of the larger leaves cordate at the base; inflorescence
thyrsiform E. cupressorum.
Blades of the leaves not or scarcely cordate at the base;
inflorescence not thyrsiform E. pycnocephaloides.
Outer phyllaries rounded or very obtuse at the apex, the inner ones
usually pale and not at all colored.
Stems villous, with obvious spreading hairs; leaves with elevated,
very conspicuous, reticulate veins beneath E. griseum.
Stems closely puberulent, the pubescence mostly appressed, the
hairs not evident and spreading; veins of the leaves not
52 FIELDIANA: BOTANY, VOLUME 24
conspicuously elevated and reticulate beneath E. schultzii.
Heads small, mostly 5 mm. long or less.
Plants shrubs to about 1 m. tall; leaves densely gland-dotted below.
E. scoparioides.
Plants herbs, often or usually low, rarely elongate but then soft-
stemmed throughout; leaves not densely gland-dotted below.
Pedicels glabrous or merely viscid; corollas white.
Pedicels viscid; leaves glabrous beneath E. viscidipes.
Pedicels glabrous, not viscid; leaves usually abundantly pubescent
beneath E. multinerve.
Pedicels puberulent or pilosulous, usually conspicuously so, but the
pubescence sometimes very minute; heads white or purple.
Heads clustered in small rounded glomerules, short-pedicellate or
subsessile, the glomerules forming small or often large, leafy
panicles E. pycnocephalum.
Heads not glomerate, either on long slender pedicels forming open
lax inflorescences or on short pedicels forming subracemose
panicles.
Heads on short pedicels forming subracemose panicles.
E. solidaginoides.
Heads on long slender pedicels forming open and often lax
inflorescences.
Stems villous with lax spreading hairs E. antiquorum.
Stems glabrous or closely puberulent.
Heads about 3 mm. long; outer phyllaries acute or
acuminate E. sinclairii.
Heads about 5 mm. long; outer phyllaries obtuse or
subacute E. microstemon.
SERIES D. The involucre normally campanulate or turbinate, not cylindric, rarely
more than twice as long as thick and usually much shorter; phyllaries
subequal in length, in about two scarcely imbricated series, usually 1-3 of the
outermost phyllaries shorter than the others; rarely the pappus none.
Pappus none E. sodali.
Pappus present.
Leaves divided into numerous linear segments E. carletonii.
Leaves entire or dentate.
Leaf blades acute or acuminate at the base.
Plants trees or shrubs.
Leaves triplinerved E. heydeanum.
Leaves penninerved.
Flower heads 9-15 mm. long.
Leaves prominently reticulate-veined, shining and glabrous below.
E. molinae.
Leaves not prominently reticulate-veined, dull and sordid pubescent
below E. caeciliae.
Flower heads about 6-7 mm. long E. semialatum.
Plants herbaceous.
Phyllaries acute or acuminate.
WILLIAMS: FLORA OF GUATEMALA 53
Leaves 5-plinerved well above the base; heads 5 mm. high or less; plants of
middle elevations E. carmonis.
Leaves 3-plinerved from the base; heads 7 mm. high or more; plants of
high elevations E. nubivagum.
Phyllaries obtuse or rounded at the apex.
Leaf blades narrowly decurrent to the base of the petiole, very densely
short-pilose or villosulous beneath E. incomptum.
Leaf blades not long decurrent on the petiole, thinly villous beneath.
E. huehuetecum.
Leaf blades cordate or rounded or very obtuse at the base.
Heads 3-5-flowered; pappus present or absent E. sodali.
Heads with 8-15 or more numerous flowers; pappus always present.
Heads with usually 8-10 flowers.
Leaf blades rounded or very obtuse at the base, glabrous E. albicaule.
Leaf blades truncate or cordate at the base, at least the principal ones
puberulent on the nerves.
Heads about 5 mm. long E. solidaginoides.
Heads mostly 7-10 mm. long.
Leaves obtuse or narrowed to an obtuse apex.
Lower leaf surface tomentellose-glandular E. tomentellum.
Lower leaf surface glabrate E. ovillum.
Leaves acuminate, with a very acute tip.
Upper side of leaves sparsely villosulous with pale or thick hairs.
E. mimicum.
Upper side of leaves puberulent along the nerves E. coulteri.
Heads with 12-15 or often more numerous flowers, at least with more than
10 flowers in all or most of the heads.
Pedicels much elongated, usually several times as long as the heads; heads
broadly campanulate, 6-7 mm. long; plants low herbs with usually
simple stems.
Phyllaries and pedicels glandular pubescent E. bettidifolium.
Phyllaries and pedicels not glandular pubescent.
Leaves cordate, 3-nerved from the apex of the petiole ...E. anchisteum.
Leaves acute or at most obtuse at the base, penninerved.....E. muelleri.
Pedicels short, usually little if at all exceeding the heads, generally
aggregate; heads not broadly campanulate, or if so only 3 mm. long.
Flowering branches leafless, very thick, woody throughout, the leaves
deciduous E. crassirameum.
Flowering branches leafy, relatively slender, often herbaceous; leaves
not deciduous.
Leaf apices rounded or very obtuse; a shrub, woody throughout, with
pink heads E. tomentellum.
Leaf apices acute to long-acuminate or, if rarely obtuse, the plants
herbaceous.
Leaves 3-5-nerved, the nerves arising at or very close to the base of
the blade.
Heads all sessile or nearly so; leaves short-petiolate, thick and
firm E. areolare.
Heads all or mostly conspicuously pedicellate; leaves long or
short petiolate, thin and soft.
54 FIELDIANA: BOTANY, VOLUME 24
Heads small, 3-5 mm. long, petioles long and slender.
Heads about 3 mm. long; plants annual E. jejunum.
Heads about 5 mm. long; plants perennial.
Leaves coarsely crenate, with few crenations; plants
herbaceous throughout; heads on long, slender, almost
filiform pedicels E. capiliipes.
Leaves not crenate or, if so, the crenations very numerous;
plants often suffrutescent; heads usually densely
fastigiate on short, rather stout pedicels.
E. aschenbornianum.
Heads large, mostly 7-11 mm. long.
Plants low, herbaceous; leaves mostly 3-4 cm. long.
E. prunellae folium.
Plants tall shrubs; leaves mostly 6-12 cm. long or larger.
Leaves subcordate or obtuse at the base; petioles less
than 1 cm. long, densely villous E. cupressorum.
Leaves not subcordate to obtuse at the base; petioles much
more than 1 cm. long, glabrous to sparsely villous.
Leaves deltoid, villous beneath on the nerves.
E. zunilanum.
Leaves not deltoid, the nerves beneath with short,
mostly appressed hairs E. pazcuarense.
Leaves with 3-5 or more numerous nerves arising well above the
base of the bland.
Stems appearing glabrous but with a minute appressed tomen-
tum, no spreading hairs present.
Leaves subtending inflorescences 5 cm. long or less, shorter
than the inflorescence; heads more than 10 mm. long.
E. mairetianum.
Leaves subtending inflorescences 15 cm. long or more, longer
than the inflorescence; heads 6-7 mm. \ong....E. ovandense.
Stems with variable pubescence but always with evident
spreading hairs.
Plants herbaceous.
Lower side of leaves villous or hirsutulous, at least on the
nerves E. aschenbornianum.
Lower side of leaves almost glabrous, the hairs of the nerves
very short and appressed.
Leaves large, mostly 5-9 cm. long; plants tall, often 1
m. high E. pazcuarense.
Leaves small, 1-2 cm. long, plants low, 25 cm. high or less.
E. saxorum.
Plants large shrubs or trees.
Branches of the inflorescence bearing numerous stipitate
glands E. chiapense.
Branches of the inflorescence without gland-tipped hairs, but
the hairs often viscid.
Heads lilac or purple; phyllaries usually purplish, hardly
longer than the achenes E. monticola.
WILLIAMS: FLORA OF GUATEMALA 55
Heads white; phyllaries green, not tinged with purple,
nearly as long as the florets E. pazcuarense.
Eupatorium albicaule Sch.-Bip. ex Klatt, Leopoldina 20: 89.
1884. Koanophyllon albicaulis King & Robinson, Phytologia 22:
149. 1971. Ixhotz (Peten, Maya, fide Lundell).
Moist or wet open forest, at or little above sea level; Peten.
Southern Mexico; British Honduras; Honduras.
A shrub or tree 3-9 m. high, sometimes subscandent, the branches terete, usually
ochraceous, when young sparsely puberulent or fulvous-tomentulose; leaves on short,
slender petioles, oblong or lance-oblong, 5-12 cm. long, 1.5-4 cm. broad, falcate-
acuminate, rounded or obtuse at the base, triplinerved well above the base, with the
lateral nerves prolonged almost to the apex, serrate or subentire, chartaceous,
glabrous or nearly so, usually blackish when dried; heads white, 7-8-flowered, 6-7 mm.
long, densely crowded in small rounded clusters, these forming dense, leafy, rounded
corymbs, short-pedicellate, the branches of the inflorescence appressed-puberulent or
tomentulose; phyllaries lance-linear, subequal, with a few very short ones at the base
of the involucre, minutely brownish tomentulose, acute, much shorter than the
florets; corollas slender, tubular; achenes glabrous, blackish; pappus ample,
fulvescent.
Known in British Honduras as "old woman's walking stick,"
"soscha," and "xoltexnuc" (Maya); "zactocaban," "xicin" (Yucatan,
Maya); "putinin," "tine— cordel" (Honduras). An unattractive
shrub, used in Honduras as a green dye for twine, cloth, and other
textiles.
Eupatorium amygdalinum Lam. Encycl. 2: 408. 1786.
Ayapana amygdalina King & Robinson, Phytologia 20: 211. 1970.
Probably in open pine forest, little above sea level; Izabal (Los
Amates, W. A. Kellerman 7605). Honduras; Costa Rica; Panama;
Colombia; Venezuela.
Plants suffrutescent, usually 75 cm. high or less, stout, simple or sparsely
branched, glabrous or sparsely pubescent, often somewhat viscid; leaves opposite,
sessile, coriaceous, lance-oblong to elliptic-oblong or ovate-oblong, 5-10 cm. long, 2-4
cm. broad, usually obtuse, cuneate or obtuse at the base, subentire or crenate-serrate,
with conspicuous venation; panicles terminal, somewhat fastigiately branched, 3-10
cm. broad or larger; heads numerous, about 40-flowered, 7-8 mm. high, rose-pink or
red-purple; involucre turbinate-campanulate; phyllaries about 40, linear, very
unequal, graduate, purplish, puberulent; achenes blackish, puberulent on the angles.
Eupatorium anchisteum Grashoff & Beaman, Rhodora 71:
567. 1969. Ageratina anchistea King & Robinson, Phytologia 24: 80.
1972.
56 FIELDIANA: BOTANY, VOLUME 24
In open oak-pine woods, 900-1,800 m.; Jalapa; Chiquimula;
Guatemala; Sacatepequez; Chimaltenango. Honduras (type, Stand-
ley 27471).
Erect or decumbent, perennial herbs to 1 m. tall, the stems short pilose with
multicellular hairs, becoming glabrescent above; leaves ovate-cordate, crenate, acute
or short-acuminate, sparsely crisped pubescent on both surfaces, 3-7 cm. long and 2-6
cm. broad, 3-nerved from the apex of the petiole, petiole pilose, to 4 cm. long, slender;
inflorescence a few-many-flowered, cymose panicle, pedicels long, subfiliform,
glabrous, to 15 mm. long; heads campanulate, about 5 mm. high and as broad;
phyllaries linear-oblong, obtuse, apex sometimes erose, glabrous, scarious; achenes
about 1.5 mm. long, prismatic, black, sparsely pubescent on the ridges; pappus
bristles about 3 mm. long, barbellate; corolla white, about 2.5 mm. long.
Closely related to E. muelleri and perhaps only a variety of it.
Eupatorium angulifolium Rob. Contr. Gray Herb. 65: 46.
1922. Eupatoriastrum angulifolium King & Robinson, Phytologia
21: 306. 1971. Madre contrahierba (fide Aguilar); flor de algodon
( Quezaltenango) .
Moist or wet, shaded, usually vertical banks or cliffs, mostly on
the sides of quebradas or along stream banks, 500-1,800 m.;
Escuintla; Guatemala; Sacatepequez (type from Barranco Hondo,
Salvin & Godman 265); Chimaltenango; Suchitepequez; Quezal-
tenango. Mexico (Chiapas).
Plants perennial from a cluster of rather thick, more or less woody roots,
herbaceous, almost wholly glabrous, the stems short, usually 25 cm. long or less,
bearing few or numerous, often crowded or almost whorled leaves; leaves opposite,
very large and thin, on long slender petioles, irregularly rounded and more or less
angulate, mostly 10-18 cm. long and broad or sometimes larger, glabrous, 7-9-
plinerved from near the base, laxly reticulate-veined, slightly paler beneath, the
angles acute, coarsely dentate, the base of the blade rounded or shallowly cordate;
inflorescence terminal, corymbose, usually long-pedunculate, rather flat-topped,
open, the slender pedicels 3 cm. long or less, glabrous; heads very few or rather
numerous, lavender, about 1 cm. long; 75-100-flowered; involucre campanulate;
phyllaries numerous, lanceolate, 3-4-seriate, laxly imbricate, striate, acute, ciliolate;
achenes pale brown, about 1.4-2 mm. long, sparsely scaberulous on the angles; pappus
bristles few, white, equaling the corolla, caducous.
The plants grow plentifully in the foothills along the roads
between Antigua and Escuintla, especially in the vicinity of the
original locality. They form colonies along the vertical banks and
cliffs with Begonias and other plants. During the rainy season they
are conspicuous and rather handsome on the roadsides, but the
leaves droop and quickly wither after the dry season begins.
WILLIAMS: FLORA OF GUATEMALA 57
Eupatorium antiquorum Standl. & Steyerm. Field Mus. Bot.
22: 302. 1940.
Moist or rather dry, shaded, mountain slopes, 1,500-1,800 m.;
endemic; Escuintla (between Santa Maria de Jesus and Palin);
Sacatepequez (type collected at Antigua, Standley 60304, growing in
a grove of planted Cupressus).
An erect perennial herb almost 1 m. high, the stems solitary or several, simple or
sparsely branched, densely villous with long, spreading, white, several-celled hairs;
leaves opposite, remote, on slender petioles 2-3 cm. long, membranaceous, deltoid or
ovate-deltoid, 5-6.5 cm. long, 3.5-4.5 cm. broad, narrowly long-acuminate, truncate or
shallowly cordate at the base, closely and evenly crenate, 5-nerved from the very
base, densely villous above with lax spreading hairs, long-villous beneath over the
whole surface; inflorescence cymose, the panicles 15-17 cm. long and as broad, the
branches densely villous, the pedicels 6-12 mm. long; heads small, numerous, white or
purplish white; involucre 5 mm. high, campanulate; phyllaries 2-3-seriate, the inner
ones linear-oblong, greenish, glabrous, ciliate, obtuse or rounded at the apex, the
outer phyllaries ovate or ovate-oblong, acute or subacute; flowers about 15, scarcely
exceeding the involucre, the corolla glabrous, 2 mm. long; achenes blackish, 1.3 mm.
long, minutely scaberulous or almost glabrous; pappus white, 2 mm. long.
A robust member of a group of species near E. microstemon
Cass.
Eupatorium araliaefolium Less. Linnaea 6: 403. 1831. E.
araliaefolium f. heterolepis Robinson ex Donn.-Sm. Enum. PI.
Guat. 6: 22. 1903, nomen. E. heterolepis Robinson, Proc. Am. Acad.
35: 335. 1900 (type from Alta Verapaz, S. Watson 65). Neomirandea
araliaefolia King & Robinson, Phytologia 19: 307. 1970. Matapalo
(San Marcos).
Moist or wet, mixed forest, 2,000 m. or lower; Peten; Izabal;
Alta Verapaz; El Progreso; Suchitepequez; Quezaltenango; San
Marcos. Southern Mexico; British Honduras; Honduras; Nicaragua;
Costa Rica.
A large shrub or a small tree, sometimes erect and 10 m. high, usually lower,
frequently an epiphytic shrub, pendent or with stems attached to the supporting tree
trunk, glabrous throughout except in the inflorescence, the stout branches somewhat
hexagonous or subterete; leaves opposite, on long slender petioles, fleshy when fresh,
usually black and very opaque when dried, oblong-ovate to lance-oblong, mostly 11-
20 cm. long, acute to long-acuminate, subacute to very obtuse or almost rounded at
the base, penninerved, entire, the nerves very inconspicuous; heads usually very
numerous, white, 6-9 mm. long, slender-pedicellate, about 25-flowered, forming small
or usually large, corymbiform, almost naked, dense or open panicles or corymbs;
inner phyllaries equal, linear, obtuse or acute, glabrous, caducous, the outer
phyllaries several, very short, persistent, ovate or lanceolate, obtuse or acute;
branches of the inflorescence usually brownish tomentulose; achenes slender,
glabrous; pappus yellowish white.
58 FIELDIANA: BOTANY, VOLUME 24
This species is said to have been introduced into cultivation in
European hothouses. It is the type species of the generic name
Neomirandea.
Eupatorium areolare DC. Prodr. 5: 169. 1836. E. tubiflorum
Benth. PL Hartweg. 76. 1841 (type from Zunil, Quezaltenango,
Hartweg 534). E. brevisetum DC. Prodr. 5: 169. 1836. Piptothrix
areolare King & Robinson, Phytologia 19: 169. 1970. Flor de San
Diego (Chimaltenango).
Moist or dry, often rocky, brushy hillsides, or probably most
often open, pine-oak forest, 800-2,700 m., mostly at the higher
elevations; Alta Verapaz; Baja Verapaz; Zacapa; Jalapa; Guate-
mala; Sacatepequez; Chimaltenango; Solola; El Quiche; Huehue-
tenango; Quezaltenango. Southern Mexico.
A slender shrub 1.5-4.5 m. high, the branches often recurved or arching, brown,
terete, villosulous on the stems, lower leaf surface, and inflorescence with usually
purplish, multicellular hairs; leaves opposite, short-petiolate, rather thick, ovate to
ovate-oblong or ovate-lanceolate, mostly 6-13 cm. long and 3-6 cm. broad, narrowly
long-acuminate, rounded or shallowly cordate at the base, rather rough on the upper
surface, 5-nerved, crenate-serrate; heads 12-20-flowered, white, 7 mm. long,
pedicellate, the pedicels densely pubescent; phyllaries rather few, unequal, linear-
oblong, acute, pubescent, rather lax; inflorescence corymbose-paniculate, very leafy,
generally dense and rounded; corollas with a long tubular throat, much exceeding the
involucre; achenes glabrous or nearly so.
A common and characteristic plant of rather dry, oak or pine
forest in the central mountains.
Eupatorium aschenbornianum Schauer, Linnaea 19: 720.
1847. E. donnell-smithii Coulter, Bot. Gaz. 16: 95. 1891 (type from
Aceituna, Guatemala, J. D. Smith 2374). E. donnell-smithii var.
parvifolium Donn.-Sm. op. cit. 96 (type from Duenas, Sacatepequez,
J. D. Smith 2333). Ageratina aschenborniana King & Robinson,
Pytologia 19: 212. 1970. Aromito bianco (Guatemala); milimento
(Alta Verapaz, perhaps an erroneous name); sakilocuj (Coban,
Quecchi).
Wet to dry thickets, usually on mountain slopes, or commonly
in pine-oak forest, sometimes in Cupressus forest or on white sand
slopes, rarely a weed in cultivated ground, 900-3,300 m., usually at
1,300 m. or higher, more common at the higher elevations; Alta
Verapaz; Baja Verapaz; Santa Rosa; Guatemala; Sacatepequez;
Suchitepequez; Chimaltenango; El Quiche; Huehuetenango;
WILLIAMS: FLORA OF GUATEMALA 59
Totonicapan; Quezaltenango; San Marcos. Central and southern
Mexico; El Salvador; Honduras; Nicaragua; Costa Rica; Panama.
An erect herb 1 m. high or less, rarely suffrutescent below, sparsely or much
branched, copiously villous almost throughout, with spreading, usually purplish,
several-celled hairs, the stems terete; leaves membranaceous, on slender, long or short
petioles, ovate to rounded-ovate, mostly 3-8 cm. long and 2.5-6.5 cm. broad,
acuminate or acute, rounded or more or less cordate at the base, crenate with usually
numerous close teeth, usually densely and softly short-villous on both surfaces, or the
pubescence rarely almost confined to the veins; heads 20-40-flowered, white,
pedicellate, about 5 mm. long, usually densely crowded in small, rounded or flat-
topped corymbs, these forming a small or large, compound, leafy corymb; phyllaries
subequal, linear or oblong-linear, pale, subscarious, lacerate-ciliate about the apex,
puberulent or glabrate; corollas pubescent about the lower part of the limb; achenes
small, blackish, usually glabrous.
This is one the very common Eupatorium species of the
Guatemalan mountains, abundant and rather conspicuous when
there is a moderate amount of moisture. It is one of the
characteristic species of the pine-oak forests. Hemsley reported from
Guatemala on the basis of a collection by Salvin and Godman of E.
bustamenta DC., a plant described from Mexico. A type photograph
of that species indicates that it is the same as, or close to E.
aschenbornianum.
Eupatorium bartlettii Robinson, Contr. Gray Herb. 100: 11.
1932. Critonia bartlettii King & Robinson, Phytologia 22: 48. 1971.
In advanced forest, often or perhaps usually on limestone, 350
m. or less; Alta Verapaz (Cubilguitz). British Honduras (type from
San Antonio, Bartlett 13068).
A glabrous shrub, the stems rather slender, flexuous, sometimes as much as 15 m.
long, scandent; leaves opposite, slender-petiolate, subchartaceous, lustrous, oblong to
ovate-oblong, 8-11 cm. long or shorter, acuminate, often abruptly so, obtuse at the
base, entire or undulate-crenate, triplinerved from a point well above the base; heads
5-flowered, sessile in small glomerules, broadly cylindric, the glomerules forming
small, leafy panicles; phyllaries pale green, obtuse, imbricate in 4-5 series, glabrous,
striate dorsally, puberulent near the apex; corollas white; achenes glabrous.
Eupatorium bellidifolium Benth. PL Hartw. 43. 1840. Agera-
tina bellidifolia King & Robinson, Phytologia 19: 212. 1970.
Moist oak or pine forests, alt. about 2,500 m.; Zacapa; Solola;
El Quiche. Mexico.
Erect or ascending herbs to 50 cm. tall, the stems short with the leaves borne at
the base, crisped pubescent, the peduncle elongate, sparsely pubescent, leafless or
nearly so; leaves ovoid or suborbicular, crenate, somewhat acute to rounded at the
60 FIELDIANA: BOTANY, VOLUME 24
base, sparsely pubescent on both surfaces (more so below) with crisped multicellular
hairs to glabrate, with 3 principal nerves arising from the apex of the petiole, blade
1.5-4 cm. long and nearly as broad, petiole 0.5-2 cm. long; inflorescence a few-headed
corymbose-panicle or panicle, the pedicels long and glandular pubescent; heads
campanula te, to about 8 mm. high; phyllaries about equal and in 1-2 series, linear or
linear-lanceolate, acute, glandular-pubescent, about 5-6 mm. long; achenes prismatic,
5-ridged, puberulent, black, about 2 mm. long; pappus bristles barbellate, about 3
mm. long; corolla white, about 3-3.5 mm. long.
Easily distinguished from its allies, E. muelleri and E.
anchisteum, by the glandular pedicels and by the leaves in a rosette.
Eupatorium blakei Rob. Contr. Gray Herb. 61: 5. 1920.
On rocks along the borders of streams or on wet banks, 1,000 m.
or less; Zacapa (Sierra de las Minas). British Honduras; Honduras
(type from Rio Molja, Dept. Copan).
An erect or decumbent, perennial herb, 80 cm. high or less, the stems usually
several, simple or sparsely branched, terete, puberulent; leaves membranaceous, on
short, slender petioles, lanceolate, elliptic or linear-lanceolate, 2-8 cm. long, 7-15 mm.
broad, long-attenuate to each end, 3-nerved from the base, remotely serrate,
puberulent on the nerves and veins but otherwise glabrous or nearly so; cymes
compound, terminating the branches, lax and open, forming a small leafy-bracted
panicle, the pedicels very short; heads about 20-flowered, campanulate, 4-5 mm. high;
phyllaries greenish, about 3-seriate, the outermost ovate or ovate-lanceolate, usually
subacute, puberulent, the inner ones narrowly elliptic-oblong, rounded at the apex;
corollas 2.5 mm. long, glabrous; achenes glabrous, 1 mm. long, fuscous, sparsely
hispidulous; pappus white.
Eupatorium caeciliae Rob. Contr. Gray Herb. 90: 23. 1930
(type from mountains between Totonicapan and los Encuentros,
Seler & Seler 2361). E. vetularum Standl. & Steyerm. Field Mus.
Bot. 23: 190. 1944 (type from mountains above Totonicapan,
Standley 84404). Ageratina caeciliae King & Robinson, Phytologia
19: 220. 1970.
Moist or wet thickets or forests, often abundant in the montane
cloud forests; Chimaltenango; Solola; Totonicapan; Huehue-
tenango. Endemic.
A branched shrub or weak tree, 2-5 m. tall, the branches stout, terete, striate,
brown, when young densely villous-tomentose or hispidulous-tomentulose with
brown, several-celled, simple or gland-tipped hairs; leaves opposite, subcoriaceous
when dry, on stout petioles 10-18 mm. long, elliptic-oblong or ovate-oblong, 5-11 cm.
long, 2.5-4.5 cm. broad, acute at the apex, acute and often oblique at the base,
penninerved, coarsely serrate or dentate, green above, puberulent or sparsely short -
villosulous to glabrate, somewhat rough to the touch, the nerves and veins
inconspicuous, paler beneath, everywhere sparsely villosulous or sordid-tomentulose,
stipitate-glandular on the costa, epunctate or obscurely and very minutely
WILLIAMS: FLORA OF GUATEMALA 61
puncticulate; inflorescence corymbose, dense and with very numerous heads, convex,
8-13 cm. broad, the branches fastigiate, densely villosulous and glandular-pilose, the
bracts and bractlets conspicuous, numerous, linear or subulate, the pedicels short or
elongate, straight, stout; heads 9-12 mm. long, 8-15-flowered, pale pink, turbinate-
campanulate; involucre about 7 mm. long, scarcely half as long as the florets;
phyllaries subequal or slightly unequal, obscurely 3-seriate, lance-linear, subap-
pressed, attenuate-acute, densely stipitate-glandular and minutely villosulous;
corollas narrowly tubular, glabrous, 7 mm. long, about equaling the pappus; achenes
slender, fuscous to blackish, 3-4 mm. long, densely glandular-atomiferous; pappus
bristles about equaling the corolla, pale purplish.
Eupatorium campechense Rob. Proc. Am. Acad. 43: 30. 1907.
Critonia campechensis King & Robinson, Phytologia 22: 48. 1971.
Thickets in the humid forest, Ruins of Tikal, Peten (Molina
15794). British Honduras; Mexico.
Erect or decumbent, perennial herbs or subshrubs, perhaps 1 m. tall, stems
smooth or obscurely angulate or striate, stramineous, glabrous, nodes 2-4 cm. long;
leaves opposite, ovate-lanceolate, acuminate, entire or obscurely dentate, triplinerved
from near the base, glabrous, blades mostly 6-10 cm. long and 3-5 cm. broad, the
petiole slender, 0.5-1.5 cm. long; inflorescence terminal, a diffused, flat-topped
panicle, at an thesis about 10 cm. broad; involucre cylindric, 7-10 mm. long and 2-3
mm. broad, glabrous; phyllaries in 4-5 series with the lower ones quite short, the
middle and upper oblong to linear- oblong, obtuse or acute, stramineous, glabrous,
except the tips ciliolate, the mid-nerved prominent and with 2 less prominent lateral
nerves, the shorter phyllaries about 1.5 mm. long, the inner longer ones to about 8
mm. long; heads mostly 5-flowered, 10-12 mm. long; corolla tubular, but slightly
expanded upward, glabrous outside, about 5 mm. long, the lobes very small, narrowly
triangular, about 0.3 mm. long; achenes prismatic, (4-)5-angled, pubescent, black at
maturity, about 3 mm. long; pappus uniseriate, as long as the corolla, with about 30
obscurely barbellate bristles.
Allied to Eupatorium bartlettii Rob., a species also found in the
lower elevations of Guatemala. E. bartlettii is easily distinguished
by the longer, narrower phyllaries, the larger leaves and the more
open panicles.
EUPATORIUM CAPILLIFOLIUM (Lam.) Small.
Plants of this or possibly some other closely related species are,
or have been, growing in the park at Puerto Barrios (Izabal). No
species of this alliance is known from continental tropical America,
and one wonders how the plants have reached here, especially since
they are neither showy nor attractive. The plant has filiform leaves
and very small heads of white flowers.
Eupatorium capillipes Benth. in Oersted, Vid. Medd. Kjoe-
benhavn 1852: 79. 1853; L. Wms. Fieldiana, Bot. 36: 84. 1975.
62 FIELDIANA: BOTANY, VOLUME 24
Ageratina helenae King & Robinson, Phytologia 24: 90. 1972. A.
molinae King & Robinson, I.e. 93.
Moist or wet thickets or dense mixed forest, often on banks
along mountain streams or beside waterfalls, 900-2,850 m., chiefly at
the higher elevations; Santa Rosa; Guatemala; Chimaltenango;
Suchitepequez: Quezaltenango; San Marcos. Honduras; Nicaragua;
Costa Rica.
A slender, erect herb, sometimes 1 m. high but generally half as tall or lower,
probably annual, usually much branched, the stems subterete, thinly puberulent or
almost glabrous; leaves opposite, small, membranaceous, on long, very slender
petioles, broadly ovate-triangular or ovate-rhombic, 3-5-plinerved from the base,
mostly 2-6.5 cm. long, acute, truncate to broadly cuneate at the base, very coarsely
crenate, with a few scattered thick hairs on the upper surface or glabrous, puberulent
or villosulous beneath on the nerves and veins or almost glabrous; heads very
numerous, 3.5-5 mm. long, with few to numerous flowers, white, mostly on long
filiform pedicels, these puberulent; phyllaries pale green, linear, equal or subequal,
attenuate-acute, glabrous; achenes small, black, scaberulous on the angles; pappus
copious, soft, white.
Apparently quite abundant in Guatemala, perhaps the impres-
sion of abundance is due to intensive collecting. It is occasional in
the mountains from Guatemala to Costa Rica and should be found
in adjacent Mexico.
Ku pa tori urn carletonii Rob. Contr. Gray Herb. 73: 7. 1924.
Fleishmannia carletonii King & Robinson, Phytologia 19: 203. 1970.
In crevices of rocks or cliffs along streams or beside waterfalls,
250-1,500 m.; Zacapa (Sierra de las Minas). Atlantic coast of
Honduras, the type collected near Cuyamel.
A slender, perennial herb, the stems numerous, forming dense clumps, 20-30 cm.
high, simple or sparsely branched, hexagonal, glabrous or sparsely puberulent, often
purplish; leaves opposite, short-petiolate, palmately trifid, 3.5-6 cm. long, the lobes
bifid, trifid, or pinnately dissected, the segments linear, 1-1.5 mm. broad, obtuse;
inflorescence terminal, fastigiately branched, cymose, 3-6 cm. broad, the pedicels 2-7
mm. long, slender; heads few or rather numerous, lilac, 4-5 mm. high, about 20-
flowered; involucre campanulate; phyllaries laxly imbricate, minutely puberulent, the
outer 2-4 much shorter than the others; achenes blackish, 1.2 mm. long, glabrate, the
pappus bristles white, 1.5-2 mm. long.
A very distinct species, easily recognized by its small dissected
leaves.
Eupatorium carmonis Standl. & Steyerm. Field Mus. Bot. 22:
303. 1940; L. Wms. Fieldiana, Bot. 36: 85. 1975.
Dry mixed forests at end of rainy season, 1,600-2,500 m.;
WILLIAMS: FLORA OF GUATEMALA 63
Chimaltenango; Sacatepequez (type from Finca Carmona, Standley
63734).
A perennial herb almost 1 m. high, the stems simple, slender, terete, almost
glabrous, when young minutely puberulent; leaves on slender petioles 3-4 cm. long,
thin-membranaceous, ovate to rhombic-ovate or lance-ovate, mostly 8.5-12.5 cm. long
and 4.5-5.5 cm. broad, acuminate, abruptly contracted near the base and long-
decurrent on the petiole, closely and evenly serrate, almost glabrous on both surfaces,
but sparsely scaberulous with minute, whitish hairs, especially on the veins, 5-
plinerved well above the base; inflorescence corymbiform, the corymb about 3 cm.
high and 5 cm. broad, much shorter than the leaves; heads small, white, numerous,
densely crowded, sessile or short-pedicellate, the pedicels densely sordid-puberulent;
involucre 3-3.5 mm. high, campanulate; phyllaries subequal, 2-seriate, greenish, acute,
pale-marginate, sparsely puberulent or pilosulous; corollas 2.2 mm. long, glabrous;
achenes glabrous; pappus white, 2.5 mm. long.
Eupatorium chiapense Rob. Proc. Am. Acad. 35: 332. 1900.
Moist or wet, mixed forest, 1,300-3,700 m.; Huehuetenango; San
Marcos. Southern Mexico, the type from Pinabete, Chiapas.
A shrub of 1.5-2.5 m., the branches very stout, terete, densely covered with a
rather coarse, purplish or brown tomentum; leaves opposite, on long, stout petioles,
membranaceous, oblong-ovate to very broadly ovate, 6-16 cm. long, 5-10 cm. broad,
acute or short-acuminate, very obtuse to subcordate at the base, rather closely and
inconspicuously serrate or crenate, penninerved or rather several-plinerved, the
principal nerves several, arising far above the base, very densely and softly pubescent
on both surfaces, more or less tomentose beneath; inflorescence large, corymbose,
convex, 6-20 cm. broad, densely tomentose and stipitate-glandular; heads 10-24-
flowered, about 9 mm. long; phyllaries linear, attenuate, subequal, laxly imbricate,
purplish, densely stipitate-glandular; corollas dull white; achenes slender, glabrous;
pappus soft, dirty white.
Eupatorium collinum DC. Prodr. 5: 164. 1836; L. Wms.
Fieldiana, Bot. 36: 85. 1975. E. neaeanum DC. I.e. 160. Krystenia
collina Greene, Leaflets 1: 9. 1903. Chromolaena collina King &
Robinson, Phytologia 20: 208. 1970. Barriquete (San Marcos).
Figure 16.
Moist or dry, brushy, often rocky hillsides, frequently in oak or
pine forest, 450-2,000 m.; Alta Verapaz; Baja Verapaz; El Progreso;
Izabal; Zacapa; Jutiapa; Santa Rosa; Escuintla; Guatemala;
Sacatepequez; Chimaltenango; Suchitepequez; El Quiche; Huehue-
tenango; Quezaltenango; San Marcos, Mexico; Honduras and El
Salvador to Costa Rica.
A rather brittle shrub or small tree, 1-6 m. high, usually much branched, the
branches slender, terete, densely puberulent or glabrate; leaves thin, grayish, on long
slender petioles, deltoid-ovate, often broadly so, 5-15 cm. long, narrowly long-
acuminate, cuneate to usually rounded or subcordate at the base, crenate, serrate, or
64 FIELDIANA: BOTANY, VOLUME 24
almost entire, 3-nerved, puberulent or pubescent on both surfaces or in one form
(Guatemala) almost glabrous; inflorescence corymbiform, almost flat to rounded;
heads very numerous, pedicellate, about 8 mm. long, 24-46-flowered, whitish or dirty
white, somewhat fastigiately aggregate; phyllaries acute or obtuse, graduate, several-
seriate, densely pubescent or tomentulose; achenes blackish or grayish, puberulent or
scaberulous, at least on the angles.
This is a common shrub, and a variable one, widely distributed
in Mexico and south through Guatemala, El Salvador, and
Honduras. A variation, which seems to be undescribed, occurs in
Nicaragua and Costa Rica. The heads of the variety are smaller
with fewer florets and the phyllaries are almost as long as the
florets. Called "vara de cama" and "vara blanca" in El Salvador,
where the stems are used to make the bottoms of beds. Probably
they are employed in the same manner in Guatemala. Known in
Honduras as "candelilla," "crucita," "majitero," "tatascan Colo-
rado," and "vara negra."
Eupatorium coulter! Rob. Proc. Am. Acad. 36: 477. 1901. E.
ageratifolium var. purpureum Coulter, Bot. Gaz. 16: 98. 1891 (based
on the same collection as E. coulteri}. Koanophyllon coulteri King
& Robinson, Phytologia 22: 149. 1971. Sanjoncillo (fide Aguilar).
Moist or wet, mixed forests or thickets, 1,200 m. or sometimes
more; Izabal; Alta Verapaz (type from Coban, Tuerckheim 52);
Baja Verapaz; El Progreso; Zacapa; Guatemala; Suchitepequez. El
Salvador; Honduras.
A slender shrub to 3 m. tall or less, the stems terete, when young densely
puberulent or villosulous, often purplish; leaves opposite, on rather short, slender
petioles, thick-membranaceous, deltoid-ovate, mostly 3-5 cm. long, acute or
acuminate, subtruncate at the base, usually subhastate near the base, shallowly
dentate, somewhat rough and scaberulous on both surfaces, minutely punctate,
slightly pubescent on the nerves; heads pale greenish yellow, about 8-flowered, 5 mm.
long, on filiform pedicels, in rounded, axillary and terminal, thyrsoid panicles;
involucre turbinate; phyllaries subequal or the outermost very short, the principal
ones lance-oblong, acuminate, erose, puberulent or granular, striate, often tinged with
purple; corolla glabrous, slightly exceeding the pappus; achenes fuscous, hispidulous
on the angles; pappus pale yellowish.
A very common shrub in the vicinity of Coban.
Eupatorium crassirameum Rob. Proc. Am. Acad. 35: 332.
1900; L. Wms., Fieldiana Bot. 36: 85. 1975. Ageratina crassiramea
King & Robinson, Phytologia 19: 228. 1970. Pachythamnus
crassirameus King & Robinson, Phytologia 23: 154. 1972.
Moist or wet, mixed forest, 200-2,500 m.; Guatemala; Huehue-
tenango; Quezaltenango. Southern Mexico; El Salvador; Nicaragua.
WILLIAMS: FLORA OF GUATEMALA 65
A shrub of 2-5 m., terrestrial or frequently epiphytic, the branches very thick,
fleshy, brittle, leafless at anthesis, grayish or brownish; leaves opposite, petiolate,
somewhat succulent, broadly ovate or ovate-deltoid, generally 10-15 cm. long and 8-
14 cm. broad, acute or acuminate, rounded or broadly cuneate at the base, subhastate
and with few sinuate teeth or merely sinuate-angulate or coarsely sinuate-dentate,
glabrous, 5-plinerved well above the base; heads about 15-flowered, 8 mm. long, white
or purplish, forming very dense, rounded, much branched corymbs at the ends of the
branches, pedicellate, the branches of the inflorescence thinly tomentulose or almost
glabrous; phyllaries subequal, linear-oblong, subacute, glabrous, the outermost few
and very short; achenes glabrous or nearly so; pappus yellowish white.
There is some question whether or not the material that we
have placed here is all one species; it is inadequate and the only
specimen from Guatemala with leaves (no flowers) was taken at
2,500-3,000 m., which makes it suspect. At the other end of the
ecological range we have collected E. crassirameum in Nicaragua on
a dry [sun] hot lava flow. All Central American material with thick,
fleshy, brittle stems has been placed here. It seems reasonable to
suspect that more than one species is involved.
Eupatorium crocodilium Standl. & Steyerm. Field Mus. Bot.
23: 182. 1944. Fleishmannia crocodilia King & Robinson,
Phytologia 19: 203. 1970.
Open swamps or wet thickets, 300 m. or less; endemic;
Huehuetenango (type from Cienago de Lagartero, below Miramar,
Steyermark 51498); Izabal.
An erect, perennial herb, 90 cm. high or less, the stems solitary or several, simple
or branched, sparsely and minutely puberulent above or glabrate, glabrous below and
abundantly leafy; leaves opposite, often with clusters of small leaves in their axils, on
petioles 1 cm. long or less, firm- membra naceous, rhombic-ovate or oblong-ovate, 2-3
cm. long, 1-1.5 cm. broad, narrowed to the obtuse apex, acute or broadly cuneate at
the base, 3-nerved, remotely and obscurely crenate, glabrous; inflorescence small,
fastigiate-trichotomous, terminal, long-pedunculate, the heads cymose-fastigiate, the
cymules 1-3, dense, 1.5-2 cm. broad; heads campanulate, lilac, 5 mm. long, about 15-
flowered, the pedicels as much as 4 mm. long, stout, minutely puberulent or
strigillose; phyllaries about 3-seriate, purplish or green, the inner ones linear-oblong,
rounded or very obtuse at the apex and minutely apiculate, minutely puberulent or
glabrate, ciliate, the outer ones very short, oblong or ovate-oblong, obtuse or
subacute; corollas glabrous, 2.3 mm. long; achenes glabrous, black, slightly more than
1 mm. long; pappus bristles numerous, scaberulous, white, 2.5 mm. long.
Eupatorium cupressorum Standl. & Steyerm. Field Mus.
Bot. 23: 183. 1944; L. Wms. Fieldiana, Bot. 36: 86. 1975. Ageratina
cupressorum King & Robinson, Phytologia 24: 89. 1972, as A.
cupressora.
66 FIELDIANA: BOTANY, VOLUME 24
Moist mixed forest or in Cupressus forest, 1,600-3,000 m.,
mostly at about 2,000 m.; Baja Verapaz; Chimaltenango; El
Quiche; Totonicapan (type between San Francisco El Alto and
Momostenango, Standley 84001).
An arching shrub, the branches terete, striate, brownish, rather densely villous-
tomentulose; leaves opposite, short- petio late, firm-membranaceous, on thickened,
short-villous petioles 12 mm. long or shorter, ovate to oblong-ovate or lance-oblong,
the larger ones 16 cm. long and 11 cm. broad, the upper ones narrower, narrowly
long-acuminate, shallowly cordate or broadly rounded at the base, triplinerved,
remotely serrate-dentate or repand-denticulate, sparsely villosulous or puberulent
above, rough to the touch, the veinlets prominent and minutely reticulate, slightly
paler beneath, villosulous on the nerves and veins, the nerves and veins very
prominent, the veinlets prominent and very finely articulate; inflorescence thyrsoid,
mostly axillary at the apex of the leafy stem, about 10-25 cm. long and 5-10 cm.
broad, the cymules few-headed, racemose, the stout pedicels 4-8 mm. long, densely
tomentulose, the bracts and bractlets mostly linear or subulate, often recurved; heads
rather large, numerous, white, about 8 mm. long with long-exserted styles at anthesis,
campanulate; phyllaries lance-linear, acute, densely pubescent, striate, pale, about
2.5-3 mm. long, subequal; flowers about 20, the corolla glabrous, twice as long as the
involucre; achenes 2 mm. long (immature), hispidulous on the angles; pappus bristles
stiff, yellowish white, slightly shorter than the corolla.
Eupatorium daleoides (DC.) Hemsl. Biol. Cent. Am. Bot. 2:
94. 1881. Critonia daleoides DC. Prodr. 5: 141. 1836. Copalillo.
Moist or wet forest, usually in pine forest, 1,450 m. or less; Alta
Verapaz; Izabal; San Marcos. Mexico; El Salvador and Honduras to
Panama.
A shrub or tree, sometimes 9-12 m. high, the young branches densely short-pilose
or tomentulose with brownish hairs, soon glabrate; leaves opposite, on short, slender
petioles, usually drying blackish, narrowly lance-oblong to ovate-oblong, 10-20 cm.
long, acute or acuminate, cuneate at the base, penninerved, serrate, often lustrous
above, tomentulose when young but soon glabrate, conspicuously brown-tomentulose
beneath when young, glabrate in age, chartaceous, pellucid-punctate and lineate;
heads numerous, white, 5-6 mm. long, narrowly campanulate, about 5-flowered,
sessile and glomerate at the ends of the branchlets, forming very large, rounded,
leafy-bracted, terminal panicles; phyllaries stramineous, unequal and strongly
imbricate, subappressed, ovate to narrowly oblong, obtuse; corollas equaling or
somewhat longer than the involucre; achenes sparsely puberulent; pappus yellowish
white.
Known in Honduras by the names "tatascan" and "tapahorno."
Eupatorium galeottii Rob. Contr. Gray Herb. 68: 17. 1923.
Moist or wet, mixed, mountain forest, 500-1,500 m.; Santa
Rosa; Suchitepequez; Quezaltenango. Southern Mexico.
A shrub or tree, 2-8 m. high, the branches obscurely 6-angulate, sparsely
puberulent or glabrate; leaves opposite, thick-membranaceous, on petioles 1-2.5 cm.
WILLIAMS: FLORA OF GUATEMALA 67
long, narrowly elliptic-oblong or lance-oblong, mostly 11-17 cm. long and 4-6.5 cm.
broad, acuminate or long-acuminate, attenuate at the base and somewhat decurrent
on the petiole, penninerved, coarsely or obscurely serrate, opaque, puberulent above
on the nerves, sparsely pubescent or almost glabrous beneath, the veins prominent
and reticulate on both surfaces; panicles small or very large, often much branched
and rather lax, as much as 19 cm. long and 17 cm. broad, more or less leafy or almost
naked, appressed-puberulent; heads short-pedicellate, campanulate, 5 mm. long,
greenish or somewhat purplish, few-flowered; phyllaries about 13, very unequal,
imbricate, acute, greenish, the outer ones deltoid-ovate, puberulent, the inner ones
oblong or linear; corollas glabrous; achenes 2 mm. long, pale, granular; pappus
yellowish white.
Closely related to E. pittieri Klatt, and perhaps not different.
Eupatorium glaberrimum DC. Prodr. 5: 144. 1836; L. Wms.,
Fieldiana; Bot. 36: 86. 1975. E. oerstedianum Benth. in Oersted, Vid.
Medd. Kjoebenhavn 1852: 71. 1853. E. vernonioides Coulter, Bot.
Gaz. 20: 45. 1895. Eupatoriastrum opadoclinium Blake, J. Wash.
Acad. Sci. 28: 479. 1938. Chromolaena glaberrima King &
Robinson, Phytologia 20: 208. 1970. C. oerstediana King &
Robinson, I.e. C. opadoclinia King & Robinson, I.e. Eupatorium
opadoclinium McVaugh, Contr. U. Mich. Herb. 9: 388. 1972.
Moist or dry, often rocky, open or brushy hillsides, frequent in
oak-pine forests, 1,600 m. or less; reported from Peten; El Progreso;
Izabal; Chiquimula; Jalapa; Santa Rosa; Guatemala; Huehue-
tenango. Mexico; British Honduras south to Costa Rica.
A stiff erect shrub, usually 1-2 m. tall, sparsely branched, the branches terete,
striate, brownish, densely pubescent to glabrous or glabrate; leaves sessile or on very
short petioles, subcoriaceous, oblong to oblanceolate-oblong to lance-oblong, mostly
15-20 cm. long, acuminate or long acuminate, acute to narrowly subcordate at the
base, a ppressed- serrate, usually inconspicuously so, sometimes entire, penninerved,
glabrous above or nearly so, often lustrous, slightly paler beneath, often resin-dotted,
villosulous beneath, especially along the costa, or glabrate, the veins prominent and
reticulate below; inflorescence large, corymbiform, almost flat-topped, leafy-bracted;
heads very numerous, cylindric-campanulate, about 40-flowered, white or pinkish-
white, about 1 cm. long; phyllaries several-seriate, closely appressed, stramineous,
somewhat indurate, the apices rounded or obtuse and ciliate, puberulent or glabrate;
receptacle rarely paleaceous; achenes black, prismatic, glabrous or puberulent.
E. glaberrimum is one of the rather characteristic plants of dry
oak-pine forests of Guatemala and Central America. Called "lengua
de vaca" and "lengua de venado" in Honduras; "carga pino" in El
Salvador. The species is distributed from Mexico south to Costa
Rica. It is exceedingly abundant in the oak-pine forests of
Guatemala and Honduras. Variations within the species are rather
great but no way to separate them satisfactorily has been found.
68 FIELDIANA: BOTANY, VOLUME 24
The heavily pubescent phase is represented by the name E.
opadoclinium.
It is unfortunate that the well-known name, E. oerstedianum,
must be substituted by an older but less well known one.
Eupatorium griseum Coulter, Bot. Gaz. 20: 43. 1895.
Peteravenia grisea King & Robinson, Phytologia 21: 395. 1971.
Moist or dry thickets or open forest, chiefly in pine forest,
1,150-2,100 m.; Zacapa; Chiquimula; Jalapa; Santa Rosa (type from
Casillas, Heyde & Lux 4250). Honduras; Nicaragua.
A coarse, perennial herb, erect or ascending, 1 m. high or less, the stem simple or
sparsely branched, densely villous with short spreading hairs, densely glandular-pilose
above and especially in the inflorescence, terete; leaves opposite, usually with very
small, almost sessile leaves in the axils, on rather long, stout petioles, broadly ovate-
cordate, 4-12 cm. long acuminate or abruptly acute, usually conspicuously cordate at
the base, rather coarsely dentate, 3-nerved, densely short-pilose or puberulent above,
densely short-pilose or villosulous and glandular-pilosulous beneath; heads white,
broadly campanula te, 10-12 mm. high and almost equally broad, many -flowered,
pedicellate, forming small or large, leafy or naked, usually open corymbs; phyllaries
several-seriate, graduate, oval or oblong lineate, stramineous, obtuse or rounded at
the apex, viscid-puberulent, the outermost short, broadly ovate, densely glandular-
puberulent; achenes minutely pubescent on the angles; pappus white or yellowish
white.
Eupatorium hastiferum Standl. & Steyerm. Field Mus. Bot.
22: 303. 1940. Neobartlettia hastifera King & Robinson, Phytologia
21: 295. 1971. Bartlettina hastifera King & Robinson, Phytologia
22: 160. 1971.
Known only from the type, in wet forest, mountains east of
Tactic, Alta Verapaz, on the road to Tamahu, 1,500-1,650 m.,
Standley 71125.
A shrub of 2.5 m., the branches rather stout, somewhat 6-angulate, glabrous;
leaves on long, slender petioles, herbaceous, blackish when dried, the petioles 3-5 cm.
long, dilated at the base into a large, amplexicaul, herbaceous pseudostipule; leaf
blades deltoid to broadly ovate or narrowly triangular, 6.5-12 cm. long, 6-10 cm.
broad, narrowly attenuate- acuminate, broadly rounded or subcordate at the base, the
sides rounded near the base or usually angulate and acute, the blades thus more or
less hastate, 3-nerved, closely and minutely serrate, minutely puberulent above on
the nerves, almost wholly glabrous elsewhere; inflorescence terminal, corymbiform,
rounded, 10 cm. broad, sparsely leafy; heads very numerous, campanulate, about 20-
flowered, on long slender pedicels, clustered in lax rounded cymes, probably white;
involucre 6-7 mm. high; phyllaries unequal, about 3-seriate, the inner ones oblong,
rounded at the apex, almost glabrous, pale, 3-5-striate, the outermost very short,
broadly ovate or suborbicular, rounded or very obtuse at the apex; corollas almost
filiform, 5.5 mm. long, glabrous; achenes 2 mm. long, glabrous, blackish; pappus
white, 4 mm. long.
WILLIAMS: FLORA OF GUATEMALA 69
Eupatorium hebebotryum (DC.) Hemsl. Biol. Cent. Am. Bot.
2: 95. 1881; L. Wms. Fieldiana, Bot. 36: 87. 1975. Critonia
hebebotrya DC. Prodr. 5: 141. 1836.
At about 1,050 m.; Santa Rosa; reported from Suchitepequez.
Central and southern Mexico; El Salvador; Honduras; Costa Rica.
A shrub or small tree, sometimes 8 m. high, glabrous or nearly so, outside the
puberulent or tomentulose inflorescence, the branches subterete or somewhat
angulate, stramineous or brownish; leaves opposite, membra naceous, on rather short,
slender petioles, rhombic-ovate to lance-oblong, 10-20 cm. long, penninerved,
acuminate or acute, acute or attenuate at the base, crenate-serrate, pellucid-punctate
and lineolate; inflorescence large, usually as broad as long or broader, rounded, leafy-
bracted or naked, usually about 20 cm. broad; heads white, about 5-flowered, very
numerous, 8 mm. long, on short, slender pedicels but usually densely aggregate and
forming large rounded glomerules; phyllaries stramineous, glabrous, the inner ones
oblong, obtuse or rounded at the apex, the outer ones much shorter, laxly imbricate;
corollas much exceeding the involucre; achenes densely puberulent; pappus pale
yellowish.
Called "tameagua" in El Salvador.
E. hebebotryum belongs in a critical section of the genus and
studies may show that the name is incorrectly applied. It is closely
related to E. pinabetense.
Eupatorium heydeanum Rob. Proc. Am. Acad. 35: 335. 1900.
Amolinia heydeana King & Robinson, Phytologia 24: 266. 1972.
Dense, moist or wet, mixed, mountain forest, sometimes on
white sand slopes, 900-2,200 m.; Santa Rosa (type from Rio de las
Canas, Heyde & Lux 3427); Suchitepequez; Quezaltenango; San
Marcos. Mexico.
A shrub or tree, 3-9 m. high, the branches stout, subterete or often somewhat
compressed, brownish, villous-tomentulose with brownish hairs when young, in age
glabrate; leaves opposite, on slender petioles 4-9 cm. long, membranaceous, usually
blackish when dried, ovate to lance-ovate, as much as 18 cm. long and 7-10 cm.
broad, gradually or abruptly long-acuminate, rather abruptly and broadly cuneate at
the base, triplinerved well above the base, entire or nearly so, at first sordid-
tomentulose above on the nerves, glabrate in age, beneath usually densely brown-
tomentulose or finally glabrate; panicles rounded, with opposite branches, 8-12 cm.
broad, rather open, much shorter than the leaves, the branches densely brown-
tomentulose; heads about 22-flowered, 9-10 mm. long, few or numerous, on short or
elongate pedicels; phyllaries unequal, linear, acute, laxly imbricate, sordid-
tomentulose; corollas yellowish white, 5-6 mm. long; achenes slender, 5 mm. long,
hispidulous on the angles; pappus dirty white.
This species is the basis of the monotypic genus Amolinia,
named for Prof. Antonio Molina R., outstanding Central American
botanist.
70 FIELDIANA: BOTANY, VOLUME 24
Eupatorium huehuetecum Standl. & Steyerm. Fieldiana, Bot.
22: 304. 1940.
Moist thickets or forest, sometimes in Alnus forest on white
sand slopes or on steep, open banks, 1,200-2,800 m.; endemic;
Huehuetenango (type from Rio Pucal, Standley 65798); Quezal-
tenango.
An erect, perennial herb, 1 m. high or less, the stems simple or branched, terete,
brownish, sparsely and minutely puberulent or shortly villous-pilose; leaves
membranaceous, on long slender petioles, rhombic-ovate to oblong-ovate or lance-
ovate, 4.5-11 cm. long, 1.5-5 cm. broad, acute or acuminate, acute at the base and
often unequal, 5-plinerved well above the base, finely and evenly crenate-serrate,
pilose above with short, spreading or appressed hairs, rough to the touch, almost
glabrous beneath or copiously villous-pilose; heads small, white, numerous, about 30-
flowered, crowded in small, rounded cymes, the cymes forming a rounded, leafy
panicle 6-12 cm. long, the slender pedicels mostly 2-6 mm. long, puberulent; involucre
campanulate, 4 mm. long; phyllaries linear subequal, acute or obtuse, pale, sparsely
puberulent, ciliate at the apex; corollas glabrous; achenes 1.2 mm. long, glabrous;
pappus white, 2 mm. long.
Eupatorium hypodictyon Rob. Proc. Bost. Soc. Nat. Hist. 31:
250. 1904.
At 900-2,100 m. or perhaps even higher; endemic; Zacapa
(Sierra de las Minas); Huehuetenango (type from Nenton, E. W.
Nelson 3517).
A shrub of 3 m., the branches rather slender but stiff, terete, glabrous; leaves
opposite, on petioles 1-2 cm. long, chartaceous, broadly ovate, 5-9 cm. long, 4-6 cm.
broad, acuminate or acute, rounded or almost truncate at the base, obscurely
crenate-serrate or almost entire, rough-puberulent above, slightly paler beneath,
thinly pilosulous beneath and somewhat viscid, the veins prominent and closely
reticulate on both surfaces, the principal nerves 5, arising well above the base of the
blade; heads 1 cm. high, on long, stout, tomentose pedicels, probably white, forming
dense, rounded or flat corymbs, 5-10 cm. broad; phyllaries closely appressed, 3-4-
seriate, graduate, broad, obtuse, sordid-puberulent; flowers about 25 in each head, the
corollas much longer than the phyllaries, tubular; achenes black, glabrous, 3 mm.
long; pappus yellowish.
The single collection from Zacapa is in bud only. Its leaves are
slightly different in pubescence from those of the type and it is
possible that it represents a different species.
Eupatorium hypomalacum Robinson in Donn.-Sm. Bot. Gaz.
35: 4. 1903. Koanophyllon hypomalaca King & Robinson,
Phytologia 22: 150. 1971.
Known in Guatemala only from the type, Los Verdes,
WILLIAMS: FLORA OF GUATEMALA 71
Guatemala, 1,100 m., Heyde & Lux 6157. A presumed variety has
been described from the Canal Zone.
A shrub, the branches grayish, subterete, minutely appressed-pubescent; leaves
opposite, on stout petioles about 1.5 cm. long, lanceolate or narrowly lance-oblong,
12-19 cm. long, 2.5-4.5 cm. broad, narrowly acuminate, cuneately narrowed at the
base and decurrent on the petiole, entire or nearly so, penninerved, sparsely
puberulent above, slightly paler beneath, densely and softly pubescent, densely and
minutely resin-dotted; inflorescence large, much branched, thrysiform-pyramidal, 10-
17 cm. long; heads campanulate, 8-10-flowered, pedicellate, crowded, 5 mm. long;
phyllaries 10-12, very unequal, pale green, striate, sparsely pubescent, acute, the outer
ones lanceolate, the inner ones oblong; corollas white, glabrous, 2.4 mm. long;
achenes 2.2 mm. long, hispidulous on the angles; pappus pale yellowish.
Eupatorium sorensenii is a closely related species.
Eupatorium imitans Rob. Contr. Gray Herb. 68: 20. 1923.
Fleishmannia imitans King & Robinson, Phytologia 19: 203. 1970.
Llovizna (fide Aguilar).
Usually on rocks in or along the edges of streams, sometimes on
sandbars or frequently on wet, shaded banks or cliffs, along or near
streams, 200-1,950 m.; Peten; El Progreso; Zacapa; Chiquimula;
Jalapa; Jutiapa; Santa Rosa (type from Santa Rosa, Heyde & Lux
4194); Escuintla; Guatemala; Suchitepequez; Retalhuleu; El
Quiche; Huehuetenango. El Salvador; Honduras; Nicaragua; Costa
Rica.
An erect, perennial herb, usually 60 cm. high or less, the stems usually numerous
and often forming dense clumps, slender, simple or sparsely branched, puberulent or
tomentulose, soon glabrate, terete; leaves opposite, membranaceous, on petioles 7-12
mm. long, narrowly ovate or usually lanceolate to linear-lanceolate mostly 2.5-4.5 cm.
long, acute or attenuate at each end, 3-nerved, remotely serrate, puberulent,
especially on the veins, or glabrate; inflorescence terminal, pyramidal or rounded,
laxly branched, leafy, the heads arranged in small corymbs of few heads; heads about
50-flowered, 5-6 mm. long; involucre usually 3-5-seriate, campanulate; phyllaries
lanceolate or linear, subequal, green, acute, mostly bicostulate, puberulent; corollas
hispidulous toward the limb; achenes black, 1.3 mm. long, minutely hispidulous on
the angles; pappus white or yellowish white.
This species is somewhat intermediate between two groups of
Eupatorium that have been separated as Fleishmannia and
Ageratina by King and Robinson. The species is an interesting one,
since it may be found in flower in the dry season along the rivers in
some of the driest valleys of Central America.
Eupatorium incomptum DC. Prodr. 5: 173. 1836. Decachaeta
incompta King & Robinson, Brittonia 21: 280. 1969. Palpala
(Jalapa).
72 FIELDIANA: BOTANY, VOLUME 24
Moist or dry, brushy, often rocky slopes, in thickets or open
pine or oak forest, 900-1,800 m.; Zacapa; Chiquimula; Jalapa;
Jutiapa; Chimaltenango; Huehuetenango. Western and southern
Mexico.
An erect herb, 1 m. high or less, perennial, the stems slender or stout, simple or
sparsely branched, terete, tomentose or in age glabrate; leaves opposite or the upper
ones alternate, membranaceous, short-petiolate, ovate to rhombic-ovate or rhombic-
lanceolate, mostly 9-15 cm. long, acute to attenuate, gradually or abruptly narrowed
below and attenuate almost to the base of the petiole, coarsely crenate or dentate to
subentire, sometimes shallowly trilobate, 3-5-plinerved far above the base of the
blade, scabrous above and very rough to the touch, finely and softly pubescent
beneath, densely resin-dotted, the nerves and veins very prominent and closely
reticulate beneath, sometimes impressed on the upper surface; heads dirty white, 4
mm. long, 10-12-flowered, pedicellate, very numerous, forming a very dense or open,
large, thyrisiform, leafy or almost naked panicle; phyllaries subequal, narrowly
oblong, very obtuse or rounded at the apex, sometimes covered with yellow or
brownish glands; achenes glabrous.
See Eupatorium ovandense Grashoff & Beaman, which is
related to E. incomptum DC.
Eupatorium ivaefolium L. Syst. ed. 10. 1205. 1759. Chromo-
laena ivaefolia King & Robinson, Phytologia 20: 202. 1970.
Open, moist or dry places, 1,200 m. or less; reported from
Peten; Santa Rosa; Huehuetenango. Southern United States;
Mexico; Honduras; West Indies; widely distributed in South
America.
A stiff, erect, perennial herb, usually less than 1 m. high, the stems simple below,
branched above, with usually strongly ascending branches, harshly villous with
multicellular hairs; leaves opposite, narrowly lanceolate or lance-linear, 3-nerved, 3-7
cm. long, attenuate-acuminate, acute or attenuate at the base, herbaceous, the upper
ones entire, the lower coarsely dentate, very scabrous above, rough-villous beneath,
conspicuously resin-dotted; inflorescence usually large and much branched, rather
open, corymbose or broadly paniculate; heads usually very numerous, about 8 mm.
long, purple or lavender, short-cylindric, slender-pedicellate; phyllaries broad, closely
appressed, rounded or obtuse, the outer and middle phyllaries with short, green tips,
somewhat erose, the innermost phyllaries often petaloid and purplish at the apex;
achenes black, 2.5 mm. long, glabrous or with roughened angles.
The specific name was written by Linnaeus at various times as
iuaefolium, ivaefolium, and ivifolium.
Eupatorium je junum Standl. & Steyerm. Field Mus. Bot. 23:
183. 1944. Fleishmannia je juna King & Robinson, Phytologia 19:
203. 1970.
Moist or wet thickets or shaded banks, sometimes in rocky
places, 850-1,700 m.; endemic; Chiquimula; Jutiapa (type from El
WILLIAMS: FLORA OF GUATEMALA 73
Barrial, east of Jutiapa, Standley 75788); Santa Rosa; Quezal-
tenango.
A very slender, erect annual, 13-50 cm. high, simple or laxly and sparsely
branched, the stems terete, pale greenish, densely and minutely puberulent; leaves
small, thinly membranaceous, bright green, on slender petioles 1-2.5 cm. long, ovate,
rhombic-ovate, or deltoid-ovate, 2.5-4 cm. long, 2-4 cm. broad, acute or acuminate,
truncate or broadly rounded at the base, sometimes subcordate, 3-nerved, coarsely
and evenly crenate-dentate, very sparsely villosulous above, with short, spreading,
white hairs or almost glabrous, more or less puberulent beneath on the nerves and
veins; heads lilac, small, numerous, cymose, forming a large, very lax panicle, on
slender pedicels 1 cm. long or shorter, the pedicels puberulent, with linear bractlets;
involucre almost 3 mm. long, campanulate; phyllaries subequal, pale green, linear,
acute, costulate, glabrous; flowers about 40, equaling the phyllaries; corollas
glabrous, 1.5 mm. long; achenes slender, black, 1 mm. long, glabrous; pappus bristles
few, white, deciduous.
A small and rather delicate plant, probably to be found only
during the wetter months. In general appearance it is much like E.
sinclairii and E. microstemon, but the phyllaries are equal in
length, rather than graduate and imbricate as in those species.
Eupatorium laevigatum Lam. Encycl. 2: 408. 1786. Chromo-
laena laevigata King & Robinson, Phytologia 20: 202. 1970. Carga-
pino.
Usually in open, pine or oak forest, sometimes in savannas or
moist or wet meadows about lakes, 300-1,600 m.; Peten; Zacapa;
Chiquimula; Jalapa; Jutiapa. Southern Mexico; Honduras; El
Salvador; widely distributed in South America.
A shrub 1-1.5 m. high, glabrous throughout, very viscid in all parts, the branches
and pedicels angulate, the older branches brown; leaves opposite, short-petiolate,
subcoriaceous, rhombic-ovate to ovate-oblong, 8-13 cm. long, acute, cuneate at the
base, 3-nerved from the very base, usually coarsely serrate or crenate, often very
lustrous, with numerous, transverse, prominent veins beneath; inflorescence
corymbose, large and many-headed, convex or almost flat-topped; heads sessile or on
rather long, stiff pedicels, white or tinged with lilac, sweet-scented; involucre
cylindric, glutinous; phyllaries several-seriate, graduate, closely appressed, broad,
rounded at the apex, stramineous; achenes almost black, glabrous.
Called "azota-caballo" in Honduras; the name "carga-pino" is
given the species because pine needles lodge on the plant when they
fall from the overhanging trees.
It is curious that this species has not been found in southern
Central America or Panama. It is apparently widespread in South
America, skips to Honduras and occurs as far as Mexico, where it is
uncommon.
74 FIELDIANA: BOTANY, VOLUME 24
Eupatorium lanicaule Robinson, Proc. Am. Acad. 35: 336.
1900. Critonia lanicaulis King & Robinson, Phytologia 22: 49. 1971.
Figure 17.
Moist or wet thickets or forest edges, 500 m. or less; Alta
Verapaz (type, Watson 74); Izabal. British Honduras.
Shrubs to 3 m. tall and 5 cm. in diameter, the stems terete, sparsely branched,
villous with spreading hairs above, glabrate in age; leaves elliptic to oblong or
oblong-oblanceolate, acuminate, attenuate to an acute or narrowly obtuse base,
penninerved with 7-8 pairs of lateral nerves, tertiary nerves impressed, reticulate
above, prominently reticulate below, thinly appressed-pilose or glabrate except along
the principal nerves, minutely to rather coarsely serrulate, the blades mostly 15-30
cm. long and 5-8 cm. broad, the petioles very short, pilose, up to about 1 cm. long;
inflorescences usually small and condensed, sometimes openly branched, the cymules
short-pedunculate, the peduncle to 2 cm. long and mostly with few heads; heads
subsessile, 12-17 mm. long and 6-10 mm. broad; involucre about 5-seriate; outer
phyllaries striate, ovate, obtuse, ciliolate and sometimes pilosulous dorsally, about 2
mm. long, the innermost phyllaries lanceolate, obtuse, 8-10 mm. long, obscurely
ciliolate; florets about 20 or sometimes more, the corollas cylindric and somewhat
expanded upward, constricted at the throat, about 6-7 mm. long, white or flushed
with purple; achenes glabrous or puberulent; pappus about as long as, or slightly
longer than, the florets.
A distinctive species but rather uncommon in the wet lowlands
of the Atlantic slope.
Eupatorium leucocephalum Benth. PL Hartw. 86. 1841. E
mendax Standl. & Steyerm. Field Mus. Bot. 23: 185. 1944.
1,300 m. or higher; Alta Verapaz; Baja Verapaz; Santa Rosa;
Escuintla; Guatemala; Sacatepequez; Chimaltenango (type of E.
dolores; flor de dolores; noche buena; flor de sacramento; chilca;
fenferex q'en (Alta Verapaz, Quecchi).
Wet to dry thickets or mixed forest, 750-1,900 m. mostly at
1,300 m. or higher; Alta Verapaz; Baja Verapaz; Santa Rosa;
Escuintla; Guatemala; Sacatepequez; Chimaltenango (type of E.
leucocephalum from Acatenango— probably the volcano— Hartweg
588); Suchitepequez; El Quiche; Huehuetenango; Quezaltenango;
San Marcos (type of E. mendax, Volcan de Tacana, Steyermark
36341). Southern Mexico; El Salvador.
A slender shrub 1-1.5 m. high, sparsely branched, the branches suberect or
ascending, terete, glabrous or nearly so; leaves thin, bright green, on short slender
petioles, opposite, lanceolate or oblong- lanceolate, 6-10 cm. long, narrowly long-
acuminate, obtuse or cuneate at the base, triplinerved from well above the base,
coarsely serrate, often lustrous, glabrous above, usually sparsely pubescent beneath
along the nerves; heads small, mostly 4-5 mm. long, very numerous, forming a large,
ovoid, leafy-bracted, thyrsoid panicle, slender-pedicellate, the pedicels pubescent;
WILLIAMS: FLORA OF GUATEMALA 75
phyllaries ovate to oblong, about 3-seriate, unequal, white, lustrous, obtuse, not
closely appressed; flowers 6-8, white; achenes glabrous.
The plant is called "flor de plata" and "hierba de plata" in El
Salvador. It is common and well known in many parts of
Guatemala, easily recognized by its large and showy panicles of
very numerous, small, pearly white heads. It is a highly ornamental
plant, probably worthy of cultivation and is frequently planted in
Guatemalan gardens. In Guatemala E. leucocephalum is a favorite
for decorating churches, household altars, outdoor crosses, and
other objects because the branches retain their fresh appearance for
a long time, at least if placed in water. It is much used in Coban
during Lent and Holy Week.
See also Eupatorium macrwn, a species closely related to this
but with sparsely headed inflorescences and greenish phyllaries.
Eupatorium lucentifolium L. Wms. Fieldiana, Bot. 36: 88.
1975.
Forested slopes and canyons, 1,300-1,400 m. Alta Verapaz,
Standley 89769. Mexico (Chiapas on Guatemalan border, type
Breedlove 9765).
Shrubs or trees to 9 m. tall but usually less, stems slightly flattened at the
younger, somewhat immature nodes, obscurely puberulent with appressed hairs;
leaves penninerved, entire, subcoriaceous, somewhat bullate, lanceolate-ovate or
oblong-ovate, abruptly acuminate, somewhat acute or rounded at the base, shining,
glabrous except minutely puberulent on the midvein below, the blade 7-15 cm. long
and 2.5-6 cm. broad, tertiary nerves prominently reticulate, especially above, petiole
slender, appressed-puberulent, 1-2 cm. long; inflorescence a terminal, multicapitate,
obscurely puberulent, profuse, corymbose panicle, to 25 cm. long and as broad; heads
sessile or on pedicels not more than 1 mm. long, mostly about 6 mm. long and usually
with 5 florets; phyllaries 3-4-seriate with the outer ones narrowly ovate and the inner
ones oblong-ovate to linear-oblong, obtuse, the innermost about 5 mm. long and
about as long as the pappus; corolla pink or white, tubular, glabrous, about 4 mm.
long and 0.3 mm. in diameter, the lobes 0.2-0.3 mm. long and narrowly ovate; anthers
minute, about 0.6 mm. long, appendaged at the apex; style at anthesis exserted 3-4
mm. beyond the corolla, slightly thickened at the apex; achenes prismatic and 5-
ridged, glabrous, about 2 mm. long; pappus about 4 mm. long, of 30-40 bristles.
Closely related to Eupatorium nubigenum Benth., from which
it may be distinguished by the relatively broad, entire leaves, with
prominently reticulate tertiary nerves (not minutely papillate), and
by the appressed-puberulent stems and inflorescence.
Eupatorium luxii Rob. Proc. Am. Acad. 36: 480. 1901 (type,
Heyde & Lux 3387 from Quiche); L. Wms. Fieldiana, Bot. 36: 89.
1975. E. pansamalense Rob. I.e. 482 (type, Tuerckheim 1342 from
76 FIELDIANA: BOTANY, VOLUME 24
Alta Verapaz). E. oresbioides Rob. Proc. Am. Acad. 44: 618. 1909
(type, Salvin & Godman 326 from Sacatepequez). Neobartlettia
luxii King & Robinson, Phytologia 21: 296. 1971. N. pansamalensis
King & Robinson, I.e. Bartlettina luxii King & Robinson,
Phytologia 22: 161. 1971. B. pansamalensis King & Robinson, I.e. B.
oresbioides King & Robinson, I.e. B. breedlovei King & Robinson,
Phytologia 28: 286. 1974 (type from Chiapas, Breedlove 9075). B.
guatemalensis King & Robinson, I.e. 287 (type from Quiche, Skutch
1700). Ec-cul (Huehuetenango); monte negro (Huehuetenango); flor
celeste (Quezaltenango).
Moist or wet, mixed forests or thickets, coniferous forests or
occasionally on open slopes, 1,500-3,300 m., most common at 2,200-
2,600 m.; Alta Verapaz; Baja Verapaz; El Progreso; Jalapa;
Sacatepequez; Chimaltenango; Solola; El Quiche; Quezaltenango;
San Marcos; doubtless in all highland departments. Southern
Mexico; El Salvador.
Shrubs or small trees, 1-6 m. tall, the branches slender, sordid-puberulent or
tomentulose to almost glabrous; leaves elliptic-ovate to deltoid-ovate, 6-15 cm. long
and mostly 3-8 cm. broad, acuminate to caudate acuminate at the tip, acute or
attenuate to rounded or subtruncate at the base, serrate, penninerved or
subplinerved, puberulent or villosulous on the costa above, and below on the nerves,
glabrescent, slender petioles mostly 2-3 cm. long; inflorescence sordid tomentulose to
subglabrous, corymbiform, often broad, dense or quite open; heads slender, short
pedicellate, 8-10 mm. high; phyllaries very unequal, 3-4-seriate, the outer lanceolate
to lanceolate- ovate, acute, the inner phyllaries often elongate, lance-oblong, acute or
obtuse, puberulent and ciliolate to tomentulose or almost glabrous; corollas pink or
lilac, 4-6 mm. long, glabrous or obscurely pilose at the tips; the styles long exserted at
anthesis and purple; achenes black, glabrous, 1-2 mm. long but mostly 1.5 mm. long;
pappus white and about equalling the florets.
A handsome and showy shrub, often abundant in the hills of
the highlands. The species is somewhat variable, especially in the
phyllaries. These are often only half as long as the heads and
mostly lanceolate, but sometimes they are as long as the heads and
broader, with the inner phyllaries relatively narrow and obtuse. A
critical study of the group is needed.
Eupatorium macrophyllum L. Sp. PI. ed. 2. 2: 1175. 1763.
Hebeclinium macrophyllum DC. Prodr. 5: 136. 1836. Arepaxiu
(Peten).
Moist or wet thickets, usually in second growth, sometimes in
wet lowland forest, 350 m. or less; Peten; Alta Verapaz; Izabal;
Escuintla. Southern Mexico; British Honduras to Panama; South
America.
WILLIAMS: FLORA OF GUATEMALA 77
A coarse erect herb, 1-2 m. high, the stems subterete, villous-tomentose with
whitish or fulvescent hairs, the internodes often much elongate; leaves opposite, thin,
on long, slender petioles, very broadly deltoid-ovate, 10-20 cm. long or larger, often as
broad as long or broader, abruptly acute or acuminate, broadly and openly cordate at
the base, closely crenate, 3-nerved from the base, finely pubescent on both surfaces or
often tomentulose or velutinous beneath; panicles terminal, dense, leafy, broad, with
spreading branches; heads 50-75-flowered, about 7 mm. high, greenish white,
pedicellate; involucre campanulate, many-seriate, regularly graduate; phyllaries
lanceolate, acute or subacute, pale green with whitish costae; corollas slender-
tubular, sometimes tinged with lilac; achenes dark gray, the costa white, slightly
roughened near the apex.
E. macrophyllwn is a common, weedy plant in the Atlantic
lowlands of Central America. The very large and thin leaves are soft
and limp and droop unless the plants are abundantly supplied with
water. Called "oregano" in Veracruz.
Eupatorium macrum Standl. & Steyerm. Field Mus. Bot. 23:
184. 1944.
Rocky banks, 400 m.; Peten; Chiquimula (type, Jocotan,
Steyermark 31534). Honduras.
Plants herbaceous or lignescent, the branches slender, pale, terete, densely short-
pilose; leaves opposite, thin-membranaceous, on slender petioles 1-2.5 cm. long,
triangular-lanceolate or narrowly rhombic-lanceolate, 3.5-8.5 cm. long, 1.5-3.5 cm.
broad, narrowly attenuate-acuminate, with an obtuse tip, acute or attenuate at the
base and long-decurrent upon the petiole, inconspicuously undulate-dentate or
almost entire, triplinerved far above the base of the blade, green above, sparsely and
minutely puberulent, slightly paler beneath, rather densely crisped-pubescent;
inflorescence laxly paniculate, leafy, the internodes much elongated, the branches
dichotomous, very slender, the heads few, sessile in small clusters at the ends of the
branches and in the forks of the branches; heads cylindric, white, 6 mm. long, 5-
flowered; phyllaries unequal, about 3-seriate, stramineous or pale greenish, the inner
ones linear-oblong, rounded or obtuse at the apex and thin, obscurely costulate,
somewhat lustrous, the outer ones few, very short, oblong-ovate, acute, glabrous,
ciliolate; corollas glabrous, scarcely longer than the involucre; achenes blackish, 1.5
mm. long, densely scaberulous; pappus bristles white, scaberulous, almost equalling
the corolla.
The species is locally abundant in central Honduras. It is
closely related to E. leucocephalum.
Eupatorium magistri L. Wms. Fieldiana, Bot. 36: 90. 1975.
Probably a vine or a shrub in the wet forest or forest edge, at
about 1,000 m., Alta Verapaz. Known only from the type,
Tuerckheim II. 1912.
Shrubs or perhaps lianas, glabrous, the stems terete with opposite branches;
leaves ovate to elliptic-ovate, acute or short acuminate, rounded or obtuse at the
78 FIELDIANA: BOTANY, VOLUME 24
base, 5-plinerved with the lower pair of nerves less conspicuous, blade 3-7 cm. long
and 1.5-2.5 cm. broad, entire or nearly so, the petiole slender, 0.5-1.5 cm. long;
inflorescences terminal or axillary on the lateral branches, paniculate, each 2-6 cm.
long, with relatively few sessile or short pedicellate heads; heads about 8 mm. long
and 3-4 mm. broad, each with 10 florets; involucre of many phyllaries in 3-5-series;
outer phyllaries much shorter, from about 1 mm. long and broadly ovate, obscurely
ciliolate, the inner phyllaries longer, 5-7 mm. long, some becoming narrowly
lanceolate and acute; corolla about 4.5 mm. long and 0.3-0.4 mm. in diameter,
cylindric, the lobes lanceolate, about 0.5 mm. long; anthers slender, about 1.5 mm.
long, appendaged at the apex; style at anthesis somewhat thickened at the apex and
exserted about 2 mm. beyond the corolla; mature achenes glabrous, 5-ridged, about 2
mm. long; pappus about 4.5 mm. long, as long as the corolla, with 30-40 bristles,
minutely barbellate at the apex.
Named for Dr. Benjamin Lincoln Robinson, outstanding
student of the Compositae, friend of the senior author and teacher,
by example, of the junior author.
Eupatorium mairetianum DC. Prodr. 5: 167. 1836. E.
rafaelense Coulter, Bot. Gaz. 16: 97. 1891 (type from San Rafael,
Sacatepequez, Smith 2331). E. mairetianum var. adenopodum
Robinson, Proc. Am. Acad. 51: 534. 1916 (type from Cerro
Quemado, Quezaltenango, Holway 98). Ageratina mairentiana
King & Robinson, Phytologia 19: 224. 1970. A. rafaelensis King &
Robinson, I.e. 225. Sajoc (Quezaltenango).
Mostly in dense or open, pine-oak forest, sometimes in forest of
Cupressus or Abies, frequently on open, brushy hillsides, often in
rocky places, 1,200-3,300 m., chiefly at the higher elevations; Alta
Verapaz; Baja Verapaz; Santa Rosa; Guatemala; Sacatepequez;
Chimaltenango; Solola; El Quiche; Huehuetenango; Totonicapan;
Quezaltenango; San Marcos. Central and southern Mexico; El
Salvador.
A shrub of 1-4 m., much branched, the pubescence of the stems and inflorescence
usually arachnoid and appressed, the leaves also often arachnoid, especially beneath
in the axils of the nerves, or the young branches and inflorescence glandular-
puberulent, the branches terete, brown, with white pith; leaves opposite, membrana-
ceous, on long, slender petioles, lance-ovate to broadly rhombic-ovate, mostly 6-11
cm. long and 3-5 cm. broad, usually long-acuminate, rounded to broadly cuneate at
the base, serrate or almost entire, green above and glabrous or nearly so, paler
beneath and more or less pubescent, plinerved above the base; heads numerous, white
or pinkish, 12-16 mm. long, corymbose, the corymbs small and dense, generally
forming an ovoid, leafy-bracted, terminal thyrse; heads about 25-flowered; phyllaries
linear, acuminate, subequal, puberulent or glandular-puberulent, lax, the outermost
very short; achenes slender, black, puberulent on the angles.
Var. adenopodum is a form in which the pedicels are densely
glandular-puberulent, rather than tomentulose, as in the typical
WILLIAMS: FLORA OF GUATEMALA 79
phase of the species. There is much variation and intergradation in
this respect, even in the same region, and the variety is scarcely
worthy of special designation, it being difficult at times to decide to
which form a certain branch should be referred.
A common and often abundant species in the western
highlands at the beginning of the dry season in December and
January.
Eupatorium microstemon Cass. Diet. Sci. Nat. 25: 432. 1822.
E. guadalupense DC. Prodr. 5: 170. 1836.
Mixed forest at 2,200-2,800 m.; Baja Verapaz; Guatemala;
Chimaltenango; Solola; El Quiche; Quezaltenango; perhaps in other
departments of the western highlands. Mexico; El Salvador (?);
Costa Rica (?); West Indies; South America.
Plants herbaceous, probably annual, very slender, 1 m. high or less, usually much
branched, puberulent or glabrate; leaves opposite, slender -petiolate, membranaceous,
deltoid-ovate, 2.5-7 cm. long, acute or acuminate, with an often obtuse tip,
subtruncate or rounded at the base, 3-nerved, crenate-serrate, puberulent or glabrate;
heads 4-5 mm. high, white or lilac, laxly paniculate, on slender, elongate, minutely
puberulent pedicels, many-flowered; involucre campanulate; phyllaries 3-4 seriate,
pale greenish, costulate, the outer ones lanceolate, acute, the inner linear-oblong,
rounded and somewhat scarious at the apex; corollas glabrous, about equaling the
involucre; achenes slender, glabrous; pappus white, equaling the corolla.
The Maya name "xultoxiu" is recorded from Yucatan.
The "small-headed" Eupatoria are difficult to distinguish and
the author is not sure that our specimens are of this species. Dr.
Robinson, at the beginning of the century, determined material
from Yucatan that seems to be similar. However his was lowland
material, while our specimens are from the highlands. Our
description is based on Guatemalan materials.
There are some seven other species in Guatemala that are
closely related to this one and much in need of careful study.
Eupatorium mimicum Standl. & Steyerm. Field Mus. Bot. 23:
186. 1944. Koanophyllon mimica King & Robinson, Phytologia 22:
150. 1971.
Moist or wet thickets or mixed forest, 1,300-3,000 m.; known
only from Guatemala but doubtless extending into Chiapas;
Quezaltenango (type from region of Las Nubes, south of San
Martin Chile Verde, Standley 85146, collected also at several other
localities); San Marcos (volcanoes of Tacana and Tajumulco).
80 FIELDIANA: BOTANY, VOLUME 24
A slender erect shrub, 1-2.5 m. high, the branches very slender, terete, brownish
or greenish, densely villosulous or incurved-puberulent, with fulvescent or purplish
hairs; leaves opposite, membranaceous, on slender petioles 1.5-4.5 cm. long, deltoid,
rhombic-deltoid or ovate-deltoid, 3.5-8 cm. long, 2-5 cm. broad, acuminate or long-
acuminate, truncate or broadly rounded at the base, 3-nerved, coarsely and unequally
crenate or crenate-dentate, green above, sparsely villosulous, with stout, whitish,
several-celled hairs, tomentulose beneath on the nerves and usually sparsely
villosulous on the nerves and veins, minutely glandular-puncticulate; inflorescence
terminal, cymose- paniculate or subthyrsoid, leafy, scarcely more than 6 cm. long, the
bracts few, linear or almost subulate, the heads few or numerous, clustered in lax
cymes, the pedicels longer and slender, densely puberulent or tomentulose; heads
whitish or pale greenish yellow, 7-8 mm. long, about 8-flowered; involucre narrowly
campanulate, half as long as the florets; phyllaries subequal, about 2-seriate, laxly
imbricate, lance-linear, acute to long-acuminate, greenish, densely puberulent,
minutely glandular-puncticulate; corollas 4 mm. long, glabrous, slightly shorter than
the pappus; achenes black, slender, 3 mm. long, densely and minutely hispidulous;
pappus bristles numerous, pale yellowish, scaberulous.
Closely related to E. coulteri and perhaps only a variety of that
species, but the two have distinct ranges in Guatemala.
Eupatorium molinae L. Wms. Fieldiana, Bot. 36: 91. 1975.
Neomirandea ovandensis King & Robinson, Phytologia 19: 309.
1970, not Eupatorium ovandensis Grashoff & Beaman, 1969.
Epiphytic in the montane cloud forest at about 2,300 m.; San
Marcos (Williams et al. 26863). Mexico (type, Matuda 3917 from
Mt. Ovando, Chis.).
Suffrescent epiphytes, glabrous except in the inflorescence; leaves lanceolate or
lanceolate-ovate, acuminate, entire or obscurely repand, 7-10 cm. long and 3-4 cm.
broad, glabrous, the tertiary nerves forming prominent reticulations, the petioles
short, 1.5 cm. long or less; inflorescence terminal, crowded, paniculate, puberulent,
about 5-6 cm. broad; heads 10-14 mm. long, with about 10 flowers; phyllaries
subequal in length, linear or linear-oblong, acute, sparsely puberulent to glabrous, 4-5
mm. long (only slightly exceeding the maturing achenes); corollas white, glabrous,
slightly ampliate upward, about 7 mm. long; achenes prismatic, black when mature,
strigillose on the ridges or glabrous.
Closely related to E. caeciliae, a terrestrial species not
uncommon in the montane forests of the western highlands.
Eupatorium monticola L. Wms. Fieldiana, Bot. 36: 91. 1975.
Figure 18.
Common in the open oak, pine and cypress forests at 2,600-
3,700 m.; Chimaltenango; Huehuetenango; Totonicapan (type,
Williams, Molina & Williams 41503); San Marcos. Mexico (Puebla,
Oaxaca and Chiapas).
WILLIAMS: FLORA OF GUATEMALA 81
Stout shrubs or small trees 0.5-5 m. tall, the young branches terete, covered with
reddish, purplish or ferruginous tomentum of somewhat viscid hairs; leaves opposite,
membranaceous, ovate to deltoid-ovate, acute or acuminate, shortly cuneate to the
petiole, crenate-serrate, 5-7-plinerved, with some nerves arising well above the base of
the blade, glabrous to usually sparsely puberulent above, usually short villosulous
below, especially along the nerves, or glabrescent, blades 5-17 cm. long and 3-10 cm.
broad (from very high elevations 3 cm. long or even less), petioles ferruginous-
tomentose becoming glabrous, 1-4 cm. long; inflorescence corymbose, rounded, 6-15
cm. broad, dense with many heads or in age somewhat lax, with reduced leaves or
bracteolate; heads narrowly campanulate, pedicellate, with 20-30 lilac or purple
(rarely white) flowers, 8-10 mm. long at an thesis; involucre less than half as long as
the heads; phyllaries subequal, linear, acute, mostly 4-5 mm. long; corolla
subcylindric, gradually enlarged upward, about 5 mm. long, lobes lanceiform, about
0.5 mm. long; styles exserted about 2 mm. at anthesis; achenes about 3 mm. long, 5-
ridged, scaberulous on the angles, black; pappus with about 25 bristles, about 4 mm.
long.
This is a common and attractive shrub or small tree on the
highest mountains of the western highlands.
Eupatorium montigenum Standl. & Steyerm. Field Mus. Bot.
23: 258. 1947.
Moist mountain forest, 2,000-3,000 m.; endemic; El Progreso
(type from Sierra de las Minas, north of Finca Piamonte, on slopes
toward summit of Volcan de Santa Luisa, Steyermark 43532);
Jalapa (Montana Miramundo).
A shrub or small tree of 3-6 m., the branches terete, very densely villous-pilose
with brownish or sordid, spreading hairs, the internodes short; leaves on slender
petioles 1.5-3.5 cm. long, thin-membranaceous, opposite, oblong-elliptic or elliptic-
ovate, 11-17 cm. long, 5-8 cm. broad, acute or acuminate, obtuse at the base,
inconspicuously serrate-dentate, especially toward the apex, or almost entire, green
above, puberulent, villosulous on the costa and nerves, slightly paler beneath, rather
densely villous-pilosulous or subtomentose, penninerved, the lateral nerves about 5 on
each side, arcuate, irregular, the blades epunctate, the veins translucent when viewed
against a strong light; inflorescence terminal, corymbose-paniculate, rounded, the
heads very numerous, apparently white, cymulose, mostly on slender pedicels 4-7
mm. long, the pedicels and branches very densely pilosulous, with soft, spreading
hairs; mature heads not seen; involucre 5-6 mm. high, campanulate, rather
densely appressed-pilosulous, about 10-flowered; phyllaries unequal, strongly gradu-
ate, about 3-seriate, at least the outer ones tinged with lavender or pale red, the
outermost short, ovate, obtuse or subacute, the innermost oblong, obtuse or rounded
at the apex; corolla 4 mm. long, glabrous, the tube very slender, cylindraceous;
pappus bristles white, 2.5 mm. long.
Both the collections are in bud only, so the mature heads may
be somewhat larger than described.
82 FIELDIANA: BOTANY, VOLUME 24
Eupatorium morifolium Mill. Card. Diet. ed. 8. no. 10. 1768.
E. populifolium HBK. Nov. Gen. & Sp. 4: 111. 1820. Critonia
morifolia King & Robinson, Phytologia 22: 49. 1971. Palo de agua;
vara de bajareque; chople (Peten); Santa Maria (Izabal).
Moist or wet thickets or moist forest, often in stony places
along streams or ravines or sometimes on open, brushy hillsides in
rather dry places, 1,600 m. or less, chiefly at low elevations;
sometimes planted at slightly higher elevations; Peten; Alta
Verapaz; El Progreso; Izabal; Zacapa; Santa Rosa; Escuintla;
Guatemala; Sacatepequez; Suchitepequez; Retalhuleu; Quezal-
tenango; San Marcos. Mexico; British Honduras to El Salvador and
Panama; South America.
Plants chiefly herbaceous but often somewhat woody, generally 1.5-4 m. high but
sometimes even taller, simple or sparsely branched, very stout, the stems terete,
costate, fistulose and hollow, striate, glabrous or somewhat villosulous or tomentose;
leaves large, pale green, rather thick, on long or short petioles, rounded-ovate to
oblong-ovate, mostly 10-20 cm. long but sometimes much larger, acute or acuminate,
subcordate to rounded or obtuse at the base, almost penninerved but more or less
triplinerved at the base, coarsely serrate or dentate, somewhat pubescent when young
but in age glabrous or nearly so, reticulate-veined; inflorescence usually very large
and leafy, rounded-paniculate, dense or rather open; heads very numerous, greenish
white or dirty white, 5 mm. long, densely glomerate, sessile or short-pedicellate;
phyllaries pale green or stramineous, ovate, obtuse, arachnoid-pubescent or glabrate,
striate, graduate, few-seriate; achenes glabrous.
Known in El Salvador as "taco," "chimaliote," "suelda con
suelda," "vara hueca," and "carrizo"; "cerbatana" (Honduras);
"lengua de vaca" (Oaxaca). This is a very common and well known,
generally weedy plant of the Pacific slope of Guatemala and El
Salvador. It grows most plentifully on plains or in foothills but
often is planted much higher, as at Antigua, and probably is really
naturalized at the higher elevations. It makes a rather neat and
attractive hedge, although it will not withstand stock. After
flowering, the branches often die back or are cut down for half their
length or more, after which they send up several long, rod-like, erect
branches that bear many large leaves. The flowers are not at all
handsome. The long hollow stems are often used for temporary
fencing, the sides of lowland huts, and other similar purposes.
Eupatorium muelleri Schultz Bip. ex Klatt, Leopoldina 20:
90. 1884. Mallinoa corymbosa Coulter, Bot. Gaz. 20: 47, t. 5. 1895
(type from Volcan de Jumaitepeque, Santa Rosa, Heyde & Lux
4255). Ageratina muelleri King & Robinson, Phytologia 19: 215.
1970. Escobita blanca. Figure 19.
WILLIAMS: FLORA OF GUATEMALA 83
Moist or rather dry places, brushy, often rocky hillsides, or
open, pine-oak forest, 1,000-2,200 m.; Zacapa; Chiquimula; Jalapa;
Santa Rosa; Sacatepequez; Chimaltenango; El Quiche; Huehue-
tenango. Southern Mexico; Honduras.
An erect or ascending, perennial herb, the stems usually simple below the
inflorescence, terete, villous below with rather long, spreading, white hairs, glabrous
above, the upper leaves usually much reduced and distant; leaves mostly basal,
membranaceous, slender- petio late, ovate to rounded-ovate, mostly 3-6 cm. long,
obtuse or acute, usually acute at the base but sometimes rounded or merely obtuse,
rather coarsely crenate, penninerved, thinly villous on both surfaces; inflorescence
corymbose-paniculate, sometimes small and bearing only a few heads but often broad
and open and bearing numerous heads, the bracts greatly reduced; pedicels usually
several times as long as the heads, glabrous; heads white, broadly campanulate,
many-flowered, 6-7 mm. high and usually somewhat broader; phyllaries few,
subequal, oblong, very obtuse, stramineous, thin, glabrous, ciliate about the apex;
achenes very small, black, obscurely scaberulous on the angles.
This plant is the type of the genus Mallinoa, which is evidently
referable to Eupatorium. It is a characteristic plant of pine-oak
forests of the Guatemalan mountains.
Eupatorium multinerve Benth. PI. Hartweg. 76. 1841.
Fleishmannia multinervis King & Robinson, Phytologia 19: 204.
1970. Flor de dolores (Chimaltenango).
Moist or wet thickets or in mixed or pine-oak forest, rarely a
weed in cafetales or other cultivated ground, 1,500-2,100 m.;
endemic; Chiquimula; Sacatepequez; Chimaltenango; El Quiche;
Huehuetenango; Quezaltenango (type from Quezaltenango, Hart-
weg 533}.
Plants very slender, herbaceous, probably always perennial, erect and 1 m. high
or usually much lower, or sometimes as much as 2.5 m. long and straggling or
supported on other vegetation, usually paniculately much branched, the stems terete,
greenish, densely puberulent to almost glabrous; leaves small, alternate, thin-
membranaceous, on slender petioles 1-3 cm. long, broadly cordate-ovate to deltoid-
ovate or lance-ovate, 2-6 cm. long, narrowly long-acuminate, usually more or less
cordate at the base but often rounded or only obtuse in the upper leaves, 3-nerved,
coarsely or rather finely and evenly crenate, usually thinly white-villous on both
surfaces, with short or rather long hairs, sometimes glabrate; heads very numerous,
on long, slender pedicels, 4 mm. long, white or rarely purplish, 15-20-flowered,
forming numerous, very lax, often leafy-bracted cymes; pedicels usually glabrous,
rarely with a few scattered, short hairs; phyllaries oblong, pale green, glabrous, few-
seriate, laxly imbricate, usually somewhat lustrous, conspicuously bicostulate,
rounded or very obtuse at the apex, not ciliate, or the outermost, very short ones
ovate or oval and ciliate; corollas glabrous, not or scarcely exceeding the involucre;
achenes fuscous, very slender, glabrous or nearly so, 1.5 mm. long; pappus scant,
white, scarcely equaling the corolla.
84 FIELDIANA: BOTANY, VOLUME 24
A member of a complex group related to E. microstemon Cass.
Much in need of revision.
Eupatorium nubigenoides Rob. Proc. Am. Acad. 42: 42. 1906.
Fleishmanniopsis nubigenoides King & Robinson, Phytologia 21:
404. 1971.
Moist or wet, mixed forest, often or perhaps always on
limestone, 900-1,150 m.; endemic; Alta Verapaz (type from
Pansamala, Tuerckheim 928; collected also at Sasis and Chirriacte).
A weak shrub about 1 m. high, glabrous throughout except in the inflorescence,
the branches very slender, terete; leaves opposite, thin, on long slender petioles,
lanceolate, 3-5-plinerved well above the base, ovate-oblong to broadly lanceolate,
about 13 cm. long and 5 cm. broad or smaller, long-acuminate, acute to attenuate at
the base, cuspidate-dentate; corymbs small, with numerous opposite branches,
puberulent or tomentulose, leafy-bracted or almost naked; heads numerous, short-
pedicellate, white, about 10-flowered, 6 mm. high, sometimes almost or quite sessile
and clustered; involucre subcylindric-campanulate; phyllaries whitish or stramineous,
laxly appressed, unequal, strongly imbricate, about 4-seriate, striate, the outer ones
very short, ovate, subacute, ciliate, the middle ones ovate-oblong, rounded at the
apex, glabrous, the innermost narrowly oblong; corollas narrowly tubular, 3 mm.
long; achenes 1 mm. long, blackish, glabrous; pappus white or yellowish white.
The materials seen by the author are inadequate for an
understanding of the species.
Eupatorium nubigenum Benth. PL Hartweg. 85. 1841; L.
Wms. Fieldiana, Bot. 36: 94. 1975. E. hospitale Rob. Proc. Am.
Acad. 43: 32. 1907. E. microdon Rob. Proc. Am. Acad. 54: 252. 1918.
Critonia hospitalis King & Robinson, Phytologia 22: 49. 1971.
Critoniadelphus microdon King & Robinson, torn. cit. 53. C.
nubigenus King & Robinson, I.e. Varilla blanca.
Mixed forest and on open slopes in cut-over lands, sometimes in
Cupressus forests, 1,200-3,100 m.; Alta Verapaz (type of E.
microdon, Tuerckheim II. 2261); Jalapa; Zacapa; Chiquimula;
Guatemala; Chimaltenango; Suchitepequez (type of E. nubigenum
from Las Nubes, Hartweg 587); El Quiche; Quezaltanango; San
Marcos. Mexico (Vera Cruz and Chiapas).
A shrub or small tree to 9 m. tall, glabrous, the branches slender, angulate or
subangulate; leaves membranaceous, on petioles 1-2 cm. long, the blades narrowly
lanceolate to lance-oblong, 7-25 cm. long and 2-9 cm. broad, long acuminate, acute or
attenuate at the base, dentate to nearly entire, penninerved, with the nerves mostly
7-11 pairs diverging at a broad angle and conspicuous, glabrous; inflorescence
hemispheric, terminal panicles mostly 6-15 cm. broad; heads sessile in dense
glomerules, 6-8 mm. long, 3-8-f lowered; involucre subcylindric; phyllaries very
unequal, few, the outer ones lance-ovate, obtuse or subacute, puberulent or ciliolate,
WILLIAMS: FLORA OF GUATEMALA 85
short, persistent, the inner ones erose-ciliolate, somewhat stramineous, cauducous;
corollas white, slender tubular, about 4.5-5 mm. long; achenes blackish, about 2 mm.
long, glabrous or puberulent on the angles; pappus pale yellow, equaling the corollas.
The group to which this species belongs is complicated and it is
possible that a still broader concept of it may be forthcoming, when
monographic study is undertaken.
Eupatorium nubivagum L. Wms. Fieldiana, Bot. 36: 94. 1975.
Figure 20.
Known only from subalpine meadows of the Cuchumatanes at
about 3,300 m., endemic; Huehuetenango (type, Molina 21238). To
be expected in Mexico (Chiapas).
Subalpine herbs to 0.5 m. tall, erect or ascending, the stems densely crisped
puberulent, leafy, terete; leaves ovate or ovate-lanceolate, acute, serrate, sparsely
crisped puberulent on both surfaces, triplinerved from the acute base, blade to 4 cm.
long and 2.5 cm. broad, reduced upward, the petioles to about 2 cm. long;
inflorescence terminal, corymbose panicles to about 7 cm. long and nearly as broad,
the heads about 5-20 in each corymb; heads campanulate, 7-9 mm. long and nearly as
broad, with about 40 florets or fewer; involucres biseriate and lacking small basal
phyllaries; phyllaries subequal, linear-oblong or linear-oblanceolate, acute, erose
above, ciliolate and the outer ones puberulent, prominently 3-nerved, about 5 mm.
long and 1-1.5 mm. broad, much shorter than the pappus and florets; corollas tubular
to about the middle, narrowly campanulate above, sparingly pilose at the apex, 3.5-4
mm. long; lobes triangular-ovate, acute, about 0.4 mm. long; styles exserted about 1
mm. at anthesis, terete and slightly clavate; achenes black, 5-ridged, sparingly
strigillose, 2.5-3 mm. long; pappus white, barbellate, about 4 mm. long.
Related to E. muelleri which is found at middle elevations in
Guatemala, it may be distinguished by the larger heads, with the
phyllaries only about two-thirds as long, the corolla campanulate
above, and the leaves triplinerved.
Eupatorium odoratum L. Syst. Nat. ed. 10. 1205. 1759. E.
conyzoides Mill. Gard. Diet. ed. 8. no. 14. 1768. Chromolaena
odorata King & Robinson, Phytologia 20: 204. 1970. Curarina de
Jmonte; canutillo; cruz-quen (Alta Verapaz); curarina; crucetilla;
suplicio; cruz de campo (fide Aguilar).
Wet to dry thickets, common in second growth, often in rather
dry, rocky places, frequently in hedges, 1,650 m. or less, most
plentiful below 1,000 m.; Alta Verapaz; Baja Verapaz; El Progreso;
Izabal; Zacapa; Chiquimula; Jutiapa; Santa Rosa; Escuintla;
Guatemala; Sacatepequez; Suchitepequez; Retalhuleu; San
Marcos; El Quiche; Huehuetenango. Mexico; British Honduras to
El Salvador and Panama; West Indies; South America.
86 FIELDIANA: BOTANY, VOLUME 24
A coarse shrub, commonly 1-2 m. high or more, often densely branched, the
branches often recurved or drooping or frequently elongate and more or less
scandent, the ultimate branches usually divergent at a wide angle, terete, villous or
variously pubescent to almost glabrous; leaves membra naceous, slender-petiolate,
deltoid-ovate or rhombic-ovate, highly variable in size and shape, acuminate to long-
attenuate, acute or obtuse at the base, 3-nerved, the upper leaves usually narrower,
coarsely serrate or crenate or subentire, often somewhat hastately dentate near the
base, varying from glabrous to tomentose; inflorescence usually large and broad,
corymbiform, flat-topped or convex; heads cylindric, lavender or whitish, about 1 cm.
long, 20-35-flowered, generally pedicellate; achenes fuscous or black, scabrous on the
angles.
Sometimes called "chimuyo" in El Salvador; "crucito," "rey del
todo," "mejorana" (Honduras); "tocaban," "tocabal" (Yucatan,
Maya); "yaxhatz" (British Honduras, Maya). One of the most
common and widely distributed species of the genus, this is a plant
of decidedly weedy habit but usually attractive. It is abundant in
most parts of the Central American lowlands and is one of the first
plants to take possession of abandoned land. The plants are highly
variable in pubescence and form of the leaves, but the variations are
so inconsistant that they scarcely seem worthy of varietal
designation. Some use is made of the plant in the domestic medicine
of Guatemala.
Eupatorium ovandense Grashoff & Beaman, Rhodora 71:
577, fig. 1969. Decachaeta ovandensis King & Robinson, Brittonia
21: 397. 1969.
Probably from wet forests at some 2,000 m., known only from
Mt. Ovando, Chiapas, Mexico, and to be expected in Guatemala.
Probably large, suffrutescent herbs more than 1 m. tall; leaves alternate, ovate,
coarsely dentate, acute, the base obtuse or truncate, 3-5-plinerved, well above the
base, mostly 10-20 cm. long and 8-15 cm. broad, glabrous or sparsely pubescent on the
nerves, with many minute glands below, petioles mostly 3-2 cm. long; inflorescences
axillary, about as long as the subtending leaves or perhaps much longer, corymbose,
sordid-puberulent; heads 6-7 mm. long, with about 15 florets, short pedicellate;
phyllaries uni- or biseriate, subequal, or with shorter ones at the base, linear-
lanceolate, acute, puberulent, mostly 3-4 mm. long; corollas mostly 3-4 mm. long,
glabrous, slightly expanded upward; achenes prismatic, about 2 mm. long, strigillose
on the angles.
Related to E. incomptum. The description is drawn from the
type material, which is not good.
Eupatorium ovillum Standl. & Steyerm. Field Mus. Bot. 22:
305. 1940. Ageratina ovilla King & Robinson, Phytologia 19: 224.
1970.
WILLIAMS: FLORA OF GUATEMALA 87
Moist or wet thickets or mixed forest, 2,400-3,200 m.; endemic,
so far as known, but to be expected in Chiapas; El Progreso (Sierra
de las Minas); Solola; Totonicapan (type from Cumbre del Aire, on
the road between Huehuetenango and Sija, Standley 65862);
Quezaltenango; San Marcos.
An erect shrub of 1.5-2.5 m. or sometimes more elongate and subscandent, the
branches rather stout, brown, terete, densely viscid-puberulent; leaves opposite,
thick-membranaceous, on stout petioles 6-14 mm. long, broadly ovate or rounded-
ovate, 1.5-3 cm. long, 1-2.5 cm. broad, acute to very obtuse, broadly rounded or
truncate at the base, 3-plinerved a short distance above the base, remotely and
inconspicuously crenate or subentire, sparsely and minutely granular-puberulent
above or almost glabrous, paler beneath, densely and minutely granular-punctate;
heads white or purplish white, about 10-flowered, in small, rounded or corymbiform
cymes, terminating short, leafy, lateral branches and forming an elongate, leafy
panicle, the slender pedicels 7 mm. long or less, viscid-puberulent; involucre narrowly
campanulate, 7-8 mm. long; phyllaries about 10, subequal, linear, pale greenish, acute
or attenuate, 2-3-costulate, sparsely and minutely viscid-puberulent; corollas
glabrous, 5 mm. long; achenes 2.3 mm. long, glabrous; pappus white, 4 mm. long,
minutely scaberulous.
Closely related to E. tomentellum, a species of the lower slopes
of the mountains.
Eupatorium pazcuarense HBK. Nov. Gen. Sp. 4: 123. 1820. E.
subpenninervium Schultz Bip. ex Klatt, Leopoldina 20: 89. 1884. E.
skutchii Rob. Contr. Gray Herb. 104: 27. 1934. Kyrstenia pazcua-
rensis Greene, Leafl. Bot. Obs. and Grit. 1: 9. 1903. Ageratina
pazcuarensis King & Robinson, Phytologia 19: 215. 1970. A.
skutchii King & Robinson, I.e. 217.
Wet to rather dry, mountain thickets or in mixed or pine-oak
forest, often in forest of Cupressus, Abies, Alnus, or Juniperus
standleyi, frequently on limestone, 1,500-3,400 m., mostly at the
higher elevations; Alta Verapaz; Baja Verapaz; Jalapa; Chimal-
tenango (type of E. skutchii from Santa Elena, Skutch 337); Solola;
Huehuetenango; Totonicapan; Quezaltenango; San Marcos. Central
and southern Mexico.
An erect, perennial herb, usually 1 m. high or less, sometimes 1.5 m. tall, rarely
suffrutescent below, the stems terete, often much branched, puberulent or pubescent
with sparse, mostly appressed hairs or glabrate; leaves membranaceous, on long or
short, slender petioles, ovate to rounded-ovate, mostly 5-9 cm. long, acute or
acuminate, rounded or somewhat cordate at the base, about 5-plinerved from a point
slightly above the base, coarsely and closely crenate or obtusely dentate, puberulent
or sparsely villosulous above, glabrate beneath but sparsely puberulent or appressed-
pubescent on the nerves and veins; heads many- flowered, campanulate, 5-7 mm. long,
slender-pedicellate, forming small, dense corymbs, these arranged in a small or large,
often leafy-bracted, compound corymb; phyllaries subequal, pale greenish, linear or
88 FIELDIANA: BOTANY, VOLUME 24
oblong-linear, obtuse or acute, minutely puberulent or almost glabrous, erose-
denticulate about the apex; corollas white, slightly longer than the involucre; achenes
blackish, glabrous or puberulent on the angles.
Eupatorium perpetiolatum (King & Robinson) L. Wms.
Fieldiana, Bot. 36: 99. 1975. Pseudokyrsteniopsis perpetiolata King
& Robinson, Phytologia 27: 241. 1973.
Known only from dry mountain slopes near Rio Blanco, El
Quiche, Williams, Molina & Williams 22457. Endemic.
Erect herbs to about 2 m. tall, much branched, the stems terete, puberulent or
glandular-puberulent above; leaves small, trinerved from the base, ovate or ovate-
cordate, acute, sparsely glandular-puberulent on both surfaces, mostly 2-5 cm. long
and 1-3 cm. broad; petioles slender, to 3 cm. long or perhaps more, and reduced
upward, the bases thickened; inflorescences small, corymbose panicles, terminating
the several, increasingly shorter, leafy branches, pedicels very short, glandular-
pubescent; heads 12-20-flowered, mostly 8 mm. long and 3-4 mm. broad; phyllaries in
3-4 series, linear-lanceolate, acuminate, up to 8 mm. long; corolla green, 3-4 mm. long,
prominently constricted at the throat, the lobes minute, about 0.2 mm. long; achenes
prismatic, 4-5-ridged, puberulent, about 2.5 mm. long; pappus barbellate, about 5
mm. long.
This plant is the basis of the monotypic genus Pseudokyrsten-
iopsis, as indicated above.
Eupatorium phoenicolepis Rob. Proc. Am. Acad. 35: 338.
1900. E. phoenicolepis var. guatemalensis Rob. Proc. Am. Acad. 43:
34. 1907 (type from Volcan de Atitlan, Solola, W. A. Kellerman
5199).
Moist or wet, mixed, mountain forest, sometimes in oak forest
or thickets, 1,200-2,850 m.; El Progreso; Zacapa; Sacatepequez;
Chimaltenango; Solola; El Quiche; Huehuetenango; Quezaltenango.
Mexico (Chiapas); Honduras.
A rather stout shrub of 1.5-3 m., the branches terete, villosulous-tomentose;
leaves thin, large, on very long, slender petioles, ovate or broadly ovate-cordate,
mostly 10-14 cm. long, acuminate or long-acuminate, shallowly and openly cordate at
the base or broadly rounded, trinerved, crenate or almost entire, scaberulous or
glabrate above, densely whitish tomentulose and pale beneath or sometimes green
and glabrate; heads usually very numerous, on long slender pedicels, forming small,
convex or ovoid corymbs, these arranged in a large, lax, leafy panicle, about 18-
flowered, 1 cm. long, purplish pink, the pedicels densely viscid-villosulous; phyllaries
very unequal, laxly imbricate, usually intensely pink or purplish, the inner ones
obtuse, narrowly oblong or lanceolate, the outermost ovate, acute or subacute;
achenes blackish, scaberulous; pappus white.
Eupatorium pinabetense Rob. Proc. Am. Acad. 36: 482. 1901;
L. Wms. Fieldiana Bot. 36: 99. 1975. Neobartlettia pinabetensis
WILLIAMS: FLORA OF GUATEMALA 89
King & Robinson, Phytologia 21: 296. 1971. Bartlettina pinabetensis
King & Robinson, Phytologia 22: 161. 1971. Tunehuac (San
Marcos).
Dense, moist or wet, mixed, mountain forest, often on white
sand slopes, 1,400-2,850 m.; Quezaltenango; San Marcos. Mexico
(Chiapas, the type collected near Pinabete).
A shrub or tree of 2-6 m., the branches angulate, striate, glabrous; leaves
opposite, narrowly lanceolate or lance-oblong, mostly 10-16 cm. long and 2-3 cm.
broad, narrowly long-acuminate, attenuate to the base, penninerved, acutely and
closely serrate, glabrous, somewhat succulent when fresh, drying dark green or
blackish; inflorescence puberulent or minutely sordid-tomentulose, the inflorescence
much shorter than the leaves, dense or open, about 10 cm. broad or less; heads
numerous, lavender or purplish white, crowded, about 10-flowered, pedicellate, 6 mm.
long; phyllaries few, unequal, laxly imbricate, elliptic, rounded at the apex, sordid-
stramineous, erose-ciliolate; corolla glabrous or nearly so, about equaling the
involucre; achenes glabrous, almost 2 mm. long; pappus white.
A very common shrub or small tree in the deep ravines of the
higher mountains of Quezaltenango and San Marcos, highly
ornamental and handsome when in full flower. It grows in places
that are enclosed in fog at night and often also during the day.
Closely related to E. tuerckheimii and also to E. hebebotryum.
Eupatorium pittieri Klatt, Bull. Soc. Bot. Belg. 31, pt. 1: 192.
1892. Koanophyllon pittieri King & Robinson, Phytologia 22: 150.
1971. Palo negro; soyoco (San Marcos).
Moist or wet, mixed forest or thickets, 2,000 m. or less, most
frequent at less than 1,000 m.; Peten; Alta Verapaz; Izabal;
Quezaltenango; San Marcos. Southern Mexico; British Honduras;
Honduras; Costa Rica.
A shrub or tree of 3-9 m., the slender branches subterete, obscurely puberulent or
almost glabrous; leaves short-petiolate, firm-membranaceous, elliptic-oblong to
oblong-lanceolate or elliptic, 9-20 cm. long, narrowly long-acuminate, acute or
attenuate at the base, penninerved, coarsely serrate to subentire, glabrous or nearly
so on both surfaces, prominently reticulate below, opaque, often lustrous; heads
white or greenish white, sometimes tinged with purple, about 5 mm. long and 10-
flowered, pedicellate, forming larger, terminal, leafy-bracted or almost naked, dense
or rather open, rounded or pyramidal panicles, the branches minutely puberulent or
strigillose; phyllaries very unequal, about 3-seriate, ovate, obtuse or rounded at the
apex, striate, ciliolate; achenes hispidulous on the angles; pappus dirty white or pale
yellowish.
This species has been reported from Guatemala and British
Honduras as E. billbergianum Beurl.
Eupatorium platyphyllum Rob. Proc. Am. Acad. 35: 339.
90 FIELDIANA: BOTANY, VOLUME 24
1900. Neobartlettia platyphylla King & Robinson, Phytologia 21:
296. 1971. Bartlettina platyphylla King & Robinson, Phytologia 22:
161. 1971.
Moist or wet forest, 950-1,350 m.; Alta Verapaz; Suchitepequez.
Southern Mexico; Costa Rica.
A shrub as much as 5.5 m. high, the younger branches herbaceous, obscurely 6-
angulate, at first tomentulose or sordid-puberulent, in age glabrate; leaves large, thin,
opposite, very broadly deltoid-ovate, mostly 12-20 cm. long and 22 cm. broad or
narrower, usually somewhat hastate-angulate at the base, acute or acuminate,
broadly rounded at the base, crenate-dentate, 3-nerved, appearing glabrous but more
or less puberulent on the nerves and veins, the basal lobes rounded to acute; heads
white, 60-75-flowered, about 12 mm. high, slender-pedicellate, somewhat aggregate at
the ends of the branches of the large corymb, this as much as 30 cm. broad, leafy,
rounded or flat-topped; involucre broadly campanulate, often broader than high;
phyllaries several-seriate, regularly but laxly imbricate, very unequal, stramineous,
oblong or oval, obtuse, striate, scarious and erose at the apex, glabrous or nearly so;
corollas slender, 6 mm. long, little if at all exceeding the involucre; achenes glabrous,
2 mm. long; pappus white or yellowish white.
Apparently rare in Guatemala, for we have not collected this
plant there, and the species is not known between Guatemala and
Costa Rica.
Eupatorium prunellaefolium HBK. Nov. Gen. Sp. 4: 123.
1820. E. salinum Standl. & Steyerm. Field Mus. Bot. 23: 188. 1944
(type from Jutiapa, Steyermark 31775). Ageratina prunellaefolia
King & Robinson, Phytologia 19: 215. 1970. Heliotropo de monte
(Huehuetenango).
Moist or wet, mountain meadows or forests, 650-3,600 m.;
Jutiapa; Sacatepequez (Volcan de Agua); Huehuetenango. Moun-
tains of central and southern Mexico.
A perennial herb, the stems erect, terete, often purplish, 1 m. high or usually
lower, puberulent or crisp-pubescent, simple or laxly branched; leaves membrana-
ceous, on long or short petioles, opposite, broadly ovate to broadly rhombic-ovate,
often as broad as long, mostly 3-4 cm. long, rounded to subacute at the apex, usually
obtuse, broadly rounded to broadly cuneate at the base, trinerved or sometimes
triplinerved from a short distance above the base, coarsely crenate or crenate-
dentate, sparsely villous on both surfaces, with weak, whitish hairs or almost wholly
glabrous; heads few, white, 8-9 mm. long, broadly campanulate, many -flowered, on
short or very long pedicels, usually in small, leafy-bracted cymes of very few heads,
or the inflorescence often trichotomous and the cymes long-pedunculate; phyllaries
often purplish, subequal, oblong-linear or lance-linear, acute or subobtuse, densely
glandular-puberulent or short-villosulous, almost equaling the florets; corollas with a
much dilated limb, 3 mm. long, pubescent about the base of the limb; achenes
blackish, slender, 2.5 mm. long, pubescent or almost glabrous; pappus soft, pinkish.
WILLIAMS: FLORA OF GUATEMALA 91
Eupatorium pycnocephaloides Rob. Proc. Am. Acad. 51: 534.
1916. E. pycnocephaloides var. glandulipes Rob. op. cit. 535 (type
from Totonicapan, E. W. D. Holway 106). Fleishmannia pycno-
cephaloides King & Robinson, Phytologia 19: 205. 1970. Shup
(Quezaltenango); Jolomacach (Totonicapan).
Moist or wet, mixed forest or thickets, often in deep sand or in
rocky places, sometimes in forest of oak, pine, Cupressus, Juniperus,
or Alnus, 1,800-3,300 m., mostly at 2,400 m. or more; so far as
known, endemic to Guatemala, but probably found also in Chiapas;
Guatemala; Sacatepequez; Chimaltenango; Solola (type from
Solola, E. W. D. Holway 144); Totonicapan; Huehuetenango;
Quezaltenango; San Marcos.
A much branched, perennial herb, usually erect and 1.5 m. high or less,
sometimes more elongate and clambering or reclining, rarely suffrutescent below, the
stems slender, terete, striate, often flexuous, frequently purplish, pubescent or short -
villous, often glabrate; leaves opposite, on slender petioles 2-3.5 cm. long,
membranaceous, mostly deltoid-ovate or ovate, commonly 4-7 cm. long but variable
in size and shape, acuminate or long-acuminate, subtruncate or rounded at the base,
sometimes subcordate, finely or coarsely crenate-serrate or dentate, trinerved,
glabrate or thinly villosulous above, usually abundantly pilose or short-villous
beneath, at least on the nerves and veins; panicles terminal, often much branched
and very leafy, usually lax and open; heads numerous, white or dirty white, 6-7 mm.
long, 15-20-flowered, narrowly campanulate, forming dense or lax, convex or often
rounded clusters, the slender pedicels 2-8 mm. long, pubescent or glandular-
pubescent; phyllaries green or greenish, about 3-seriate, unequal, often tinged with
purple, striate with 2 thick stramineous nerves, these joined at the base, pubescent or
glabrate, the outer ones short, ovate, acuminate, the middle ones oblong, acute, the
innermost linear-oblong, obtuse; corollas glabrous, narrowly tubular; achenes dark
brown or blackish, 1.7 mm. long, hispidulous on the angles; pappus white or yellowish
white, scarcely as long as the corolla.
Var. glandulipes is a form in which the pedicels are glandular-
pubescent, while in the typical variety the pubescence is eglandular.
This character apparently is not very important, since about half
the material is referable to the variety, which is distributed almost
as widely as the typical form. Originally based on two specimens
collected in Solola, this species proves to be one of the commonest
ones of Guatemala, 50 collections having been made in recent years.
It is somewhat variable in habit, leaf form, and appearance of the
inflorescence, whether lax or dense, but the heads are remarkably
uniform in their characters.
Eupatorium pycnocephalum Less. Linnaea 6: 404. 1831. E.
schiedeanum Schrad. Ind. Sem. Hort. Goetting. 1832: 3. 1832. E.
anisopodum Rob. Proc. Am. Acad. 36: 477. 1901 (?) (type from Baja
92 FIELDIANA: BOTANY, VOLUME 24
Verapaz, Tuerckheim 1177). E. pratense Klatt, Bull. Soc. Bot. Belg.
31: 193. 1892. Fleishmannia anisopoda King & Robinson,
Phytologia 19: 202. 1970. F. pratensis King & Robinson, I.e. 205. F.
pycnocephala King & Robinson, I.e. 205. Mejorana; bretillo;
llovizna', te (Izabal); flor de Octubre (Chiquimula); inmortal
(Chimaltenango).
Moist or wet, mixed forest, or often in forest of pine, oak,
Cupressus, Alnus, or Juniperus, sometimes on limestone, wet to dry
thickets, often in rocky places or on cliffs, rarely in marshes,
frequently in pastures or other open fields, 3,300 m. or usually
much less; Peten; Izabal; Alta Verapaz; Baja Verapaz; El Progreso;
Zacapa; Chiquimula; Jalapa; Jutiapa; Santa Rosa; Escuintla;
Guatemala; Sacatepequez; Chimaltenango; Suchitepequez; Retal-
huleu; El Quiche; Solola; Totonicapan; Huehuetenango, Quezal-
tenango; San Marcos. Mexico; British Honduras to El Salvador and
Panama; northern South America.
A perennial herb, usually erect and 1 m. high or less, often not more than 35 cm.
high, the stems simple or usually branched, terete, puberulent to villous; leaves
opposite, slender- petio late, membranaceous, ovate or usually deltoid-ovate, mostly 4-
6 cm. long, acuminate or long-acuminate, rounded to subcordate at the base or in the
upper leaves cuneate, trinerved, crenate-serrate, sparsely or densely pilose, especially
beneath, rarely glabrate; inflorescence leafy, usually open and rather lax, with
spreading or ascending branchlets, each of these bearing a rounded, terminal cluster
of 7-20 or more, short -pedicellate heads; heads purple, lavender, pinkish, or rarely
white, about 25 flowered, 4-5 mm. long; involucre campanulate; phyllaries about 3-
seriate, the inner ones oblong, obtuse, bicostulate, pale or greenish, glabrous or nearly
so, ciliate, scarious at the apex, the outer ones mostly ovate and acute or subacute;
corollas glabrous, about equaling the involucre; achenes black, little more than 1 mm.
long, glabrous; pappus scant, white.
Called "acahualera" in Veracruz, this is one of the most
common and widely distributed Eupatorium species of Central
America, often becoming weedy and sometimes invading cafetales
and other cultivated ground. It is abundant in most of the
uncultivated mountain areas and occurs in a great variety of
habitats. Notwithstanding, the plants are remarkably uniform in
most of their characters. E. anisopodum may be a distinct species,
but we have seen no authentic material of it, and the original
description mentions no character that is not found commonly in E.
pycnocephalum.
Eupatorium quadrangulare DC. Prodr. 5: 150. 1836. Critonia
quadrangularis King & Robinson, Phytologia 22: 50. 1971.
WILLIAMS: FLORA OF GUATEMALA 93
Moist or wet thickets or mixed, lowland forest, frequently along
streams or in ravines, 1,400 m. or usually at lower elevations;
Zacapa; Chiquimula; Santa Rosa; Escuintla; Guatemala; Sacatepe-
quez; Solola; Suchitepequez; Retalhuleu. Mexico; El Salvador;
Honduras; Nicaragua; Costa Rica.
A coarse, stout, erect herb 1.5-3 m. high, simple or sparsely branched, the
branches green, 4-angulate, glabrous or nearly so, hollow, striate; leaves large, thick-
membranaceous, opposite, ovate, acute, mostly 20-30 cm. long or larger, abruptly
narrowed below into a broad, petioliform portion, this sessile and auriculate at the
base, coarsely serrate or dentate, in age glabrous or nearly so, when young often
villosulous along the nerves; inflorescence usually very large, thyrsiform or broad,
rounded; heads very numerous, white, 8- 10-flowered, glomerate and often forming
dense rounded clusters, pedicellate or subsessile, about 7 mm. long; phyllaries
stramineous, ovate-oblong, very obtuse, graduate, several-seriate, striate; achenes
puberulent; pappus yellowish.
Known in El Salvador by the names "tallo hueco," "chima-
liote," "chimaliote hueco." The leaves and young branches are eaten
by cattle. The plant is a common one at many places along the
Pacific plains, usually growing in or near the rather thin, open
forests.
Eupatorium quercetorum L. Wms. Fieldiana, Bot. 36: 101.
1975.
Oak woods or oak-pine woods at 900-1,200 m.; to be expected in
Guatemala. Mexico (type from Chiapas near the Guatemalan
border, Breedlove 14046).
Shrubs to 2.5 m. or more, the branches slender, terete, pubescent becoming
glabrous; leaves elliptic to elliptic-oblanceolate, acuminate, serrate, acute to the
petiolate base, penninerved with 4-5 pairs of nerves, glabrous above, puberulent along
the nerves below, the blade 4-9 cm. long and 1.5-2.5 cm. broad, the petiole slender,
puberulent, 0.4-10 mm. long; inflorescences terminal, cymose- paniculate, arachnoid-
puberulent, with 20-30 heads, to 5 cm. long and as broad; heads cylindric, 8-10 mm.
long and 2-3 mm. in diameter, with 15-20 flowers in each head; phyllaries 5-6-seriate,
puberulent, the outermost suborbicular-ovate, obtuse, about 2 mm. long, the inner
series becoming longer and narrower, the innermost linear, obtuse, about 7 mm. long,
reaching almost to the top of the pappus; corolla white, subcampanulate at the
throat, about 4.5 mm. long and 1.2 mm. broad at the throat, the lobes ovate-
triangular, acute; anthers appendaged at the apex, linear-oblong, about 1.5 mm. long;
style filiform, surpassing the corolla by about 4 mm. at anthesis; achenes glabrous, 5-
ridged; pappus about 4 mm. long, composed of about 30 nearly smooth bristles.
Eupatorium rojasianum Standl. & Steyerm. Field Mus. Bot.
23: 188. 1944.
Moist thickets, 1,200-1,650 m.; endemic; Quezaltenango (type
94 FIELDIANA: BOTANY, VOLUME 24
collected near El Muro below Santa Maria de Jesus, Standley
67221', collected also along the road above Santa Maria).
An erect, perennial herb about 1 m. high, branched, the branches slender, terete,
greenish, very minutely and sparsely puberulent or almost glabrous, striate; leaves
opposite, membranaceous, on slender petioles 2-5.5 cm. long, deltoid-ovate, 3.5-7 cm.
long, 1.5-4.5 cm. broad, narrowly long-acuminate, truncate to rounded-cuneate at the
base, trinerved, closely and evenly serrate-dentate, green and almost glabrous above,
slightly paler beneath, almost glabrous or sparsely puberulent on the nerves; heads
white, few or numerous, 5-flowered, slender-pedicellate, clustered in small cymes,
terminating short lateral branches, the cymes laxly paniculate, the pedicels sparsely
and minutely pilosulous; involucre subcylindric, 5 mm. long; phyllaries pale green,
about 3-seriate, the inner ones narrowly oblong, rounded and subscarious at the apex,
glabrous, the outer ones short, appressed, oval-ovate, obtuse; corollas glabrous,
slender-tubular, 2.5 mm. long; achenes black, 1.6 mm. long, glabrous; pappus bristles
white, equaling the corolla.
The species was named for Prof. Ulises Rojas, Director of the
Jardin Botanico of Guatemala.
Eupatorivun saxorum Standl. & Steyerm, Field Mus. Bot. 23:
189. 1944.
Dry, shaded, rocky slopes or on limestone in Juniperus forest,
2,500-4,000 m.; endemic; Huehuetenango (La Pradera, Sierra de los
Cuchumatanes); San Marcos (type collected between Sibinal and
summit of Volcan de Tacana, Steyermark 36075).
An erect, perennial herb or subshrub about 20 cm. high, the stems several or
numerous, simple or sparsely branched, very sparsely short-pilosulous or almost
glabrous, terete, brownish or purplish, rather densely leafy below; leaves small,
opposite, on stout petioles 3-5 mm. long, thick-membranaceous, ovate, deltoid- ovate,
or oblong-ovate, 1-2 cm. long, 6-15 mm. broad, narrowed to the obtuse apex, rounded
at the base, trinerved, inconspicuously crenate-serrate or almost entire, green above,
minutely scaberulous or glabrate, strigose beneath on the nerves and veins or almost
glabrous; heads few, about 6 mm. long, slender-pedicellate, about 15-flowered,
cymose-fasciculate at the ends of the branches, the inflorescence simple or of 3 long-
pedunculate cymes; phyllaries few, lance-linear, subequal, acute or acuminate, 2-3-
costulate, glabrous or glabrate, ciliolate, much shorter than the florets; corollas
slender, glabrous, 3 mm. long, white; achenes blackish, 2 mm. long, hispidulous on the
angles; pappus bristles whitish or pale purplish, scaberulous, slightly shorter than the
corolla.
In general appearance the plant suggests a reduced form of E.
pycnocephalum, but differs in that the involucre is several-seriate,
rather than having the phyllaries almost equal in length.
Eupatorium schultzii Schnittspahn, Zeitschr. Gartenb.
Darmst. 6. 1857. Peteravenia schultzii King & Robinson, Phytologia
21: 395. 1971.
WILLIAMS: FLORA OF GUATEMALA 95
Coarse, tall, sometimes lignescent herbs, 1-3 m. tall, the branches terete, closely
and finely tomentulose or glabrate; leaves usually large, thin, ovate-cordate, or
rounded-cordate, acute or acuminate, shallowly to deeply cordate at the base, 3-5-
nerved from the base, unevenly crenate-dentate, sparsely scaberulous or puberulent
above, beneath densely tomentulose or velutinous-pilosulose, (5-)15-20 cm. long or
large and nearly as broad; inflorescence of many flat-topped or rounded corymbose
panicles, these often disposed in very large, leafy panicles, with up to 100 heads in
each panicle but usually fewer; heads mostly 8-10 mm. long, campanulate, with
about 40-50 flowers; phyllaries usually with 3 prominent nerves, several-seriate, very
unequal, the outermost about 1 mm. long and ovate, the innermost linear, obtuse and
6-8 mm. long, glabrous; corollas white or usually pink, tubular and abruptly
subcampanulate at the throat, 4-5 mm. long; achenes black when mature, about 3
mm. long, 5-ridged, sometimes puberulent on the angles; pappus white or purplish,
about as long as the inner phyllaries.
This is an occasional but widely distributed species in the
mountains of Guatemala, as elsewhere in Mexico and Central
America. Dr. Robinson has divided it into the following four forms,
which are usually separable but probably of little importance. One,
f. erythranthodium, is at best a color form.
Pedicels velutinous-tomentulose, the glandular hairs none or very few f. velutipes.
Pedicels densely glandular-pilosulous or glandular-puberulent.
Phyllaries tinged with pink or purple f. erythranthodium.
Phyllaries without pink or purplish coloring.
Pedicels glandular-puberulent and with sparse, longer, eglandular hairs.
f. schultzii.
Pedicels glandular-puberulent or glandular-pilosulous, without longer, eglandular
hairs f. ophryolepis.
Eupatorium schultzii f. schultzii.
Moist or dry, brushy, often rocky hillsides or in mixed forest,
350-1,900 m.; Alta Verapaz; Baja Verapaz; Guatemala; Suchitepe-
quez; Quiche. Southern Mexico; Costa Rica.
Eupatorium schultzii f. erythranthodium Rob. Proc. Am.
Acad. 51: 536. 1916.
Moist thickets or forest, sometimes in oak forest, 350-1,950 m.;
Alta Verapaz; Baja Verapaz; Huehuetenango. Mexico.
This form is hardly worthy of special designation.
Eupatorium schultzii f. ophryolepis Rob. Proc. Am. Acad.
51: 536. 1916.
Moist forest or thickets, 500-2,150 m.; Alta Verapaz; Santa
Rosa; Escuintla; Sacatepequez; Solola (type from Volcan de
Atitlan, E. W. D. Holway 187). Known only from Guatemala.
96 FIELDIANA: BOTANY, VOLUME 24
Eupatorium schultzii f. velutipes Rob. Proc. Am. Acad. 51:
536. 1916.
Moist or wet thickets or mixed forest, 1,000-2,800 m.; Jutiapa;
Santa Rosa; Escuintla; Guatemala; Sacatepequez; Chimaltenango;
Retalhuleu; Quezaltenango. Mexico; El Salvador.
This is by far the commonest form of the species in Guatemala.
Eupatorium scoparioides L. Wms. Fieldiana, Bot. 36: 102.
1975.
Open pine savanas; known only from the type, Molina 15595,
from near Poptun, Peten. Endemic.
Small, profusely branched shrubs to about 1 m. tall, the multistriate branches
slender, terete, densely spreading puberulent, the hairs sometimes minutely dendroid
toward the apices; leaves petiolate, triangular-ovate, acute or acuminate, truncate to
subcordate at the base, trinerved from the base of the blade, glabrous above or nearly
so, puberulent on the nerves below and densely punctate-glandular over the entire
surface, blades 1.5-3 cm. long and 1-2.3 cm. broad at the base, the petioles slender,
puberulent, mostly 5-6 mm. long; inflorescences spreading puberulent, terminal and
axillary, spicate or spicate-paniculate, bracteate, to about 8 cm. long, the heads in
small cymules, pedicellate; heads 3-4 mm. long, with about 5 florets; involucre bi- or
triseriate; phyllaries 10-12, prominently glandular-dotted, outer phyllaries linear-
lanceolate to ovate, acute, mostly less than 1 mm. long, the inner phyllaries linear-
oblong, truncate or rarely acute and somewhat expanded at the apices, mostly about
2 mm. long; corolla white, cylindric but subcampanulate at the throat, about 1.5-2
mm. long, the lobes minute, linear-lanceolate, about 0.2 mm. long; styles exserted
about 1 mm. at anthesis; achenes 5-ridged, strigillose, black at maturity, about 1-1.5
mm. long; pappus of 25-30 bristles, about 1.5 mm. long.
This is one of the most distinctive species of Eupatorium in
Central America, due to the very small flowers and inflorescence,
the punctate-glandular undersurface of the leaves, and the
punctate-glandular phyllaries. The occasionally dendroid hairs on
the stems are most unusual.
Eupatorium semialatum Benth. PL Hartw. 76. 1840; L. Wms.
Fieldiana, Bot. 36: 103. 1974. E. plethadenium Standl. & Steyerm.
Field Mus. Bot. 23: 186. 1944. Ageratina plethadenia King &
Robinson, Phytologia 24: 95. 1972. Bacche, baq ce' or q'eqci kay
Bacche, (Coban, Quecchi); barretillo; chicajol (Huehuetenango).
Thickets or in mixed forest areas, most often in oak-pine forest
but sometimes in Cupressus forest, 1,000-3,300 m.; Alta Verapaz;
Baja Verapaz; El Progreso; Zacapa; Chiquimula; Jalapa; Guate-
mala; Chimaltenango (type of E. plethadenium, Standley 61100);
Solola; El Quiche; Huehuetenango; Quezaltenango (type of E.
WILLIAMS: FLORA OF GUATEMALA 97
semialatum from Zunil, Hartweg 532); San Marcos. Mexico; El
Salvador to Costa Rica.
A shrub or a small, weak tree, 1.5-6 m. tall, the branches subterete, brownish or
fuscous, densely pubescent, tomentulose or short villosulous; leaves thick, on short
petioles, oblong to lance-oblong, mostly 4-9 cm. long, acute or acuminate, acute to
attenuate at the base, often or usually strongly and broadly revolute near the base,
penninerved, remotely crenate-dentate or sometimes very coarsely salient-dentate,
deep green above, puberulent or almost glabrous, densely punctate, often lustrous,
paler beneath, sparsely puberulent or glabrate, densely puberulent or villosulous on
the costa and nerves, very densely and finely punctate; corymbs small or large,
convex, fastigiately branched, usually very dense, the branches and pedicels densely
puberulent or tomentulose with brownish hairs; heads numerous, pale pink, 6-7 mm.
long, 4-8-flowered; phyllaries subequal, scarcely half as long as the florets, a few
short ones sometimes present at the base, obtuse, minutely gland-dotted and
tomentulose; corollas glabrous; achenes slender, black, sparsely puberulent; pappus
white or sometimes tinged with pink.
E. semialatum is a very common shrub or small tree in many
regions, especially in moist or wet forest of pine and oak. It is a
rather showy plant and a very fragrant one when in flower. The
very ample Guatemalan material is remarkably uniform in foliage
and pubescence characters. One conspicuous variant has been found
in Guatemala, represented by three collections from Totonicapan
and El Progreso. In this the leaves are very coarsely salient-dentate,
in outline suggesting those of some species of Lycopus of the United
States. There are intergrading specimens from other regions, and
this form probably is scarcely worthy of special designation. The
foliage is said to be bitter, and it is reported to be used in
Guatemala in domestic medicine, particularly in Alta Verapaz, for
the treatment of intestinal disturbances.
Most material of this species has been determined as E.
ligustrinum DC. in years past. The author believes that that name
represents a north Mexican species quite different from ours.
Eupatorium sexangulare (Klatt) Rob. Proc. Am. Acad. 43:
35. 1907. Piptocarpha sexangularis Klatt, Leopoldina Bot. Beibl. 1.
1895.
Wet, mixed forest or tall thickets along streams, 1,400-1,600 m.;
Alta Verapaz (Rio Frio near Tactic); Baja Verapaz. Honduras;
Nicaragua; Costa Rica.
A coarse, glabrous shrub or herb, 1.5-2 m. high, the stems hollow, sharply 6-
angulate and sulcate; leaves large, on rather short, marginate petioles, chartaceous,
olive-green when dried, lustrous, ovate-lanceolate or lance-oblong, mostly 10-25 cm.
long, acute or acuminate, acute and long-decurrent at the base, penninerved,
obscurely sinuate-dentate or almost entire; heads white or purplish-white, 5-flowered,
98 FIELDIANA: BOTANY, VOLUME 24
7-8 mm. long, cylindric-campanulate, forming large, open, corymbiform panicles, the
individual heads sessile or nearly so and densely glomerate; phyllaries multiseriate,
subcoriaceous, stramineous, closely imbricate, the inner ones caducous, ovate-
lanceolate, obtuse, striate, glabrous, the outer ones short, ovate; corollas glabrous;
achenes 3 mm. long, glabrous; pappus stiff, yellowish white.
Principally a species of the high, wet forests, scarce in
Guatemala but abundant in Honduras. A similar species with
triplinerved leaves from Peten is E. molinae.
Eupatorium sinclairii Benth. in Oersted, Vid. Medd. Kjoeben-
havn 1852: 79. 1853. Fleishmannia sinclairii King & Robinson,
Phytologia 19: 206. 1970.
Wet to dry thickets, shaded banks, often in rocky places,
sometimes in open fields or along the borders of streams, 2,200 m. or
less; Peten; El Progreso; Santa Rosa; Escuintla; Sacatepequez;
Solola; Retalhuleu; Quezaltenango. Costa Rica; Panama; probably
ranging more widely.
Plants herbaceous, slender, erect, usually much branched, annual or perhaps
sometimes perennial, usually finely puberulent almost throughout, sometimes short-
pilose, the stems terete; leaves membranaceous, on long, slender petioles, opposite,
deltoid-ovate or rhombic-ovate, mostly 2-5 cm. long, acuminate, rounded to broadly
cuneate at the base, 3-nerved, finely or coarsely crenate, green, finely puberulent or
glabrate; heads many-flowered, usually purple or lavender, 3 mm. high, forming lax,
leafy panicles, on long, slender, minutely puberulent pedicels; involucre campanulate;
phyllaries very unequal, the inner ones linear, greenish, acute or acuminate, costulate,
glabrous or nearly so, the outer ones short, lanceolate, acuminate; corollas glabrous,
slender-tubular, equaling the involucre; achenes very short, black, glabrous; pappus
white, equaling the corolla.
A close relative of E. microstemon Cass. but with very small
heads, perhaps not really distinct.
Eupatorium sodali L. Wms. Fieldiana, Bot. 36: 103. 1975.
Piqueria standleyi Rob. Contr. Gray Herb. 104: 4. 1934, not
Eupatorium standleyi Rob. Koanophyllon standleyi King &
Robinson, Phytologia 22: 151. 1971. Figure 21.
Dry to wet thickets or rather open forest, often in pine-oak
forest, 200-1,650 m.; Jalapa; Jutiapa; Santa Rosa; Escuintla;
Sacatepequez; Suchitepequez. El Salvador, the type from Sierra de
Apaneca.
A slender shrub 1.5-3 m. high, usually much branched, the stems striate, densely
and minutely appressed-puberulent; leaves on slender petioles 1.5-3 cm. long, ovate-
triangular, mostly 6-12 cm. long, narrowly attenuate-acuminate or caudate-
acuminate, broadly cuneate to truncate or shallowly cordate at the base, often
subhastate, coarsely and irregularly dentate or crenate-dentate, very thin, 3-nerved,
WILLIAMS: FLORA OF GUATEMALA 99
green and almost glabrous above, sparsely and minutely puberulent beneath; heads
white, 3-5-flowered, 4.5 mm. long, sessile or nearly so, crowded into dense cymules
about 1 cm. in diameter or in somewhat more elongated, dense spikes, these forming
leafy panicles; involucre about 4 mm. long; inner phyllaries about 6-7, linear-
lanceolate, acute, puberulent, subequal, the 2-3 outer phyllaries much smaller;
achenes about 2 mm. long, minutely puberulent; pappus present or none.
The present species illustrates the artificiality of the clas-
sification among some Compositae and more especially of the tribe
Euaptorieae. This species was originally placed with quite unlike
plants in the genus Piqueria by one of the most competent students
of American Compositae. The deciding factor in placing it there
undoubtedly was the lack of a pappus, but pappus bearing
specimens are now known and the species fits quite well into
Eupatorium, although it does disrupt in a minor way the pappus
character ascribed to Eupatorium.
Eupatorium solidaginoides HBK. Nov. Gen. & Sp. 4: 126.
1820. E. filicaule Schultz Bip. ex Gray, Proc. Am. Acad. 21: 384.
1886. E. decussatum Klatt, Bull. Soc. Bot. Belg. 35: 295. 1896.
Koanophyllon solidaginoides King & Robinson, Phytologia 22: 151.
1971. Sasajan (fide Aguilar), saqi lokab q'eqi lokab, (Alta Verapaz,
Quicchi).
Moist or rather dry, mixed forest or thickets, frequently in oak
forest, 300-1,850 m.; Peten; Santa Rosa; Escuintla; Guatemala;
Sacatepequez; Chimaltenango; El Quiche. Southern Mexico; British
Honduras; Honduras; El Salvador; Costa Rica; northwestern South
America.
A slender shrub, 1.5-4 m. high, often with recurved branches; leaves very thin,
dull green when dried, on long, slender petioles, rather narrowly triangular-ovate, 5-
12 cm. long, narrowly long-acuminate, usually openly and shallowly cordate at the
base, often somewhat hastate- lobate, with rounded lobes, dentate or subentire,
sparsely short-villosulous above or almost wholly glabrous, puberulent beneath or
somewhat tomentulose on the nerves; heads white or greenish white, 10-15-flowered,
about 5 mm. long, on short slender pedicels, in small rounded glomerules or in
elongate narrow racemes, forming a large, open, leafy, terminal panicle; phyllaries
greenish, lanceolate, acute, puberulent, unequal, laxly imbricate in few series; florets
equaling or scarcely exceeding the phyllaries; achenes small, blackish, minutely
hispidulous or puberulent on the sides as well as on the angles.
Robinson (in Standley, Trees and Shrubs of Mexico] describes
this species as a "calciphile," but in Guatemala it very rarely, if
ever, occurs on calcareous soils.
Eupatorium sorensenii (King & Robinson) L. Wms. Field-
100 FIELDIANA: BOTANY, VOLUME 24
iana, Bot. 36: 104. 1975. Koanophyllon sorensenii King & Robinson,
Phytologia 23: 395. 1972.
In pine woods or thickets, known only from the type, Sorensen
7129, from "Pine Ridge," British Honduras.
Low shrubs to 0.5 m. tall, the stems densely sordid-puberulent; leaves lanceolate,
acuminate, base acute, mostly 5-7-penninerved, glabrous and glandular above,
densely short-puberulent and glandular-dotted below, the blade to 10 cm. long and
3.5 cm. broad, petioles short, to about 1 cm. long, sordid-puberulent; inflorescences
axillary in the upper leaves, paniculate, hardly longer than the subtending leaves;
heads 7-8 mm. long; phyllaries about 3-seriate, the outer phyllaries mostly about 2
mm. long, narrowly ovate, puberulent dorsally, the inner phyllaries about 4 mm. long,
narrowly ovate to lanceolate, acute, puberulent at the tip or glabrous; corollas
tubular, about 3 mm. long; achenes prismatic, 5-ridged, puberulent on the angles or
glabrous; pappus about 3.5 mm. long.
Perhaps too closely related to Eupatorium hypomalacum Rob.,
which is known from the eastern slope of Guatemala only by the
type collection made three quarters of a century ago.
Eupatorium tomentellum Schrad. Ind. Sem. Hort. Goett.
1833: 3, t. 3. 1833. Ageratina tomentella King & Robinson,
Phytologia 19: 227. 1970.
Dry, brushy, usually rocky hillsides, 900-2,100 m.; El Quiche;
Huehuetenango. Southern Mexico.
A stout shrub of 1-2 m., often much branched, the branches terete or obtuse-
tetragonous, closely tomentulose, with yellowish or pale brownish tomentum; leaves
thick, on rather long, stout petioles, broadly deltoid-ovate or rounded-deltoid, often
fully as broad as long, 2-5 cm. long, rounded or very obtuse at the apex, shallowly or
rather deeply cordate at the base, with rounded basal lobes, 3-5-plinerved from near
the base, crenulate or almost entire, grayish green and puberulent above, yellowish
tomentulose beneath and conspicuously glandular; inflorescence corymbose, small,
very dense, convex, leafy-bracted at the base, 4-6 cm. broad; heads very numerous,
pink or purplish, 7 mm. long, with 5-7 flowers, short-pedicellate; phyllaries narrowly
oblong, acute, subequal, few, glandular-puberulent; corollas twice as long as the
involucre; achenes blackish, scabrous on the angles.
In Guatemala this shrub is confined to the lower foothills of
the Cuchumantanes, where it grows in the most exposed and dryest
places, frequently where there is no soil except in rock crevices; a
closely allied species, E. ovillum, is on the high mountain ridges and
tops.
Eupatorium tuerckheimii Klatt, Leopoldina 20: 95. 1884.
Neobartlettia tuerckheimii King & Robinson, Phytologia 21: 297.
1971. Bartlettina tuerckheimii King & Robinson, Phytologia 22: 162.
1971.
WILLIAMS: FLORA OF GUATEMALA 101
Moist or wet, dense, mixed forest or often in pine forest, 1,250-
3,000 m.; Alta Verapaz (type from Coban, Tuerckheim 77); El
Progreso; Zacapa; Jalapa; El Quiche. Mexico (Chiapas); Honduras.
A shrub of 1-3 m., sometimes a tree of 5-6 m., erect or weak and subscandent,
glabrous throughout, the branches rather stout, brownish, striate; leaves opposite, on
long, slender petioles, lanceolate or lance-oblong, 22 cm. long and 7 cm. broad or
smaller, narrowly long-attenuate, acute or attenuate at the base, cuspidate-
denticulate, usually remotely and inconspicuously so, with minute, digitaliform
processes, penninerved, the veins translucent under a strong light; heads pale purple
or lilac, about 50-flowered, 8 mm. long and usually broader, broadly campanulate,
long-pedicellate, forming small or large, usually dense, teriminal, rounded, somewhat
umbelliform corymbs; phyllaries unequal, lanceolate, strongly imbricate, acute;
achenes glabrous; pappus white or nearly so.
This species is very closely related to E. pinabetense.
Eupatorium tunii L. Wms. Fieldiana, Bot. 36: 106. 1975.
Forests or thickets at about 400 m. or less; Peten (type, Tim
1306); British Honduras (Schipp 137). Mexico.
Glabrous vines to 2.5 cm. in diameter, the branches opposite; leaves lanceolate-
ovate to ovate, acute or very shortly acuminate, glabrous, triplinerved, with the
nerves beginning 4-5 mm. above the base of the blade, the blade 4.5-9 cm. long and 2-
5 cm. broad, petioles slender, mostly about 1 cm. long; inflorescence axillary or
terminal, usually very compact cymules, 2-3 cm. in diameter and each containing up
to about 50 heads; heads sessile or nearly so, somewhat more lax in fruit, bearing 4-6
florets, 7-11 mm. long and 2-2.5 mm. in diameter; involucre 4-6-seriate, phyllaries
very unequal, noticeably puberulent at the apices, outermost phyllaries ovate and
about 1 mm. long, the innermost lanceolate to narrowly elliptic, acute and 5-6 mm.
long; corolla cylindric and slightly expanded upward, about 6 mm. long, the lobes
lanceolate, acute, about 0.5 mm. long; achenes obscurely hirtellous on the angles,
becoming glabrous, brownish black, 5-ridged, about 2.5-3 mm. long; pappus 5-6 mm.
long, equaling the corollas, very obscurely barbellate or smooth.
Named for Rolando Tun Ortiz, who has made some 2,000
collections from the department of Peten, thereby greatly increas-
ing our knowledge of the flora of that remote part of Guatemala.
Eupatorium ultraisthmium McVaugh, Contr. Univ. Mich.
Herb. 9: 388. 1972. Eupatoriastrum nelsonii Greenm. Proc. Am.
Acad. 39: 93. 1903, not Eupatorium nelsonii Rob. 1900. Eu-
patoriastrum nelsonii var. cardiophyllum Rob. & Greenm. Proc.
Am. Acad. 41: 277. 1905 (type from Los Pinos, Chiapas).
Dept. Guatemala, without locality, about 1,500 m., Aguilar 577.
Southern Mexico; El Salvador.
A tall herb, or perhaps sometimes suffrutescent, the stems thinly or densely
villosulous or glabrate, pale, striate, subterete; leaves short -petiolate, membrana-
102 FIELDIANA: BOTANY, VOLUME 24
ceous, broadly ovate or rounded-ovate, 10-20 cm. long and 7-13 cm. broad or larger,
acute or rather abruptly short-acuminate, broadly rounded or subcordate at the base,
denticulate or sinuate-dentate, 5-7-nerved from the base, thinly villosulous or
glabrate, the veins often prominent and closely reticulate; heads 1.5-2 cm. broad,
subglobose in age; inflorescences lateral, mostly of 3 heads, the upper ones few,
forming an open panicle, the individual heads on stiff, spreading peduncles; phyllaries
lance-oblong, acute, pubescent; pales narrow, slightly broadened toward the apex;
achenes 1.5-2 mm. long, fuscous, hirtellous on the angles; pappus dirty white, about
equaling the corollas.
Known in El Salvador as "sunsumpate" and "raiz barbona."
The roots are said to have an agreeable odor and a somewhat biting
taste, and are employed in that country in domestic medicine.
Eupatorium viscidipes Rob. Proc. Am. Acad. 36: 484. 1901.
Fleishmannia viscidipes King & Robinson, Phytologia 19: 206. 1970.
Usually in wet forest, sometimes cloud forest, wet thickets or in
oak-pine forest, 1,700-2,900 m.; Baja Verapaz; Solola; Huehue-
tenango; El Quiche (type from Chicaman, Heyde & Lux 3397);
^Totonicapan; Quezaltenango; San Marcos. Endemic.
Plants slender, erect or ascending, herbaceous, perennial or perhaps annual, 1 m.
high or less, usually much branched, the stems terete, often purplish, minutely
puberulent and very viscid; leaves opposite, slender-petiolate, deltoid-ovate or
rhombic-ovate, 2-5 cm. long, narrowly long-acuminate, subtruncate to broadly
cuneate at the base, 3-nerved, crenate-serrate, minutely puberulent above, slightly
paler and punctate beneath; heads numerous, 4 mm. long, white, about 18-flowered,
forming large, lax, leafy-bracted, corymbiform panicles, the pedicels very slender, 3-13
mm. long, viscid; involucre turbinate-campanulate; phyllaries several-seriate, very
unequal, stramineous or pale greenish, the outer ovate-oblong, the inner oblong,
subscarious, striate, obtuse or rounded at the apex, glabrous or slightly glandular,
viscid; corolla glabrous, 2.3 mm. long; achenes black, columnar, hispidulous on the
angles; pappus white, slightly shorter than the corolla, sometimes wanting or present
in the same head.
A member of a small group of species related to Eupatorium
microstemon Cass. and not easily distinguished from it.
Eupatorium vitalbae DC. Prodr. 5: 163. 1836. Heterocondylus
vitalbae King & Robinson, Phytologia 24: 393. 1972, as vitalbis.
A vine over other shrubs or sometimes a liana much resembling
Clematis, from wet, lowland thickets to mixed, oak-pine forests,
mostly near sea level but occasionally up to 1,000 m. Not known in
Guatemala but to be expected in Izabal and Peten. Honduras;
Nicaragua; Costa Rica; Panama; over much of the lowland tropics
of South America; perhaps Mexico.
Suffrutescent vines or lianas, the branches opposite, terete or obscurely
quadrangular, sordid-puberulent, becoming glabrous; leaves lanceolate-ovate to
WILLIAMS: FLORA OF GUATEMALA 103
lanceolate, acuminate at the apex, the base rounded or subtruncate, serrate,
triplinerved from the base or slightly above, sordid-puberulent but soon glabrous, 3-12
cm. long, the petioles 1.5 cm. long or less, glabrescent; inflorescence terminal in the
axils of reduced leaves, often a cymose panicle 15 cm. broad; heads large, 1.5 cm. long
or less, with about 50 flowers; phyllaries in three series, the outer phyllaries ovate or
lanceolate-ovate, acute, sordid-puberulent, 5-8 mm. long, the inner ones linear to
linear-oblong, acute or the apex crisped and ciliolate, as long as the pappus; flowers
white or pinkish, at anthesis about as long as the pappus; pappus barbellate, white.
A widely distributed species in the lowland tropics of South
America.
Eupatorium zunilanum Standl. & Steyerm. Field Mus. Bot.
23: 191. 1944. Ageratina zunilana King & Robinson, Phytologia 19:
218. 1970.
Moist or wet forest, at least sometimes with Abies, 2,500-3,600
m.; endemic; Quezaltenango (type from Volcan de Zunil, Steyer-
mark 34744; also on Volcan de Santo Tomas).
A shrub, 1 m. high or less, or perhaps only suffrutescent, sparsely and laxly
branched, the branches terete, greenish, densely villous with long, slender, spreading,
gland-tipped hairs; leaves opposite, membranaceous, on slender petioles 2.5-6.5 cm.
long, deltoid or ovate-deltoid, 4-7.5 cm. long, 3-6 cm. broad, acute or acuminate,
truncate at the base, 3-nerved, coarsely and almost evenly crenate, sparsely pilose or
villosulous about, sparsely villosulous beneath over the whole surface; inflorescence
corymbiform, sparsely and laxly branched, about 12 cm. high and broad, the branches
elongate, the heads long-pedicellate, laxly cymose, the pedicels mostly 1-2 cm. long,
rather densely glandular-pilose; heads few, white, 10-12 mm. long, campanulate, 30-
35-flowered; phyllaries subequal, pale green, broadly linear, acute, scarious above,
densely glandular-pubescent, sometimes purplish, costulate; corollas glabrous, 5-6
mm. long, scarcely longer than the involucre; achenes black, rather stout, very
minutely puberulent or almost glabrous; pappus bristles few, white, minutely
scaberulous, deciduous.
EXCLUDED SPECIES
EUPATORIUM GUATEMALENSE Regel, Linnaea 24: 231. 1851.
Based on plants said to have come from Guatemala.
Section Subimbricata. Suffruticose, erect, paniculately branched, the younger
branches hirsutulous; leaves opposite, long-petiolate, deltoid-ovate, 3-nerved, acu-
minate, entire at the base and apex, crenate-dentate along the middle, the teeth
obtuse; corymbs lax, pedunculate, axillary and terminal, bearing few heads;
peduncles slender, hirtellous, bearing 2-3 heads; heads short-pedicellate, ovate-oblong,
about 30-flowered; phyllaries laxly imbricate, oblong, glabrous, scarious at the apex,
ciliate, the inner ones obtuse, the outer ones smaller, acute; corollas white.
In habit and leaves similar to E. schiedeanum, but differing in
the hirsute young branches, more lax inflorescence, about 30-
104 FIELDIANA: BOTANY, VOLUME 24
flowered heads, and white corollas. We have seen no representation
of this species, which is not included in the key to species. The
description seems to be a rather good and ample one, but we have
been unable to decide to what plant it should apply.
EUPATORIUM LIGUSTRINUM DC. Prodr. 5: 181. 1836. Ageratina
ligustrina King & Robinson, Phytologia 19: 233. 1970.
This name has been used on a great mass of Central American
(and Mexican) collections of Eupatorium. However, the type of E.
ligustrinum seems to represent a different species of north Mexico.
The name E. semialatum Benth. is used in this flora for most of the
specimens formerly referred to E. ligustrinum.
EUPATORIUM MACROCEPHALUM Lessing, Linnaea 5: 136. 1830.
Campuloclinium macrocephakim DC. Prodr. 5: 137. 1836; King &
Robinson, Phytologia 24: 172. 1972.
King & Robinson give the range of this species as Mexico,
Honduras, Colombia, Bolivia, and Paraguay. If the range given is
correct, the species should be found in Guatemala, but we have seen
no specimens from Central America. Although the species is
reported from Mexico, we have not seen specimens from there
either.
EUPATORIUM VERNALE Vatke & Kurtz, Ind. Sem. Hort. Berol.
1871: App. 2. 1871; Gartenfl. 1873: 36, t. 750. 1873; Robinson in
Standley, Contr. U. S. Nat. Herb. 23: 1461. 1926.
This plant was grown out in the garden in Berlin from seed
thought to have originated in Mexico. It has not been identified
with any species known to occur there. There apparently was no
type in the Berlin Herbarium for we have no photograph and if
Macbride missed it, which is not likely, then it may have been
destroyed during World War II.
The name has sometimes been identified, from the characters,
with the plant here described as Eupatorium monticola L. Wms.,
but the characters of the phyllaries shown, nearly as long as the
flower heads, indicate that it must have been some other species.
The illustration shows leaves with both penninerves and plinerves, a
condition not likely to occur in a single species.
ISOCARPHA R. Brown
Annual herbs, perhaps sometimes more enduring, divaricately branched, usually
softly pubescent; leaves opposite or the upper ones alternate, sessile or petiolate, the
WILLIAMS: FLORA OF GUATEMALA 105
margins entire or dentate; heads homogamous, discoid, in fruit ovoid-conic, on long
or short peduncles, sometimes solitary, sometimes several, sessile or short-pedicellate
in small corymbs at the ends of the peduncles; flowers numerous, all hermaphrodite
and fertile; involucres short; phyllaries 2-3-seriate, subequal or the outer ones
shorter; receptacle conic (in ours), the pales complicate, embracing the flowers;
corollas regular, tubular, white or greenish white, the limb 5-dentate; anthers entire
and truncate at the base; style branches slender, ending in long, subulate, pubescent
appendages; achenes glabrous, 4-5-angulate, truncate; pappus none.
About five species, in tropical America, with only two in
Mexico and Central America. This genus has usually been placed in
the tribe Heliantheae, but Lessing, in 1830, placed it in the tribe
Eupatorieae, where it seems to belong.
Leaf blades ovate to rhombic-ovate, not triplinerved, at least the lower ones
petiolate, the petioles more or less winged, dilated and auriculate at the base;
heads always pedunculate or pedicellate, never sessile I. atripUcifolia.
Leaf blades lanceolate or linear-lanceolate, conspicuously triplinerved, sessile or
subsessile; heads often sessile in dense clusters of 2 or 3 /. oppositifolia.
Isocarpha atriplicifolia (L.) R. Br. Trans. Linn. Soc. 12: 110.
1816. Bidens atriplicifolia L. Cent. PL 2: 30. 1756.
Damp thickets, rocky river banks, or on saline flats, sea level to
600 m.; Jutiapa; Santa Rosa. Southern Mexico; Honduras;
Nicaragua; Costa Rica.
Erect annuals, 30-60 cm. high, much branched, sparsely or densely puberulent or
pubescent throughout, the stems pale, subterete, the branches mostly spreading, very
leafy; lower leaves on winged petioles, the blades thin, broadly rhombic-ovate, mostly
2.5-6 cm. long, the margins rather coarsely sinuate-dentate, obtuse or acute, broadly
cuneate at the base, sometimes glabrate on the upper surface, more or less pubescent
beneath; upper leaves much smaller, petiolate or sessile, the petioles, when present,
broadly winged, abruptly dilated at the base into 2 conspicuous auricles; heads
always pedicellate or pedunculate, never sessile, 5-7 mm. high, broadly ovoid; pales
greenish and conspicuous; phyllaries numerous, linear-lanceolate, attenuate, hispi-
dulous; corollas white, about 2 mm. long; achenes black, about 1 mm. long.
Isocarpha oppositifolia (L.) R. Br. Trans. Linn. Soc. 12: 110.
1818. Santolina oppositifolia L. Syst. ed. 10. 1207. 1759. /. echioides
Less. Linnaea 5: 141. 1830. Figure 22.
Damp or dry, often brushy plains or hillsides, frequently in
rocky places, near sea level to 1,100 m.; Baja Verapaz; Chiquimula;
Guatemala; Jalapa; Zacapa. Texas; Mexico; Honduras and El
Salvador to Panama; West Indies; northern South America.
Erect herbs to about 1 m. high, simple or sometimes much branched, usually
abundantly hirsute or hispidulous, the whole plant grayish, the stems very slender,
the internodes elongated; leaves sessile or subsessile, the blades lanceolate or linear-
lanceolate, mostly 3-10 cm. long, 1-2.5 cm. wide, the margins entire or nearly so,
106 FIELDIANA: BOTANY, VOLUME 24
acuminate, acute at the base, conspicuously triplinerved, often scabrous; heads
turbinate, 6-10 mm. high, solitary at the ends of long, naked peduncles or often
clustered at the ends of long peduncles, there sessile or pedicellate; phyllaries
biseriate, obovate-oblong, acute or obtuse, hispidulous and glandular, pale, 3-4 mm.
long; pales oblong-spathulate, mucronulate, ciliate and glandular, about 3.5 mm.
long; corollas 2-2.5 mm. long; achenes black, oblong, about 5-angulate, narrowed at
the base, 1.5-2 mm. long.
Known in Yucatan by the names "chahancan," "habancan,"
"extocaban amargo," and "tocabal amargo" (fide Standley).
MACVAUGHIELLA King & Robinson
Perennial herbs or shrubs, pubescent or almost glabrous, branched; leaves
opposite, broad, thin, dentate, 3-nerved or triplinerved; heads small, white,
homogamous, many-flowered, densely crowded in small, terminal corymbs; involucre
cylindric; phyllaries about 2-seriate, narrow, rather rigid, subequal or the outermost
shorter; receptacle convex or conic; corollas regular, the tube slender, the limb
narrow-campanulate, 5-fid; anthers appendaged at the apex, obtuse and entire at the
base; style branches rather long, subobtuse; achenes narrow, black, pubescent or
glabrous, compressed, not costate; pappus bristles 2, inserted on a cartilaginous
annulus, without intermediate paleae.
One other species is known. It is found in Mexico.
Macvaughiella standleyi (Steyerm.) King & Robinson, Sida
3: 282. 1968; L. Wms. Fieldiana, Bot. 36: 107. 1975. Schaetzellia
standleyi Steyerm. Field Mus. Bot. 23: 107. 1944. Figure 23.
Dry, rocky slopes, 30-1,400 m.; Chiquimula; Jalapa; Jutiapa
(type from Lago de Retama, Steyermark 32041). El Salvador;
Honduras.
Plants erect, somewhat woody, 2 m. high or less, branched, the young branches
densely tomentulose or velutinous-pilosulous; leaves on petioles 6-35 mm. long, thin,
the blades broadly deltoid-ovate, 1.5-8 cm. long, 1-5 cm. broad, 3-nerved, obtuse or
acute to acuminate, cuneate or broadly rounded at the base, coarsely crenate,
minutely and densely grayish velutinous on both surfaces to subglabrous, densely
puncticulate beneath; inflorescences small and very dense, the heads short-
pedicellate, 16-25-flowered; involucre turbinate-cylindric, 3.5-4 mm. high; phyllaries
about 10, grayish green, the outer ones slightly purplish at the apex, spathulate-
oblanceolate, mucronate, conspicuously 3-nerved, glandular-pilosulous; corollas
white, almost 3 mm. long, glandular; awns of the pappus 2 and 2.5 mm. long,
barbellate; achenes black, linear-oblong, upwardly hispidulous on the margins, 2 mm.
long.
This species may not be distinct from M. mexicana, known
from Vera Cruz. Specimens from near sea level along the coast of El
Salvador are more luxuriant than those that we have seen from
middle elevations. The species is often abundant in the open, pine-
oak forest regions of central Honduras.
WILLIAMS: FLORA OF GUATEMALA 107
MIKANIA Willdenow
Reference: B. L. Robinson & J. M. Greenman, Synopsis of the
Mexican and Central American species of the genus Mikania, Proc.
Am. Acad. 32: 10-13. 1896.
Herbs or shrubs, scandent, glabrous or pubescent; leaves opposite, generally
petiolate, usually broad, often cordate at the base, mostly palmate- nerved, entire or
dentate; heads small, homogamous, 4-flowered, spicate, racemose, corymbose, or
cymose and paniculate, usually white; phyllaries 4, equal, commonly narrowly
oblong, sometimes with a much smaller outermost one (bractlet); receptacle small,
naked, glabrous; corollas tubular, the throat gradually or abruptly enlarged,
turbinate or campanulate, 5-dentate; anthers rounded to emarginate at the base,
with an ovate or oblong apical appendage; style branches elongate, clavellate or
filiform; achenes 5-angulate, prismatic or narrowed below; pappus bristles capillary,
often slightly scaberulous or barbellate, never truly plumose, free or very slightly
connate at the base, white to fulvous.
Species about 275, one in the Old World tropics, the others
American, chiefly in the tropics and almost half of them Brazilian.
One species extends into temperate, eastern North America. A few
others are known from Central America. Mikania Willdenow is a
conserved name.
Inflorescence spicate or racemose.
Heads sessile.
Branches winged; nerves of the leaves arising at or near the base of the blade.
M. pterocaula.
Branches not winged; nerves, at least the principal ones, arising far above the
base of the blade M. leiostachya.
Heads pedicellate.
Achenes glabrous; rachis of the inflorescence rather densely pilose.
M. houstoniana.
Achenes glandular-puberulent; rachis of the inflorescence glabrous or nearly so.
M. houstoniana var. guatemalensis.
Inflorescence of cymose panicles, never spicate or racemose.
Leaves acute to rounded at the base.
Heads in globose glomerules; leaf blades abruptly contracted at the base, not
decurrent M. aromatica.
Heads in cymes; leaves long-decurrent at the base M. guaco.
Leaves conspicuously cordate at the base, or at least subcordate.
Heads small, scarcely 5mm. long M. micrantha.
Heads all or mostly 8-12 mm. long.
Stems densely setose-pilose with long, spreading, brownish hairs.
M. pyramidata.
Stems glabrous or pubescent, but never setose-pilose.
Involucres densely or sparsely pilose or villous.
Leaves deeply cordate at the base; pubescence white or gray.
M. cordifolia.
Leaves subcordate or broadly rounded at the base; pubescence brown.
M. petrina.
108 FIELDIANA: BOTANY, VOLUME 24
Involucres glabrous or practically so.
Leaves somewhat hastat^-angulate; phyllaries obtuse M. vitifolia.
Leaves not at all angulate.
Leaves subcoriaceous; heads sessile or nearly so M. concinna.
Leaves membranaceous; heads mostly pedicellate M. cordifolia.
Mikania aromatica Oersted, Overs. Dansk. Vidensk. Selsk.
Forh. 10. 1863. Willughbaea globosa Coulter, Hot. Gaz. 20: 46. 1895.
Mikania globosa Coulter, /. c., in synon.
Mostly in dense, wet, mixed forest, 500-2,400 m.; Alta Verapaz;
Zacapa; Santa Rosa (type from Santa Rosa, Heyde & Lux 3430);
Suchitepequez. Southern Mexico; El Salvador; Honduras; Costa
Rica(?).
A large vine, glabrous throughout or nearly so, the stems terete; leaves slender-
petiolate, coriaceous or subcoriaceous, often lustrous, rounded-ovate to lance-ovate,
mostly 10-16 cm. long, acuminate or caudate-acuminate, obtuse to broadly rounded
at the base or rarely subcordate, 5-plinerved, the inner nerves arising far above the
base, entire or obscurely and remotely undulate-denticulate, slightly paler beneath;
heads greenish white, in globose glomerules, arranged in lateral or terminal panicles;
phyllaries 3-3.5 mm. long, truncate at the apex, enveloping the flowers, not imbricate,
sparsely puberulent, hardly longer than the achenes; achenes glabrous or nearly so;
pappus dirty white.
Closely related to this and perhaps not distinct, is M. tonduzii
Robinson, described from Costa Rica.
Mikania concinna Standl. & Steyerm. Field Mus. Bot. 23: 260.
1947.
Known only from the type, Zacapa, Sierra de las Minas,
climbing over mossy banks, middle and upper, southern slopes of
Volcan Gemelos, about 3,000 m., Steyermark 43292.
A glabrous, herbaceous vine, the stems subterete, dark purple; leaves small,
coriaceous or subcoriaceous, somewhat lustrous, on slender petioles 1-2.5 cm. long,
deltoid-ovate or rounded-ovate, 2.5-5.5 cm. long, 2-4.5 cm. broad, acute or acuminate,
broadly and rather shallowly cordate at the base or subtruncate, green above, the
nerves prominulous, the veins obsolete, paler beneath, obscurely puncticulate, 5-
nerved, the veins obsolete, the margin entire or nearly so, usually with about 3
minute, repand teeth on each side; heads few, in fruit 12 mm. long, densely aggregate-
cymose, sessile or subsessile, the bracts subtending the heads ovate or elliptic, acute,
ciliolate, shorter than the involucre; phyllaries tinged with purple, 7-8 mm. long,
glabrous or glabrate, ciliate, oblong or broadly oblong, obtuse and apiculate, dirty
brown when dried, costate-nerved; corolla glabrous, shorter than the pappus; achenes
4.5-5 mm. long, sordid-olivaceous, glabrous; pappus bristles brownish, 5 mm. long.
Mikania cordifolia (L. f.) Willd. Sp. PL 3: 1746. 1804. Cacalia
cordifolia L. f. Suppl. PL 351. 1781. M. gonoclada DC. Prodr. 5: 199.
WILLIAMS: FLORA OF GUATEMALA 109
1836. M. huitzensis Standl. & Steyerm. Field Mus. Bot. 23: 260. 1947
(type from Huehuetenango, Steyermark 48662).
Moist or wet thickets or mixed forest, 1,800 m. or less; Alta
Verapaz; Escuintla; Guatemala; Sacatepequez; Chimaltenango;
Suchitepequez; Huehuetenango; Quezaltenango; San Marcos. Mex-
ico; El Salvador and Honduras to Panama; tropical South America.
A small or large, herbaceous vine, the stems much branched, 6-angulate or
subterete, glabrous to grayish-villosulous; leaves long-petiolate, ovate or broadly
ovate, 5-15 cm. long, acute or acuminate, deeply cordate at the base, with rounded
basal lobes, membranaceous, subentire to undulate-dentate, glabrate or usually
rather densely short-villous, grayish, 3-nerved from the base of the blade; corymbs
terminal and lateral, convex, the pedicels 1-2 mm. long; phyllaries 6-8 mm. long,
acute, rather densely pubescent to glabrous; corollas greenish white or dirty white;
achenes glabrous, 3.5 mm. long; pappus dull white or in age rufescent.
A somewhat variable species and a wide ranging one, mostly at
middle elevations. It perhaps should include M. petrina, a species
distinguished principally by its abundant, sordid pubescence.
Mikania guaco Humb. & Bonpl. PI. Aequin. 2: 84, t. 105. 1809.
Wet, mixed forest, at or near sea level; Izabal. Southern
Mexico; Honduras to Panama; tropical South America.
A large, coarse, herbaceous vine, the stems terete, soon glabrate, often fistulose,
when young somewhat sordid-tomentulose; leaves slender-petiolate, ovate or
rhombic-ovate, 12-20 cm. long, usually long-acuminate, cuneate-decurrent at the
base, undulate and denticulate or subentire, 5-7-plinerved far above the base or
practically penninerved, thin, inconspicuously puberulent or scaberulous above,
tomentulose beneath along the nerves or glabrate; heads greenish white, about 1 cm.
long, mostly sessile, in large, convex or rounded corymbs, long-pedunculate in the
upper leaf axils and forming large, ovoid panicles; phyllaries oblong, thinly
puberulent or glabrate, rounded at the apex; achenes 3.3 mm. long, minutely
roughened; pappus bristles pale brownish or fulvous.
The specific name is derived from the vernacular name,
"guaco," used in many parts of tropical America for this and other
species of the genus. This name is given generally to plants
considered remedies or antidotes for the bites of poisonous snakes.
Molina says that the plants, stems as well as roots, are used on the
north coast of Honduras in fishing. The plant is called either
"guaco" or "pate."
Mikania houstoniana (L.) Robinson, Proc. Am. Acad. 42: 47.
1906. Eupatorium houstonianum L. Sp. PL 836. 1753. E. houstonis
L. Syst. Nat. ed. 10. 2: 1204. 1759. M. houstonis Willd. Sp. PL 3:
1742. 1804.
110 FIELDIANA: BOTANY, VOLUME 24
Moist or wet thickets, 250-2,000 m.; Alta Verapaz; Chiquimula;
Suchitepequez; Retalhuleu; Quezaltenango. Southern Mexico;
Honduras; Costa Rica; tropical South America.
A large, coarse, herbaceous vine, the stems terete, puberulent or sparsely pilose
with rather long, spreading hairs, in age glabrous or nearly so; leaves thin, on long,
slender petioles, very broadly ovate to ovate, mostly 7-13 cm. long but sometimes
larger, acuminate, rounded at the base, entire, 5-plinerved from near the base,
glabrous or practically so; inflorescence a large, compound racemose, panicle, the
pedicels scattered, 2-3 mm. long, puberulent; heads pale greenish yellow or greenish
white, 4-6 mm. long; phyllaries oblong, subacute, sparsely puberulent or almost
glabrous, 3-4.5 mm. long; achenes glabrous; pappus dirty white.
Mikania houstoniana var. guatemalensis (Standl. & Stey-
erm.) L. Wms. Field Mus. Bot. 38: 108. 1975. M. guatemalensis
Standl. & Steyerm. Field Mus. Bot. 23: 105. 1944.
Wet thickets or mixed forest, sometimes Manicaria swamps,
from near sea level to 600 m.; Alta Verapaz; Izabal (type from
Puerto Barrios, Deam 40). British Honduras; Honduras; possibly
Costa Rica.
Differs from the typical variety in having the inflorescence
glabrous, not puberulent; the achenes puberulent and somewhat
gland-dotted, not glabrous and gland-less.
The variety is a minor one.
Mikania leiostachya Benth. PI. Hartweg. 201. 1845.
Wet forest, at or little above sea level; British Honduras
(Middlesex, Schipp 513); to be expected in Izabal. Honduras;
Nicaragua; Costa Rica; Panama; northwestern South America.
A large, coarse vine, sometimes 9 m. long, almost glabrous throughout, perhaps
sometimes woody below, the stems as much as 5 cm. thick, terete; leaves short-
petiolate, ovate or lance-ovate, 6-20 cm. long, acuminate or long-acuminate, rounded
at the base, subcoriaceous, often lustrous, 5-plinerved, the inner nerves arising far
above the base, the veins usually prominent and reticulate on both surfaces,
somewhat puberulent beneath or in age glabrous; inflorescence a large panicle; heads
7 mm. long, greenish white, sessile, spicate; phyllaries ovate-oblong, obtuse, pubescent
or glabrate, 4 mm. long; achenes fuscous, lustrous, glabrous, 2 mm. long; pappus dirty
white.
Mikania micrantha HBK. Nov. Gen. & Sp. 4: 134. 1820. Name
de raton (fide Aguilar); bejuco llouizna (fide Aguilar).
Moist or wet thickets, often in second growth, frequently in
roadside hedges, 1,700 m. or less, mostly at less than 1,000 m.; Alta
Verapaz; Baja Verapaz; Izabal; Zacapa; Chiquimula; Jutiapa;
Santa Rosa; Escuintla; Guatemala; Sacatepequez; Suchitepequez;
WILLIAMS: FLORA OF GUATEMALA 111
Retalhuleu; Quezaltenango; Huehuetenango; probably in all the
lowland departments. Mexico; British Honduras to El Salvador and
Panama; West Indies; tropical South America.
A small or large, herbaceous vine, sparsely pubescent or almost glabrous, the
stems slender, terete, often much branched and interlaced; leaves slender-petiolate,
thin, pale green, broadly ovate or deltoid-ovate, mostly 5-13 cm. long, acuminate,
deeply cordate at the base and somewhat hastately or saggitately lobate, subentire to
coarsely crenate-undulate, 3-7-nerved from the base, almost glabrous or sparsely
pubescent beneath; panicles of compound cymes, terminal and lateral; heads dirty
white or greenish white, 4-5.5 mm. high, on filiform pedicels 5 mm. long or less;
phyllaries obovate-oblong, acute or short-acuminate, greenish white; achenes black,
1.7 mm. long, sparsely glandular-atomiferous; pappus white or yellowish.
A common, weedy plant in many parts of the Central American
lowlands. It is a variable plant both in the leaves and in the heads
and flowers.
Mikania petrina Standl. & Steyerm. Field Mus. Bot. 23: 261.
1947.
Known only from the type, Solola, in wet cloud forest near the
summit of Volcan de San Pedro, northern slopes toward Lago de
Atitlan, above San Pedro, 2,500-2,800 m. Steyermark 47257.
A scandent herb, the stems obscurely hexagonal or subterete, densely brown-
pilosulous; leaves rather small, on slender brown-pilosulous petioles 2-3 cm. long,
ovate or oblong-ovate, about 6 cm. long and 3.5 cm. broad, or often smaller,
acuminate, truncate at the base or subcordate, rather closely serrate-dentate,
scabrous or scaberulous on both surfaces, rough to the touch, brown-pilose beneath
when young, 5-nerved from the base, the veins evident beneath; fruiting heads 13
mm. long, numerous, cymose-corymbose, sessile, densely aggregate, the corymbs
rounded, about 5.5 cm. broad, bearing reduced leaves; bracts subtending the heads,
lanceolate or oblong-lanceolate, acute or acuminate, shorter than the involucre;
phyllaries about 7 mm. long, green, dirty brown when dried, narrowly lance-oblong,
acute or abruptly acute, densely brown-pilosulous or sometimes glabrate; corolla
glabrous, slightly shorter than the pappus; achenes narrowly prismatic, pale-
olivaceous, glabrous or very sparsely puberulent, 5 mm. long; pappus bristles
fulvescent, 6-7 mm. long, minutely scaberulous.
Mikania petrina is perhaps only a variation of M. cordifolia
that is somewhat more sordid pubescent than the typical form.
Mikania pterocaula Sch.-Bip. ex Hemsl. Biol. Cent. Am. Bot.
2: 103. 1881, hyponym; Klatt, Leopoldina 20: 91. 1884.
Dense, wet, mixed forest or thickets, 1,300-1,650 m.; Alta
Verapaz; Baja Verapaz; Chimaltenango; San Marcos. Southern
Mexico.
A stout, herbaceous vine, the stems 6-angulate or sometimes 4-angulate, with
narrow green wings along the angles, glabrous throughout or nearly so; leaves
112 FIELDIANA: BOTANY, VOLUME 24
slender-petiolate, thin, broadly ovate or rounded-ovate, mostly 7-10 cm. long,
acuminate or abruptly acuminate, rounded at the base, sometimes abruptly
contracted and short-decurrent, obscurely undulate-denticulate or almost entire, 5-7-
plinerved; inflorescence a large, open, somewhat leafy panicle; heads dirty white, 4
mm. long, in short dense spikes; phyllaries narrowly oblong, very obtuse, glabrous,
about 4 mm. long; achenes fuscous, glabrous; pappus dirty white.
Mikania pyramidata Donn.-Sm. Bot. Gaz. 13: 188. 1888.
Moist or wet, mixed forest, 2,500 m. or less; Alta Verapaz (type
from Coban Tuerckheim 1106); Baja Verapaz; Izabal; Zacapa;
Chiquimula(?); Huehuetenango; San Marcos(?). Mexico; Honduras.
A large, coarse, herbaceous vine, perhaps sometimes woody below, the stems
terete, densely setose-pilose with spreading, brown hairs; leaves thin, long-petiolate,
lance-ovate to very broadly ovate, mostly 10-20 cm. long, long-acuminate, cordate at
the base and usually somewhat hastate-lobate, undulate-dentate or subentire, 5-
plinerved, the inner nerves arising far above the base, thinly hirsute on both surfaces;
panicles large, lax, compound, pyramidal, the branches setose-pilose; heads laxly
cymose-corymbose, slender-pedicellate, 8 mm. long; phyllaries linear-oblong, 4 mm.
long, obtuse, pubescent at the apex or almost glabrous; achenes brown or fuscous, 3
mm. long, sparsely hispidulous, pappus yellowish white.
Mikania skutchii Blake, of Costa Rica, is very closely allied
and possibly synonymous.
Mikania vitifolia DC. Prodr. 5: 202. 1836. M. punctata Klatt,
Bull. Soc. Bot. Belg. 31, pt. 1: 195. 1892. Figure 24.
Moist or wet thickets or mixed forest, 1,650 m. or less; Peten;
Alta Verapaz; Escuintla; Retalhuleu. Southern Mexico; British
Honduras, along the Atlantic coast to Panama; tropical South
America.
A large vine, sometimes 8 m. long, occasionally woody below, the stems terete,
somewhat pubescent with spreading hairs or almost glabrous; leaves on long, slender
petioles broadly ovate to triangular or broader, 12-16 cm. long or even larger,
acuminate, deeply cordate at the base and often with distinct, acuminate lobes,
punctate above, rufous-pubescent beneath, especially along the nerves and veins, or
in age glabrate, commonly 5-plinerved; panicles usually large, compound, more or less
leafy, puberulent or brownish pubescent, the pedicels short; heads white or creamy
white, subglomerate at the ends of the branchlets; phyllaries linear-oblong, obtuse, 5
mm. long, pubescent toward the apex; achenes fuscous, glabrous, 3.5 mm. long;
pappus bristles dull white, in age often with a reddish tinge.
OXYLOBUS Mocino
Reference: B. L. Robinson, Revision of the genus Oxylobus,
Proc. Am. Acad. 49: 483-487. 1913.
Perennial herbs or low shrubs, branched or simple, viscid-pubescent; leaves
opposite, thick, persistent, obtuse, crenate, reticulate-veined, not punctate; heads
WILLIAMS: FLORA OF GUATEMALA 113
small, white, homogamous, in lax or dense, compound cymes; involucre cylindric-
campanulate or funnelform-campanulate; phyllaries ovate-oblong or lance-oblong,
acute, subherbaceous, glandular-pubescent, often costate; receptacle flat or nearly so,
naked; corolla tube slender, the limb campanulate-cylindric, the lobes oblong or
narrowly ovate, subacute, spreading; anthers rounded at the base, the apical
appendage ovate, membranaceous; achenes slender-prismatic, 5-angulate, scabrous on
the angles, callous at the base; pappus scales 5-10, unequal, usually acute and
fimbriate, much shorter than the achene.
Four species, in Mexico, Guatemala, and Venezuela.
Leaves mostly basal, the cauline ones about 3 pairs, remote; leaves broadly
spathulate, acute or attenuate at the base O. adscendens.
Leaves mostly cauline, numerous, oblong, elliptic, or ovate-oblong, often rounded or
obtuse at the base.
Leaves mostly glabrous, 1-3 cm. long, 3.5-12 mm. broad; heads 20-25-flowered.
O. arbutifolius.
Leaves pubescent, 3-5 cm. long, 10-20 mm. broad; heads 50-75-flowered.
O. glanduliferus.
Oxylobus adscendens (Sch.-Bip.) Rob. & Greenm. Proc. Am.
Acad. 41: 272. 1905. Ageratum adscendens Sch.-Bip. ex Hemsl. Biol.
Cent. Am. Bot. 2: 80. 1881. Bretonica.
Moist or dry, open, alpine slopes, sometimes among pine trees,
3,300-4,000 m.; Huehuetenango; San Marcos. High mountains of
southern Mexico.
A decumbent, perennial herb with hard, elongate rootstocks, 25-50 cm. high, the
basal portion of the stem elongate and radicant, the stems terete, with elongate
internodes, glandular-puberulent; basal leaves spathulate, 3-4 cm. long, 1-2 cm. broad,
obtuse or rounded at the apex, narrowed at the base into a short broad petiole,
crenulate, venose, glabrate above, puberulent beneath, at least on the veins, the
cauline leaves about 3 pairs, oblanceolate-oblong, sessile; cymes terminal, compound,
about 4 cm. broad; involucre 5-6 mm. high; outer phyllaries narrowly oblong, acute,
glandular-pubescent, the inner ones linear; corollas 3-7 mm. long, glabrous; achenes
2.8 mm. long, scabrous on the angles; pappus scales several, narrow, fimbriate, 0.5
mm. long.
Oxylobus arbutifolius (HBK.) Gray, Proc. Am. Acad. 15: 26.
1879. Ageratum arbutifolium HBK. Nov. Gen. & Sp. 4: 149. 1820.
Open, alpine areas, 3,700 m.; Huehuetenango (Sierra de los
Cuchumatanes, Steyermark 50161). High mountains of southern
Mexico.
A small, suffrutescent plant, usually erect, but sometimes trailing, often much
branched, mostly 20-40 cm. high, the stems often purplish and glandular-tomentulose,
the branches densely leafy; leaves mostly subsessile or the lowest subpetiolate, elliptic
or elliptic- oblong, firmly membranaceous to subcoriaceous, rich green above, pale
green below, 1-3 cm. long, 3.5-12 mm. broad, obtuse, rounded to cuneate at the base,
crenate, mostly glabrous, or the margin glandular-puberulent; cymes terminal,
114 FIELDI ANA: BOTANY, VOLUME 24
compound, 2-3 cm. broad, usually dense; involucre 5-5.5 mm. high; outer phyllaries
lance-oblong, glandular-puberulent, the inner ones linear-lanceolate; corollas white,
3.5-4 mm. long; achenes 2.2-2.5 mm. long; pappus scales several, oblong or linear, 0.7
mm. long, fimbriate.
Oxylobus glanduliferus (Sch.-Bip.) Gray, Proc. Am. Acad. 15:
26. 1879. Ageratum glanduliferum Sch.-Bip. ex Hemsl. Biol. Cent.
Am. Bot. 2: 82. 1881. Figure 25.
Open, rather dry, rocky slopes of the highest mountains, or in
open, pine or Juniperus forest, alpine situations, 3,000-3,800 m.;
Totonicapan; Huehuetenango (Sierra de los Cuchumatanes); Que-
zaltenango (Volcan de Santa Maria); San Marcos (volcanoes of
Tajumulco and Tacana). High mountains of southern Mexico;
Venezuela.
A stout, erect shrub, 1 m. high or less, or herbaceous and suffrutescent below,
often much branched, glandular-pubescent throughout and very viscid, the branches
densely leafy; lower leaves petiolate, usually with winged petioles, the upper ones
sessile, oblong or ovate-oblong, 2-5 cm. long, 1-2 cm. broad, obtuse, rounded to
cuneate at the base, crenate, villous-pilose; cymes compound, trichotomous, the
heads few or numerous, pedicellate or subsessile; involucre 6 mm. high; outer
phyllaries ovate-oblong, glandular-puberulent, the inner ones oblanceolate-linear;
corollas white, 5.5 mm. long; achenes 3-3.5 mm. long; pappus scales 5 or more,
oblong, 0.7 mm. long, lacerate.
A very common and typical plant of the Sierra de los
Cuchumatanes, also on the higher slopes of the western volcanoes.
PIQUERIA Cavanilles
Reference'. B. L. Robinson, Revision of the genus Piqueria,
Proc. Am. Acad. 42: 4-16. 1906; L. Wms. Fieldiana, Bot. 36: 103.
1975.
Annual or perennial herbs or shrubs, the stems erect or decumbent, leafy,
branched; leaves opposite or alternate, petiolate or subsessile, mostly serrate or
dentate, sometimes angulate; heads small, homogamous, 3- many -flowered; involucre
ovoid, cylindric, or campanulate; phyllaries usually few, subequal, laxly imbricate or
in almost a single series; receptacle flat or slightly convex, naked; corollas tubular,
white, the tube short, pilose or glandular-puberulent, the limb dilated, dentate;
achenes prismatic, 5-angulate, naked at the apex or with a deciduous annular disc,
rarely with a few short setae.
About 20 species, in the mountains of tropical America. Only
the following is known from Central America. The genus was
named for Andres Piquer, a Spanish physician of the 18th century.
See Eupatorium sodali which was originally ascribed to this
genus.
GRASHOFF: FLORA OF GUATEMALA 115
Piqueria trinervis Cav. Icon. 3: 19, t. 235. 1794. P. trinervis
var. luxurians Kuntze, Rev. Gen. 355. 1891 (type from Volcan de
Irazu, Costa Rica). P. luxurians Robinson ex Volkens, Verb. Bot.
Ver. Brandenb. 65: 118. 1923. Sauquillo (fide Aguilar). Figure 26.
Open, moist or dry, often rocky hillsides or in thin or dense,
moist or wet forest, sometimes on limestone, often with Alnus,
Pinus, or Abies, 1,200-3,300 m.; Chimaltenango; Solola; El Quiche;
Totonicapan; Huehuetenango. Mexico; Costa Rica; Panama; Hispa-
niola.
An erect herb, 1 m. high or less, almost glabrous, annual or sometimes perennial,
the stems terete, pale, puberulent in 2 lines, very leafy; leaves slender-petiolate, thin,
lanceolate to ovate, 9 cm. long or less, 3-5-nerved, long-acuminate, acute to almost
rounded at the base, serrate, nearly glabrous; heads white, mostly 4-flowered, 4 mm.
long, slender-pedicellate, laxly cymose-corymbose; phyllaries pale green, elliptic,
rounded and mucronate at the apex, scarious-marginate, erose-denticulate near the
apex; corolla tube short, pilose; achenes dark olivaceous, 5-angulate, 2-2.4 mm. long,
oblique and callous at the base; pappus none.
The plant is common in moist places throughout the
northwestern highlands, often growing abundantly in areas closely
grazed by sheep, and apparently not eaten by those animals, being
one of very few plants that they do not molest when pressed by
hunger. The plants wither soon after the rains end. In Mexico they
are much used in domestic medicine, especially as a febrifuge. In the
United States this plant sometimes is cultivated, usually under the
name Stevia, to be used as a filler in bouquets and set pieces.
STEVIA Cavanilles
By JEROLD L. GRASHOFF
Annuals, rhizomatous perennials, shrubs, or rarely plants with woody caudices
and herbaceous stems, the stems usually erect, glabrous to floccose-tomentose, the
trichomes sometimes viscid or glandular or both; leaves opposite, alternate or
scattered, filiform to orbicular to deltoid-reniform but most commonly spathulate,
lanceolate or ovate, sessile, petiolate, or, most commonly with a winged petioliform
base, surfaces usually sessile- or punctate-glandular or glandular-pilose;
inflorescences (in our species) terminal panicles or corymbs, the plants sometimes
scapiform; heads rayless, 3-18 mm. long, sessile to long- pedunculate; involucres
cylindrical to narrowly campanulate, uniseriate; phyllaries 5 in a quincuncial
arrangement; florets 5, in 1 whorl; corollas funnelform or with a more or less
expanded throat, actinomorphic to rather zygomorphic, with the adaxial lobes
shorter and more erect than the 3 outer lobes, pilulose just below the level of the
sinuses within, white, pink, magenta, violet, lavender, or greenish, the white flowers
often a clear, pale yellow when dried; lobes 5; achenes glabrous, hispid or sessile-
glandular, tapering to a small calloused base, 5-sided, the sides concave, the ribs
usually 5, sometimes white and calloused; pappus of awns or scales or both, often
116 FIELDIANA: BOTANY, VOLUME 24
coronulate, apparently uniseriate; awns (in our species) up to 5 (rarely 6) per achene,
frequently variable in number among the achenes in a single head, barbellate, tubular
or flattened, stiff or somewhat flexuous, never caducous; pollen spherical,
tricolporate and moderately echinate in sexual plants, variously deformed in
apomicts.
1. Herbs 2.
I. Shrubs 14.
2. Inflorescences lax, peduncles mostly equaling or exceeding the involucres 3.
2. Inflorescences more or less compact, peduncles mostly shorter than the
involucres or heads sessile in clusters 5.
3. Leaves alternate or crowded and scattered over the stem S. viscida.
3. Leaves opposite 4.
4. Leaves glandular throughout, not tomentose nor villous; phyllaries 5.5-6 mm.
long S. elatior.
4. Leaves glabrous to villous or tomentose, not stipitate-glandular; phyllaries less
than 5.5 mm. long S. lehmannii.
5. Leaves alternate or scattered, spathulate to sublinear S. serrata.
5. Leaves opposite, oblong to broadly ovate 6.
6. Leaves sessile, oblong to oblong-rhombic, nearly glabrous, often connate at base.
S. connata.
6. Leaves petiolate (petiole often winged), usually ovate, usually pubescent, blades
not connate 7.
7. Involucres stipitate-glandular 8.
7. Involucres sessile-glandular or glandless, often puberulent 9.
8. Inflorescence large and open, its branches elongate, arising from the axis at an
angle of 45-90 degrees, often beset with numerous, sessile, elliptic bracts;
largest leaves at the base of the stem, decreasing in size steadily upward;
achenes often aristate S. caracasana.
8. Inflorescence moderately compact, its branches short, arising at an angle of 45
degrees or less from the axis, never beset with elliptical bracts; leaves of the
major portion of the stem subequal in size; achenes always exaristate.
S. incognita.
9. Leaves in a basal rosette or limited to the lower one-fourth of the plant 10.
9. Leaves more or less evenly spaced throughout stem 11.
10. Leaves glabrous to sparsely pilose, reddish beneath, crenate to entire, rounded
at the apex S. seemannii.
10. Leaves densely pilose, green or suffused reddish, serrate, pointed at the apex.
S. hirsuta and vars.
II. Leaves densely tomentose or subtomentose beneath 12.
11. Leaves glabrous to puberulent beneath 13.
12. Leaves lanceolate, stems reddish, flowers diffuse pink S. suaveolens.
12. Leaves ovate to broadly ovate, stems green or yellow, flowers white.. ..S. triflora.
13. Florets white, achenes aristate or exaristate S. ovata.
13. Florets pink, achenes exaristate S. jorullensis.
14. Leaves chartaceous; heads about 6 mm. high; florets often yellowish when
dried S. tephrophylla.
14. Leaves membranaceous; heads over 8 mm. high; florets not yellowish when
dried 15.
15. Pappus a (usually incomplete) crown of lacerated scales, 0.5-1.5 mm. high.
S. microchaeta.
GRASHOFF: FLORA OF GUATEMALA 117
15. Pappus a complete crown of entire to minutely dentate scales to 0.5 mm. high,
rarely with short aristae interspersed 16.
16. Leaves glabrous, glutinous S. lucida var. oaxacana
16. Leaves arachnoid-tomentose at least on the veins or in the axils of the veins
beneath, not glutinous 17.
16. Leaves arachnoid-tomentose at least on the veins or in the axils of the veins
beneath, not glutinous 17.
17. Achenes exaristate; leaves arachnoid-tomentose at least on the veins and usually
completely beneath; corolla lobes puberulous without S. polycephala.
17. Achenes with minute aristae; leaves arachnoid only in the axils of the principle
veins beneath; corolla lobes glabrous without S. chiapensis.
Stevia caracasana DC. Prodr. 5: 121. 1836. S. elongata HBK.
as the name has been commonly used but not as to type (which
equals S. elatior HBK.). S. hirtiflora Sch.-Bip. Linnaea 25: 274.
1853. S. elongata HBK. var. caracasana (DC.) Robins. Contr. Gray
Herb. II, 90: 141. 1930.
Usually in oak or pine forests from 300-3,000 m. in altitude;
Alta Verapaz; Escuintla; Guatemala; Huehuetenango; Zacapa.
Sinaloa, Mexico to Colombia and Venezuela. It blooms from
December to March.
Rhizomatous, perennial herbs to 1.5 m. tall, the stems usually solitary,
puberulous to sparsely pilose, stipitate-glandular above; leaves opposite, petiolate,
thin-membranaceous, frequently blackening when dried, blade ovate (rarely lanceo-
late or elliptic), 3-10 cm. long, 2-5 cm. wide, serrate, crenate, or dentate, the apex
acute to obtuse, the base broadly cuneate to truncate (rarely acute or subcordate),
upper surface glabrous to obscurely pilose, typically without glandular punctation,
very rarely with sessile and/or stipitate glands, lower surface sparsely puberulous to
sparsely pilose along the veins, otherwise subglabrous, very rarely stipitate- and /or
sessile- glandular, veins raised below, whitish; inflorescence an open, often elongate,
divaricately branched corymb, branches opposite except for the small distal ones,
stipitate-glandular and often puberulous to pilose; bracts numerous, the lower mostly
petiolate, ovate, serrate with pointed apices, the upper sessile, elliptic, entire, seldom
linear; heads about 7-10 mm. high on somewhat flexuous, short peduncles in groups
of from 3-7, the groups typically subtended by several elliptic bracts; involucres 5-6
mm. high, stipitate-glandular, green or tinged purple; corollas 4.5-5.5 mm. long, pink,
sometimes pale; achenes heteromorphic or homomorphic, 3-4.7 mm. long excluding
pappus, hispid; pappus a low crown of united scales 0.1-0.3 mm. high, serrate; four
achenes frequently bearing 3 awns each, the awns alternating with the scales.
Stevia chiapensis Grashoff, Brittonia 26: 348. 1974.
Not yet known from Guatemala, where it is to be expected.
Mexico, (Southern Chiapas). It blooms from November to January
but nothing is know of its habitat preferences.
Shrubs, known only from the apical 2 dm. portion, the bark brown, glabrous,
smooth; leaves opposite, petiolate, lanceolate to lance-ovate, 5-10 cm. long, 2.5 cm.
wide, crenate to crenate-dentate, apex acute and acuminate, base cuneate, acuminate
118 FIELDIANA: BOTANY, VOLUME 24
at the petiole, upper surface obscurely puberulous along the veins, otherwise glabrous,
densely glandular-punctate, lower surface arachnoid tomentose in the vein axils
toward the base of the blade, otherwise glabrous, densely glandular-punctate; petiole
2.0-4.5 cm. long, expanded and keel-shaped at the base, glabrous; inflorescences
terminal corymbs, to 15 cm. across, branches at first opposite then alternate,
arachnoid tomentose in the axils, otherwise glabrous; heads ca. 8 mm. long;
involucres glabrous, 5 mm. long; corollas white (?), ca. 4 mm. long, glabrous; achenes
4 mm. long; pappus of several translucent scales, 0.3 mm. high alternating with 1-3
short awns, 0.8-1.2 mm. long.
This species is currently known only from a restricted portion
of the Chiapan Cordilleras.
Stevia connata Lag. Gen. & Sp. Nov. 27. 1816.
Trans-mexican volcanic belt south to Nicaragua, from 610-2,450
m. It blooms from August to November.
Rhizomatous, perennial herbs to 1 m. tall, the stems few, glabrous or sparsely
puberulous; leaves opposite, sessile, linear-oblong to oblong, rarely narrowly
lanceolate, only rarely proliferating in the axils, 4.5-11 cm. long, 3.5-15 mm. wide,
usually conduplicate, often reflex- curved, shallowly crenate-serrate, narrowly acute
at apex, acute to truncate to auriculate at base and usually connate; upper surface
glabrous, not glandular-punctate but bespeckled with numerous minute, white dots,
lower surface glabrous, paler, glandular-punctate, veins 3-5, paralleling the margins;
inflorescence a compact, terminal, rounded corymb, 7-19 cm. in diameter, branches
opposite and decussate, subglabrous or fuscous- puberulous; heads about 8-10 mm.
high, short-pedunculate; involucres 4.5-6 mm. high, glabrous to minutely puberulous,
with or without glandular punctations; corollas white, about 4-6 mm. long; achenes
2.5-3.7 mm. long excluding pappus, heteromorphic; pappus of the 4 adelphocarps of 3
awns, 4-5 mm. long, alternating with 3 scales, 0.5 mm. long; pappus of the idiocarp a
crown of separate, fimbriate scales 0.3-0.7 mm. high.
Stevia elatior HBK. Nov. Gen. & Sp. 4: 113. 1818. S. elongata
HBK. loc. cit. (as to type but not as commonly used from 1820 to
1970). S. elatior var. dissoluta Robins. Contr. Gray Herb. II, 90: 13.
1930. S. elatior var. podophylla Robins, loc. cit.
In Mexico and Central America, S. elatior is found at 1,300-
2,650 m. in and near pine and oak forests, usually in sunny sites and
frequently on red clay soil. It usually flowers from July to October.
In South America the species may reach higher altitudes and is said
to grow in fields, grassy areas, and near villages just below the
paramo; Chimaltenango; Huehuetenango; El Quiche: Sacatepe-
quez; Santa Rosa; Solola. Eastern and southern Mexico, Central
America, and northern South America.
Rhizomatous, perennial herbs to 1.5 m. tall, the stems solitary or few, erect,
unbranched to the inflorescence, densely glandular-pilose and often sparsely pilose
with long, tapering hairs; leaves ovate to ovate-deltoid (rarely elliptic), sessile,
subpetiolate or long-petiolate, crenate, the apex acute to obtuse to rounded, the base
GRASHOFF: FLORA OF GUATEMALA 119
obtuse to cordate (or cuneate if petioliform), blades 2-6 cm. long, 1.5-4 cm. wide,
triplinerved, glandular-punctate and stipitate-glandular on both surfaces, lower
surface slightly paler; petioles 0-3 cm. long, usually widely winged and indistinct from
the leaf blades when short; inflorescences terminal, open panicles, the branches
stipitate-glandular; heads about 10-11 mm. high on peduncles up to 2.2 cm. long,
immature heads borne on very short peduncles; involucres 4.5-6 mm. high, stipitate-
glandular, frequently purple or purple-tinged; corollas about 5 mm. high, reddish
purple with white or pale pink lobes; achenes heteromorphic (rarely homomorphic
and exaristate), 4-4.5 mm. long excluding pappus, rather densely hispid; adelphocarps
4, usually with 3 awns alternating with as many scales, occasionally with up to 5
awns; idiocarp single, exaristate with a crown of united or separate scales 0.3-0.6 mm.
high.
Stevia hirsuta DC. Prodr. 5: 121. 1836.
The species is divided into two varieties:
1. Achenes 3-3.5 mm. long, idiocarp hispid; involucres 4-5 mm. high; inflorescence
compact; heads approximate. Huehuetenango var. hirsuta.
I. Achenes 2-2.5 mm. long, idiocarp glabrous; involucres 3.5-4 mm. high; inflorescence
slightly open; heads often somewhat separate. Chiquimula var. chortiana.
S. hirsuta var. hirsuta.
In sunny locations on steep, rocky slopes in pine and oak forest
from 1,800-2,600 m., blooming in October and November; Huehue-
tenango. Widely scattered but locally abundant from Aguasca-
lientes and Hidalgo in Mexico to western Guatemala.
Rhizomatous, perennial herbs to 8 dm. high, the roots densely grey-tomentose,
the stems solitary, subsimple, pilose below, pilulose and puberulous above; leaves
usually confined to the lower one-third to one-half of the plant, opposite, ovate-
deltoid, 2.5-4.5 cm. long, 1.5-4 cm. wide, crenate, obtuse at apex and base, upper
surface sparsely to moderately pilose, with tapering or nontapering hairs, with or
without glandular punctations, lower surface pilose, especially along the veins,
glandular-punctate; petiole 1-2.5 (-4.5) cm. long, winged, pilose; inflorescence a
rounded corymb, branches opposite, densely puberulous, sparsely pilose; heads 7-9
mm. high; involucres 4-5 mm. high, green suffused with purple (especially at the
apex), puberulous; corollas 4.5-5.5 mm. high, purple or pink; achenes heteromorphic,
3-3.5 mm. long (the idiocarp somewhat shorter); pappus of the 4 adelphocarps,
triaristate; pappus of the idiocarp a crown of scales.
S. hirsuta var. chortiana (Standl. & Steyerm.) Grashoff,
Brittonia 26: 354. 1974. Stevia chortiana Standl. & Steyerm. Field
Mus. Bot. 23: 108. 1944.
Differing from var. hirsuta in its slightly more open
inflorescence (each head short-pedunculate), its smaller involucres
and achenes and its glabrous idiocarp. Endemic, known only from
the department of Chiquimula.
120 FIELDIANA: BOTANY, VOLUME 24
Stevia incognita Grashoff, Brittonia 26: 357. 1974.
In coniferous forests from 1,600-3,400 m.; Chimaltenango; El
Progreso; Guatemala; Huehuetenango; Jalapa; Quezaltenango; El
Quiche; Sacatepequez; San Marcos; Totonicapan. Mexico;
Honduras; Colombia; Venezuela.
Rhizomatous, perennial herbs to 1.5 m. tall, the stems purple, puberulent, often
stipitate-glandular above; leaves opposite, petiolate, ovate; blade 3-5 cm. long, 17-30
mm. wide, serrate to dentate (rarely crenate), the apex acute, the base usually acute
to obtuse (very rarely truncate), decurrent at the petiole, upper surface subglabrous
or crisped-puberulent and slightly scabrid, usually sessile- or punctate-glandular,
lower surface puberulent to sparsely hirsute, especially along the veins, veins usually
not contrasting greatly with the rest of the blade, the petiole winged, 1-3 cm. long,
increasingly less distinct in the upper leaves; inflorescence a few-to-many-branched
corymb, the branches opposite, the upper alternate in large inflorescences, sometimes
all alternate, stipitate-glandular and often puberulous; heads about 9-12 mm. high,
sessile to subsessile, mostly borne in groups of 15 or more; involucres 6.5-8.5 mm.
long, purple at least at the apex, stipitate-glandular (rarely also puberulous); corollas
pink to purple, about 5-6.5 mm. long; achenes homomorphic, exaristate, hispid, 3.5-4.5
mm. long excluding pappus; pappus a crown of separate to united scales, usually
about 0.5 mm. high, rarely up to 1 mm. high, serrate to fimbrillate.
This species can be somewhat variable in Guatemala. Except
for a few populations in Chiapas, it is represented only by apomictic
plants.
Stevia jorullensis HBK. Nov. Gen. & Sp. 4: 112. 1818.
Occasionally common in pine or oak forests from 1,000 to 3,200
m., in bloom from July to November; Guatemala; Solola;
Huehuetenango; Totonicapan. Nuevo Leon, Mexico to Guatemala
with outlying populations in western Mexico.
Rhizomatous, perennial herbs to 1 m. tall, the stems one to several, subsimple,
erect, purple (rarely green), pilose or crisped-puberulous; leaves opposite, usually
petiolate (rarely sessile or subsessile), the blade usually ovate but at times elliptic,
deltoid or rarely lanceolate, 1.5-5 (-8) cm. long, 1-3 (-5) cm. wide, usually serrate or
dentate with pronounced teeth but rarely crenate, upper surface glabrous to
puberulous, glandular-punctate, lower surface glabrous to pilose, glandular-punctate;
petiole 0-15 mm. long, tapering-winged; inflorescence a terminal, rounded corymb, the
branches opposite, puberulous to pilose; heads variable in size, 6-10.5 mm. high,
sessile to subsessile in fan-shaped or hemispherical clusters; involucres 5-8 mm. high,
puberulous, usually purple at least in part; corollas bright pink to bright purple or
lavender (white in some apomicts), 3-5 mm. long; achenes homomorphic, exaristate,
3-4 mm. long; pappus a crown of united to separate scales, 0.4-0.8 mm. high.
This species equals S. ovata in its variability, especially in leaf
shape and size and in head size. It can usually be separated from S.
ovata by virtue of the colored florets but a few apomictic clones in
Guatemala have apparently lost the coloration of the florets and
GRASHOFF: FLORA OF GUATEMALA 121
stems. These white flowered plants have heads larger than
homomorphic plants of S. ovata and in other respects resemble
colored clones and populations near to them. The species can be
distinguished from S. incognita by its lack of stipitate glands.
Stevia lehmannii Hieron. in Engler, Bot. Jahrb. 28: 562. 1901.
Pine or oak woods at about 1,000-1,500 m. in Mexico and
Central America and in unspecified habitats from 1,600-2,400 m. in
South America; Zacapa (Steyermark 29714, the only known
collection from Guatemala). Mexico (Guerrero, Puebla); Honduras;
Colombia; Venezuela.
Rhizomatous, perennial herbs, the stems solitary or few, sparsely pilose and
puberulous or densely glandular-pilose; leaves opposite, petiolate, the blade
lanceolate to ovate to triangular (rarely subreniform below), triplinerved, crenate, the
apex obtuse to rounded (rarely acute), the base cuneate to truncate to cordate, upper
surface scabrous or sparsely or moderately pilose, rarely stipitate-glandular at the
margin, obscurely glandular-punctate, lower surface tomentose or subtomentose,
densely but often obscurely glandular-punctate, petiole 0.6-2.5 cm. long, narrowly to
widely winged; inflorescences terminal, open panicles, often very diffuse, the branches
stipitate-glandular; heads 6-9 mm. high on peduncles up to 3 cm. long; involucres 4.5-
6 mm. long, stipitate-glandular, green; corollas pinkish or greenish with white lobes,
3.5-4 mm. high; achenes 2.5-3 mm. long excluding pappus, usually homomorphic and
exaristate but rarely with one or a few awns present in some of the heads; pappus a
crown of united regular scales, 0.2-0.7 mm. long, minutely serrate, rarely partially
expanded into short awnlike projections, rarely interrupted by a single awn.
This rarely collected species occurs in widely separated
populations. Each collection seems to have a slightly different
combination of morphological features.
Stevia lucida Lag. Gen. & Sp. Nov. 28. 1816.
All our specimens are of the following variety:
Stevia lucida var. oaxacana (DC.) Grashoff, Brittonia 26:
365. 1974. S. lucida var. oaxacana DC. Prodr. 5: 116. 1836.
Oak and pine woodlands in Mexico and Central America but in
the south most commonly encountered in subparamo regions,
blooming all year long, 1,800-3,200 m.; Chimaltenango; Huehue-
tenango; El Quiche; Quezaltenango. Mexico (Oaxaca, Chiapas);
Costa Rica and Panama; Colombia and Venezuela.
Shrubs to 2 m. tall, the stems many, bark smooth, brown; leaves opposite,
narrowly lanceolate to ovate, 2.5-12 cm. long, 0.8-4 cm. wide, serrate, glabrous,
obscurely glandular-punctate, glaucous when fresh, vernicose when dried, green, often
darkening on drying, apex narrowly acute, often acuminate, base acute (rarely
obtuse); petiole about one-fifth the length of the leaf blade; inflorescences terminal,
122 FIELDIANA: BOTANY, VOLUME 24
tripartite, somewhat open corymbs, the major inflorescence branches alternate or
subopposite, obscurely puberulous to pilulose, sometimes arachnoid, sometimes
vernicose and glabrous; heads about 10-12 mm. (rarely 8 mm.) long short-
pedunculate; involucre vernicose, green to yellow-green, narrow, sometimes
constricted towards the apex, glabrous, 4-7 mm. long, often purple-tinged, obscurely
punctate-glandular; corollas pink, 5-6 mm. long, totally glabrous; achenes 3.5-5.5 mm.
long; pappus a low crown of dentate to entire, united or separate scales 0.2 mm. high.
For reasons yet unknown, some races of this variety and the
typical variety sometimes have heads smaller than usual (8 mm.
long vs. 10-12 mm.). The plants occurring from Oaxaca to
Guatemala are often more robust than those from Costa Rica
southward. Plants of this variety are often difficult to distinguish
from subvernicose forms of S. polycephala var. polycephala where
they occur together in Guatemala. Four collections from four
different departments of Guatemala, vis. Steyermark 43466 (El
Progreso), Steyermark 50939 (Huehuetenango), Steyermark 33061
(Jalapa), and Standley 62376 (El Quiche), appear to be hybrids
between these taxa. The fact that plants of one or both of the
parent species have not yet been collected from El Progreso, Jalapa,
or El Quiche attests either to a more generalized distribution of
these species in earlier times, or to the inadequate botanical
knowledge of the areas.
Stevia microchaeta Sch.-Bip. Linnaea 25: 291. 1853. S.
vulcanicola Standl. & Steyerm. Field Mus. Bot. 23: 27. 1943 (type,
Steyermark 34757 from Quezaltenango).
In cloud forest region, just below the tree-fern zone at about
2,300-3,300 m.; Quezaltenango; San Marcos. Widely scattered in
portions of Veracruz, Oaxaca, and Chiapas, Mexico and in
Guatemala. It has been collected in bloom from June through
August and December through February. The latter season is
apparently the peak season of bloom.
Large shrubs to 4 m. tall, the stems few, sometimes weak, smooth-barked,
crisped-puberulent to glabrous, pith area greater than one half the diameter of the
stem, the branches few, opposite; leaves opposite, lanceolate, mostly over 10 cm. long
(up to 20 cm. long), and 3.5 cm. wide (up to 9 cm. wide), crenate to dentate, apex
acuminate, base acute (rarely obtuse); upper surface bright green, short-puberulous
on the veins, otherwise glabrous, obscurely glandular-punctate; lower surface paler,
glabrous except along the veins which are arachnoid-pilose, glandular-punctate;
immature leaves arachnoid-wooly beneath; petiole 3-6 cm. long, winged; inflorescence
a large, flat to rounded corymb 8-45 cm. across, lower branches opposite, upper
alternate, viscid arachnoid-pilose; heads short-pedunculate, sometimes somewhat
loosely disposed, about 12 mm. long; involucre light green, about 8 mm. long, sessile-
glandular, glabrous; corolla lavender, about 4.5-5 mm. long; achene about 6.2 mm.
long excluding pappus; pappus of irregular, translucent scales to 1.5 mm. long
GRASHOFF: FLORA OF GUATEMALA 123
(usually shorter), separate, frequently narrowly and deeply incised to lacerate,
resembling short awns, the ring of pappus scales is frequently incomplete.
Stevia microchaeta approaches S. polycephala var. polycephala
in Guatemala, where there is almost certainly some hybridization
occurring (cf. Standley 67388). As recognized here, S. microchaeta
consists of tall, often treelike shrubs with large leaves seldom under
10 cm. long. Specimens of S. polycephala have a pappus of low,
united or separate scales which are entire, or, if fimbrillate, then not
lacerated as is the case with S. microchaeta. The pappus crown is
always complete in S. polycephala, and the achenes are moderately
hispid.
Stevia ovata Willd. Enum. Hort. Berol. 855. 1809. S.
paniculata Lag. Gen. & Sp. Nov. 27. 1816. S. rhombifolia HBK.
Nov. Gen. & Sp. 4: 112. 1818. S. rhombifolia var. uniaristata (DC.)
Sch.-Bip. Linnaea 25: 279. 1853.
Common in rocky areas along roads and in forested areas,
usually above 1,200 m.; Chimaltenango; Guatemala; Huehue-
tenango; Jutiapa; Sacatepequez; San Marcos; Solola; Zacapa.
Northern Mexico to Ecuador. It blooms from July to January.
Rhizomatous, perennial herbs to 2 m. tall, the stems one to several, lignescent at
the base, usually simple to moderately branched, usually purple, crisped-puberulent
to weakly crisped- pilose to viscid-pilose; leaves opposite, petiolate or subpetiolate,
triplinerved, the blade ovate to rhombic (rarely lance-ovate or elliptic), (2.5-) 3-9 cm.
long, 1.5-5 cm. wide but sometimes larger or smaller in apomictic aberrants, crenate,
dentate or serrate, the apex acute to obtuse, base acute to obtuse, decurrent at the
petiole, upper surface subglabrous to obscurely puberulous, glandular-punctate, lower
surface puberulous along the major veins, otherwise usually glabrous, but
occasionally purple-viscid-pilose throughout, glandular-punctate; petiole widely
tapering- winged; inflorescence a terminal, rounded corymb of variable size, the
branches opposite or opposite below and subopposite or alternate above, puberulous
to tomentulose or viscid-pilulose (the color of the pubescence white, grey, or purple,
never yellow); heads about 7-10 mm. high, short-pedunculate; involucre 3.5-7 mm.
long, subglabrous to weakly crisped- or spreading- pilose or puberulous or viscid-
pilulose, green or purple-tinged; corollas 3-5 mm. long, white or rarely pink-tinged;
achenes about 3-4 mm. long excluding pappus, heteromorphic or homomorphic;
pappus of the 0-4 adelphocarps of 1-4 awns equaling the corolla, alternating with
small, dentate scales 0.4 mm. or less long; pappus of the 1-5 idiocarps a crown of
separate to united scales about 0.3-1 mm. long.
All of our plants are of var. ovata. This species is one of the
most variable in the genus. It is largely apomictic and contains
numerous clones or microspecies often interdispersed in the range,
or in parts of the range, of the species. Apomictic intermediates
between this and other species of the genus are frequently found. Of
124 FIELDIANA: BOTANY, VOLUME 24
particular frequency in Guatemala are intermediates with S.
jorullensis and S. triflora.
White flowered forms of S. jorullensis (apomicts) may be
confused with S. ovata. Stevia ovata, however, has heads smaller in
size than these "albino" plants.
Stevia polycephala Bertol. Novi Comm. Acad. Sci. Inst.
Bonon. 4: 432. 1840 (type: Bertoloni 26 from Sacatepequez). S.
arachnoidea Robins. Proc. Am. Acad. 35: 326. 1900 (type: Donnell-
Smith 2327 from Sacatepequez). Figure 27.
Oak, coniferous, or montane cloud forests or on gravelly soil on
hills and along roadsides from 2,600-3,200 m.; Chimaltenango;
Guatemala; Huehuetenango; Sacatepequez; San Marcos; Solola;
Suchitepequez; Totonicapan. Mexico (Chiapas). It has been collect-
ed from June through March and perhaps blooms throughout the
year.
Shrubs to 2 m. tall, the stems one to several, bark smooth, dull brown, branches
opposite, furrowed, puberulent, soon glabrate; leaves opposite, lanceolate, about 7-18
cm. long, 1.7-5.3 cm. wide, glandular-punctate, entire to serrate, apex acuminate, base
acute to acute-acuminate, upper surface grass-green, subglabrous to obscurely
puberulent, lower surface subglabrous to tomentulose between the veins, puberulous
to tomentose to arachnoid-wooly along the veins; immature leaves heavily pubescent
beneath (pulvo-puberulous to tomentose); inflorescences terminal, rounded corymbs,
6-20 cm. across, branches opposite (alternate above, puberulous to arachnoid-wooly;
heads short-pedunculate, about 12 mm. long; involucres glandular-punctate, glabrous
to puberulent to sparsely arachnoid-wooly, about 6.5 mm. long, green or with purple
tinges; corolla white, pink, or lavender, about 6 mm. long; achenes about 5.5 mm.
long; pappus usually a short crown of united, equal, minutely dentate scales, about
0.2 mm. high, occasionally separate, unequal or equal scales, as large as 1 mm. long,
the crown never incomplete nor lacerated.
Our specimens are all of var. polycephala.
Stevia polycephala is the most widely encountered shrub of its
genus in Guatemala, occurring in the Sierra Madres from the
Mexican border to Guatemala City, and in at least one area in the
Sierra de los Cuchumatanes. It is possible, however, that the
collections said to be from the Department of Guatemala are falsely
labeled, since all such collections are by one collector (Aguilar)
whose labels are frequently inaccurate, and the mountainous
regions near Guatemala City are not high enough to make this
species a likely inhabitant. Judging from morphologically inter-
mediate specimens, S. polycephala hybridizes with some degree of
facility with S. lucida var. oaxacana and S. microchaeta (which
see).
GRASHOFF: FLORA OF GUATEMALA 125
Stevia seemanii Sch.-Bip. in Seemann, Bot. Voy. Herald 298.
1856.
Rocky slopes in pine or oak forests from 1,000-2,500 m.;
Huehuetenango; El Quiche. Known with certainty only from
Oaxaca, Mexico and Guatemala. Blooming from August to
November.
Rhizomatous, perennial herbs to 0.5 m. tall, the roots tomentulose above, the
stems single, simple, erect, tomentose at base, sparsely pilose and sparsely puberulous
to the inflorescence, where it becomes densely puberulous or viscid-pilose; internodes
very contracted at the base (1-10 mm. long), greatly elongated above (to 20 cm. long);
leaves chiefly basal, opposite, petiolate, palmately nerved or triplinerved, blade ovate
to orbicular, sometimes rather spathulate, 1.5-4.5 cm. long, 1.2-3.5 cm. wide, crenate,
the upper leaves much smaller, elliptic, oblanceolate or spathulate, crenate to
subentire, petiolate or sessile, the apex rounded, rarely obtuse, the base cuneate to
truncate, decurrent at the petiole, upper and lower surfaces sparsely pilose, glandular-
punctate, lower surface deep purple, upper surface green or suffused with purple
especially along the veins and margins, veins raised below, pilose; inflorescence borne
well above the leaves, corymbose, 1-3 (-5) parted, the branches puberulous to viscid-
pilose, opposite; heads 10-11 mm. high, short-pedunculate to subsessile; involucres 7-
7.3 mm. high, purple, puberulous; corollas 6-6.5 mm. high, purple; achenes
homomorphic, 4 mm. long excluding pappus; pappus of 3-5 awns, 7 mm. long, and 3
scales 0.8-1 mm. long.
Stevia serrata Cav. Icon. & Descrip. Plant. 4: 33. 1797. S.
serrata var. ivaefolia (Willd.) Robins. Contr. Gray Herb. II, 90: 123.
1930.
The species grows along roadsides and in pastures in various
habitats from Yucca-Opuntia scrub to pine forests, usually over
1,500 m. in altitude. The plants prefer sunny, stony, well-drained
places but also grow in moist pastures and other flat areas.
Chimaltenango; Guatemala; Huehuetenango; Quezaltenango; El
Quiche; Sacatepequez; Solola. Southern Arizona to southern Texas
to northern Oaxaca, Chiapas to Honduras, Colombia to Ecuador.
Rhizomatous, perennial herbs to 8 dm. tall, the stems single to many, alternately
branched above, puberulous and often moderately to densely pilose, usually green;
leaves scattered and often crowded, sessile to subsessile, proliferating in the axils,
linear-spathulate to spathulate to oblanceolate, 2.5-6.5 cm. long, 2-15 (-20) mm. wide,
flat to carinate to conduplicate, serrate toward the apex or rarely subentire, apex
acute to rounded, base narrow, upper surface glabrous (except for the obscurely
puberulous margin) to subscabropuberulous to puberulous to pilulose, glandular-
punctate, lower surface subglabrous to puberulous to pilulose, glandular-punctate;
inflorescence a rounded to hemispherical to ovoid corymb, the branches alternate,
puberulous and often pilose; heads 5-9 mm. high, short-pedunculate; involucres 3.5-6
mm. high, green (rarely slightly purple-tinged), moderately to densely glandular-
punctate, appressed-puberulous to pilulose; corollas white, 3-5 mm. long; achenes
usually heteromorphic, 2.2-4.2 mm. long excluding pappus, hispid; pappus of the 4
126 FIELDIANA: BOTANY, VOLUME 24
adelphocarps with 3-5 awns equalling the corolla and alternating with 3-5 scales, 0.2-
0.7 mm. long; pappus of the idiocarp a crown of separate to united scales, 0.3-0.7 mm.
long.
All our specimens are of variety serrata.
Stevia suaveolens Lag., Gen. & Sp. Nov. 27. 1816. S.
nepetifolia ("nepetaefolia") HBK. Nov. Gen. & Sp. 4: 146. 1818. S.
nepetifolia var. leucantha (Schlecht.) Sch.-Bip. Linnaea 25: 279.
1853.
In pine-oak woods from 2,000-3,100 m. alt; Chimaltenango; San
Marcos; El Quiche. Mexico (Mexico) to Guatemala, with outlying
populations in Durango, Costa Rica, Colombia, and Ecuador.
Rhizomatous, perennial herbs to 7 dm. tall, the stems numerous, erect,
subtomentose-crisped-puberulent, often purple; leaves opposite, petiolate, the blade
lanceolate to lance-ovate (rarely ovate), 2.5-6 cm. long, 1.3-3.5 cm. wide, serrate
(rarely crenate), triplinerved, its apex usually acute, its base acute to obtuse, not
truncate, upper surface appressed-puberulous, glandular-punctate, lower surface
subtomentose-crisped-pilulose, sessile-glandular, grey, green, or brown, not white, soft
to the touch; petiole 5-20 mm. long, winged at least toward the blade; inflorescence
of terminal, well-branched corymbs, the branches at first opposite, then alternate,
subtomentose; heads about 8-9 mm. tall, sessile to subsessile in groups of about 5;
involucre subtomentose, sessile-glandular, often purple tinged, 5-7.5 mm. long;
corollas pale pink, 4-5 mm. long; achenes either heteromorphic or homomorphic and
exaristate, 3-4 mm. long excluding pappus; pappus of the adelphocarps of several
short scales and from 1-3 awns, 4.5 mm. long; pappus of the idiocarps a low crown of
separate to united scales, 0.3-0.7 mm. long.
The species is currently known only from apomictic material.
Stevia tephrophylla Blake, Contr. U. S. Nat. Herb. 22: 590.
1924. S. williamsii Standl. Ceiba 1: 95. 1950.
Not yet known from Guatemala. Chiapas, Mexico and
Honduras, in rocky places in pine forests at elevations of 780-1,350
m. It can apparently be in bloom at any time.
Small shrubs, 6 dm. high or less, the stems varying from tan to dark grey,
frequently with vestiges of a tomentose covering, shallowly furrowed, branches
numerous, opposite, grey- or white-tomentulose; leaves opposite, sessile or winged-
petiolate, lanceolate, oblanceolate or spathulate, 2-5 cm. long, 5-20 mm. wide, crenate
to subentire, apex rounded, base attenuate, upper surface bright green (frequently
drying to brown), glabrous or sparsely puberulous to subtomentulose, lower surface
white or grey owing to a dense tomentulum; veins pinnate or leaves subtriplinerved,
sunken above, raised below; inflorescences terminal and lateral corymbs 2-5 cm.
across, the branches few, short, opposite, tomentulose; heads about 6 mm. tall, short-
pedunculate; involucre about 4 mm. high, white-tomentulose over sessile glands;
corollas white (often appearing yellow when dried), about 3 mm. long; achenes
heteromorphic or homomorphic, 1.7-2 mm. long, excluding pappus; pappus of all
GRASHOFF: FLORA OF GUATEMALA 127
achenes a crown of united scales or one or more achenes with one to 3 slender awns
3.5 mm. long.
Stevia triflora DC. Prodr. 5: 115. 1836. S. compacta Benth. PI.
Hartweg. 197. 1847. S. rhombifolia HBK. var. stephanocoma Sch.-
Bip. Linnaea 25: 279. 1853.
Sunny and grassy sites and usually found near oak or pine
forest regions, blooming from September to December; Alta
Verapaz; Chimaltenango; Chiquimula; Guatemala; Huehuetenango.
Scattered from Jalisco, Mexico to Ecuador.
Rhizomatous, perennial herbs to 1.5 m. tall, the stems usually several, erect,
densely ochraleucous-puberulent, rarely floccose-fuscous-tomentose, green or brown
or yellowish, never purple; leaves opposite, short-petiolate, or with a winged petiole
one-third the length of the blade, the blade ovate (rarely lance-elliptic), triplinerved,
about 4-8 cm. long, about 2-5 cm. wide, crenate to serrate, apex acute, base acute,
often decurrent into the petiole, upper surface puberulent, glandular-punctate, lower
surface moderately to densely ochraleucous-puberulent to tomentose on and between
the major veins; petiole 4-20 mm. long, unwinged if short, broadly winged when long;
inflorescence a rounded corymb of (usually) crowded clusters of heads, lateral
inflorescences numerous, the branches opposite, ochraleucous-puberulous; heads
about 7 mm. long, short-pedunculate; involucre about 5 mm. high, green or yellowish,
puberulous, glandular-punctate; corollas about 3 mm. long, white; achenes
homomorphic exaristate, 2.5 mm. long excluding pappus; pappus a crown of united
scales about 0.3 mm. long.
The specific epithet is a misnomer. Stevia triflora has five
florets per head, although they may not all develop at the same
time.
Stevia viscida HBK. Nov. Gen. & Sp. 4: 110. 1818.
Pastures and roadsides on rather level sites from southern
Arizona and Texas to western Guatemala at elevations of 1,100-
2,200 m.; Huehuetenango (Steyermark 52044, the only known
Guatemalan collection). Southwestern United States; Mexico. It
blooms from August through early November.
Rhizomatous, perennial herbs to 1 m. tall, the stems single to many, viscid-pilose
and pilose-glandular; leaves alternate and often crowded, usually proliferating in the
axils, linear-spathulate to oblanceolate, 2-8 cm. long, 3-10 mm. wide, entire to
obscurely crenate to serrate, apex obtuse to rounded, base narrow, densely glandular-
punctate and glabrous above, stipitate- and punctate-glandular below; inflorescence a
large terminal panicle, typically over 3 dm. high and 2 dm. wide, the branches
alternate, stipitate-glandular; peduncles stipitate-glandular, mostly longer than the
involucre (but often greatly shortened in apomictic individuals with many or most
heads appearing sessile in clusters); heads 12-15 mm. high; involucres about 7-10 mm.
high, stipitate and sessile-glandular, green or partially or totally purple; corollas
purple to pinkish purple, about 6-9 mm. long; achenes about 5 mm. long,
128 FIELDIANA: BOTANY, VOLUME 24
homomorphic, black, hispid; pappus of 3-6 awns, 7-9 mm. long, alternating with
small, transparent scales to 0.5 mm. long.
This species has been collected only once in Guatemala and
except for a single site in extreme western Oaxaca, the nearest
populations are in Guerrero, Mexico. Except for plants nea_
Guadalajara, Mexico, the species is apomictic.
TRIBE III. ASTEREAE
By DOROTHY L. NASH
Annual or perennial herbs, shrubs, or small trees (Baccharis), often arching,
suffrutescent vines (Archibac charts); leaves (in ours) alternate; inflorescences
varying from a single terminal head to long, spicate inflorescences, to panicles,
corymbs, or cymes; involucres usually of numerous phyllaries (involucral bracts) in a
single series or 2-several-seriate; heads occasionally cylindric, but more often broadly
campanulate or hemispheric, usually heterogamous, but sometimes homogamous by
suppression of ray flowers; disc flowers always yellow; ray flowers (usually present)
heterochromatic, but never distinctly yellow or red; receptacle usually naked, anthers
obtuse at the base and entire or (rarely) sub-sagittate; style branches flattened,
appendaged (except Baccharis and Archibaccharis); florets several to many in a
head, usually small; pappus usually present but occasionally none (Achaetogeron) or
much reduced, most often of uniseriate or biseriate bristles.
The tribe is most abundant in north temperate regions; the two
predominantly tropical genera, Baccharis and Archibaccharis, are
prominent in the Guatemalan highlands and Baccharis vac-
cinioides is often the predominant shrub (or small tree) on the high
mountains of the highlands.
Conyza species are often weedy in Guatemala, as elsewhere,
and one or two species of Baccharis tend to invade disturbed
roadsides or fields.
Solidago, so abundant in species and often in individuals in
more temperate regions, has not been seen or collected by us in
Guatemala, although Dr. Lundell (in litt.) reports that one of his
collectors found a Solidago in Peten. There are several species in
Mexico.
Plants dioecious or polygamo-dioecious; pistillate flowers filiform, the limb truncate
or nearly so, rarely distinctly ligulate, if ligules present these always small and
inconspicuous.
Plants usually shrubs or small trees (rarely suffruticose), normally dioecious, the
heads wholly staminate or pistillate; leaves often triplinerved; achenes of
pistillate heads 5-10-costate Baccharis.
Plants herbs, shrubs, or vines, normally polygamo-dioecious (but largely function-
ally dioecious), the pistillate heads commonly heterogamous with 1-26 usually
NASH: FLORA OF GUATEMALA 129
sterile, central disc flowers; leaves seldom triplinerved; achenes of outer
pistillate flowers 2-5-7-costate Archibaccharis.
Plants neither dioecious nor polygamo-dioecious; pistillate flowers frequently ligulate.
Receptacles conic to ovoid.
Plants annual, usually viscid; leaves shallowly or deeply lobate, dentate,
pinnatifid, or bipinnatifid; pappus reduced to a cartilaginous cupule or a
narrow but evident, uneven crown Egletes.
Plants frequently perennial, not viscid; leaves (in ours) essentially entire; pappus
absent or reduced to a minute and inconspicuous ring of setae less than 0.1
mm. high Astranthium.
Receptacles flat or convex.
Ray flowers yellow.
Pappus well developed in both ray and disc flowers, uniseriate, 3-5 mm. long,
equal, or if unequal, not divided into 2 lengths.
Heads (in ours) solitary on long peduncles; ligules about 8 mm. long; pappus
more or less unequal Haplopappus.
Heads numerous, disposed in spiciform, racemose, or paniculate
inflorescences; ligules about 3 mm. long (in ours); pappus equal.
Solidago.
Pappus developed only in disc flowers, biseriate, the inner bristles about 7 mm.
long, the outer series much shorter, of minute scales or bristles, that of the
ray flowers absent or vestigial Heterotheca.
Ray flowers white, pink, blue, violet, or purple, never yellow (in ours); sometimes
the rays minute and inconspicuous or rarely, none.
Pappus none or reduced to a short, viscid beak or ring, or an inconspicuous
crown of minute scales or short setae.
Plants glutinous; leaves narrowly linear or rarely linear-lanceolate;
involucres narrowly ovoid-campanulate, the phyllaries 4-5-seriate and
unequal Gymnosperma.
Plants not glutinous; leaves orbicular or ovate to obovate or elliptic;
involucres hemispheric to open-campanulate, the phyllaries 2-3-seriate
and subequal.
Ray flowers 2-3-seriate, the ligules conspicuous, spreading, about 8 mm.
long (in ours); pappus reduced to an inconspicuous corona of short
setae Achaetogeron.
Ray flowers uniseriate, the ligules short, reflexed, less than 3 mm. long (in
ours); pappus reduced to a short viscid beak Lagenophora.
Pappus 1-3-seriate, the principal or inner series composed of slender bristles.
Plants frequently perennial; exterior flowers 1-2-seriate, the ligules
conspicuous, longer than the tubes, equalling or exceeding the pappus.
Inflorescences usually of solitary, long-pedunculate heads (rarely the
heads disposed in corymbiform arrangement); phyllaries subequal in
length; ray flowers pistillate, usually numerous, mostly more than 40;
style tips short, triangular Erigeron.
Inflorescences usually racemose or paniculate; phyllaries (in ours) very
unequal in length; ray flowers hermaphrodite, usually less than 40;
style tips elongated, slender and acuminate to ovate and acute.
Aster.
Plants usually annual or biennial, often weedy; exterior flowers 2-many-
130 FIELDIANA: BOTANY, VOLUME 24
seriate, the ligules absent or if present, inconspicuous, shorter than the
tubes, scarcely if at all exceeding the pappus Conyza.
ACHAETOGERON Gray
Usually perennial herbs with slender, striate stems, simple or branched; leaves
alternate, the margins usually entire; heads long-pedunculate; involucres hemis-
pheric; phyllaries usually biseriate, numerous, subequal, narrow, herbaceous or
subherbaceous; receptacles flat or slightly convex, foveolate or subalveolate; ray
flowers numerous (more than 60), usually in 2-3 series, narrow, pistillate, fertile, the
tube glabrous, the ligules spreading, 2- 3- denticulate; disc flowers hermaphrodite,
fertile, yellow, slender funnelform, 5-dentate; anthers obtuse at the base; styles of the
disc flowers short; achenes oblong, somewhat compressed, often widest about the
middle, tapering at each end, 2-nerved; pappus absent or reduced and simple, the
very short setae forming a minute corona.
Less than 10 species, distributed from Arizona through Mexico
and with one in Guatemala.
Achaetogeron guatemalensis (Blake) DeJong, Publ. Mich.
St. Univ. Biol. 2: 489. 1965. Astranthium guatemalense Blake,
Brittonia 2: 335. 1937. Figure 28.
Known only from the type locality, open, pine woods, Sierra de
los Cuchumatanes, 3,000-3,400 m., Huehuetenango, Skutch 1239,
type.
Perennial herbs, the stems slender, arching, procumbent, or prostrate, sometimes
rooting at the lower internodes, striate and sparsely pilose with short, spreading hairs;
cauline leaves alternate, sessile or nearly so, thin, mostly 1-3 cm. long, obovate or
obovate-oblong to elliptic, rounded or obtuse, bluntly and minutely apiculate,
gradually narrowing to the cuneate base, subentire or with 1-2 coarse teeth on each
side, more or less hirsutulous or pilose with spreading hairs, the basal leaves obovate
or spathulate, the blades mostly 2-3 cm. long, narrowly long-decurrent on the petiole,
this 2-3.5 cm. long; heads solitary on long, slender peduncles somewhat thickened just
below the head; involucres hemispheric; phyllaries biseriate, subequal, 4-5 mm. long,
lance-linear, acute or acuminate, hispidulous, greenish with a very narrow, pale,
scarious margin, the tips sometimes purplish, ciliolate at the apex; ray flowers 60-70,
3-seriate, pistillate, white, the ligules about 8 mm. long; disc flowers numerous, fertile,
yellow, about 3 mm. long; achenes 1.5-1.8 mm. long, essentially glabrous or sparsely
hispidulous toward the apex, with a very short corona of minute setae.
ARCHIBACCHARIS Heering
References: S. F. Blake, Hemibaccharis, a new genus of
Baccharidinae, Contr. U. S. Nat. Herb. 20: 543-554, tt. 48-51. 1924.
John D. Jackson, Notes on Archibaccharis, Phytologia 28: 296-302.
1974.
Herbs or shrubs, erect, arching, or scandent; leaves alternate, penninerved, sessile
or short-petiolate, usually serrate; plants polygamo-dioecious; inflorescences cymose-
NASH: FLORA OF GUATEMALA 131
paniculate; heads discoid or rarely radiate; involucres hemispherical or campanulate;
phyllaries 3-6-seriate, linear to ovate, narrowly scarious-marginate and more or less
ciliolate; receptacle flat, alveolate; pistillate heads heterogamous, the outer flowers
few to many, pistillate and fertile, the corollas with a filiform tube, subtruncate or
with a very short, erect, often bidentate limb, rarely distinctly ligulate; central 1-15
flowers hermaphrodite but often sterile, the corollas tubular, 5-dentate; staminate
heads with 5-93 hermaphrodite flowers, the corollas tubular, regular, 5-dentate, and
sometimes a few small, inconspicuous, sterile ray flowers present; stamens subentire
or rarely minutely sagittate at the base, the terminal apical appendages elliptic,
rather long; achenes compressed, rarely trigonous, hispidulous; pappus uniseriate, the
bristles of the pistillate pappus capillary; bristles of the staminate pappus often
dilated at the apex.
Twenty or more species, in tropical America, with 12 in
Guatemala.
Leaf bases abruptly contracted and narrowing into broadly winged petioles, these
dilated and clasping at the base A. blakeana.
Leaf bases not narrowing into broadly winged petioles, the blades either sessile or
petiolate, the petioles neither dilated nor clasping at the base.
Leaf blades linear-lanceolate or very narrowly lanceolate, often less than 1 cm.
wide, commonly 7-8 times longer than broad; stems, leaves, and pedicels
glabrous A. androgyna.
Leaf blades never linear-lanceolate; if lanceolate, the principal leaves more than 1
cm. wide; stems, leaves, and pedicels more or less pubescent, puberulent, or
tomentose (except the lower leaf surface of A. salmeoides only obscurely
puberulent).
Lower leaf surfaces velutinous or densely white-tomentose.
Leaf bases rounded and then becoming rather abruptly cuneate; cymes
disposed in large, open panicles; heads usually small, the involucres 2-3
mm. high; ray flowers white .A. serratifolia.
Leaf bases rounded to subacute; cymes disposed in dense, more or less
corymbiform panicles; heads larger, the involucres 3-4 mm. high; ray
flowers pale pink or purplish A. corymbosa.
Lower leaf surfaces not velutinous nor white-tomentose.
Leaf blades mostly elliptic or elliptic-lanceolate, attenuate to the base or
sometimes cuneate.
Plants more or less scandent; pistillate heads commonly with 16-20 (-25)
pistillate flowers; staminate heads with 10-18 (-21) hermaphrodite
flowers A. flexilis.
Plants usually erect, sometimes arching; pistillate heads with 25-35 pistillate
flowers; staminate heads with 22-30 hermaphrodite flowers.
Stems of inflorescence very minutely and densely puberulent with fine,
short hairs -A- aequwenia.
Stems of inflorescence more or less pubescent with spreading hairs.
A. asperifolia.
Leaf blades mostly ovate or obovate to oblanceolate, rounded, subcordate, or
subacute at the base or rounded and then abruptly cuneate.
Stems (at least the older ones) fractiflex.
Heads small, commonly about 2 mm. high, rarely to 3 mm.
A. hirtella var. taenotricha.
132 FIELDIANA: BOTANY, VOLUME 24
Heads larger, commonly 3-6 mm. high.
Leaf blades coriaceous, essentially glabrous above, shining, any
indument confined to costae and veins; pistillate heads with only 8-
12 (-22) flowers; phyllaries obtuse or subacute A., salmeoides.
Leaf blades chartaceous, thinly pilosulous above; pistillate heads with
28-30 (-50) flowers; phyllaries acute or acuminate A. schiedeana.
Stems not fractiflex.
Indument of stems and pedicels composed of multiseptate, acuminate,
primarily eglandular hairs; pistillate heads with 17-29 filiform flowers;
staminate heads 4-6 mm. high A. subsessilis.
Indument of stems and pedicels composed entirely of gland-tipped hairs;
pistillate heads with 30-40 filiform flowers; staminate heads about 7
mm. high A. lineariloba.
Archibaccharis aequivenia (Blake) D. Nash, Fieldiana:
Botany 36 (9): 73. 1974. A. standleyi Blake var. aequivenia Blake,
Brittonia 2: 340. 1937.
Damp thickets, 900-1,300 m.; Quezaltenango; Suchitepequez
(type from Finca Moca, Skutch 2056). Mexico (Chiapas).
Erect or arching shrubs to about 2 m. tall, the stems slender, often flexuous, very
minutely and densely puberulent; upper leaves sessile, the lower ones on very short
petioles, the blades thin, lanceolate to lance-oblong or elliptic, mostly 4-13 (-16) cm.
long, 1.5-3.5 cm. wide, long-acuminate, attenuate to a cuneate base, the margins
remotely serrulate, minutely scaberulous above, glandular-punctate and sparsely and
minutely puberulent beneath; inflorescences cymose-paniculate, the heads oh slender,
densely tomentulose pedicels; involucres densely and minutely puberulent, those of
the pistillate heads 3-4 mm. high, those of the staminate heads 2-3 mm. high;
phyllaries 3-4-seriate, whitish with dark green midribs, linear, acute or acuminate, the
margins inconspicuously ciliolate; pistillate heads with 26-31 filiform pistillate flowers
and 1-3 hermaphrodite flowers; staminate heads with 1-3 inconspicuous, sterile,
filiform pistillate flowers and 22-25 hermaphrodite flowers; achenes 3-5-costate,
hispidulous, about 1.8 mm. long; pappus white or sordid, that of the pistillate flowers
3-3.5 mm. long, that of the hermaphrodite flowers 2-3 mm. long.
Archibaccharis androgyna (Brandg.) Blake, Contr. U. S.
Nat. Herb. 23: 1509. 1926. Baccharis androgyna Brandegee, Univ.
Calif. Publ. Bot. 6: 77. 1914. Hemibaccharis androgyna Blake,
Contr. U. S. Nat. Herb. 20: 552. 1924. Copalillo.
Damp or wet thickets or open forest, 2,200-3,000 m.; Huehue-
tenango; San Marcos. Mexico (Chiapas).
Erect shrubs, 1-2 m. tall, glabrous throughout, the branches subterete; leaves on
very short petioles, the uppermost ones subsessile, the blades narrowly lanceolate or
linear-lanceolate, mostly 4-9 cm. long, 0.7-2 cm. wide, narrowly long-attenuate,
almost rounded or subacute at the base, the margins inconspicuously serrate or
serrulate; inflorescences cymose-paniculate, the panicles usually large and dense with
numerous, short-pedicellate heads; involucres 2-3 mm. high; phyllaries 4-5-seriate,
linear-lanceolate, acuminate or acute, glabrous, pale greenish- white with a slightly
NASH: FLORA OF GUATEMALA 133
darker midrib and the apex tinged with red or purple, ciliolate; pistillate heads with
17-30 filiform pistillate flowers and 1-15 (commonly 4) tubular hermaphrodite
flowers; achenes somewhat shiny, pubescent, 1-1.4 mm. long; pappus white or sordid,
that of the pistillate heads 1.5-3 mm. long; no staminate plants seen.
Archibaccharis asperifolia (Benth.) Blake, Contr. U. S. Nat.
Herb. 23: 1509. 1926. Baccharis asperifolia Benth. PL Hartweg. 86.
1841. Conyza asperifolia Benth. & Hook, ex Hemsl. Biol. Centr.
Amer. Bot. 2: 126. 1881. B. scabridula Brandg. Univ. Calif. Publ.
Bot. 6: 77. 1914. Hemibaccharis asperifolia Blake, Contr. U. S. Nat.
Herb. 20: 552. 1924. Figure 29.
Wet to dry thickets or open forest, often in pine-oak forest,
rarely in wet meadows, sometimes along roadsides, 1,200-3,400 m.;
Alta Verapaz; Baja Verapaz; Chimaltenango; Guatemala; Huehue-
tenango; El Progreso; Quezaltenango; El Quiche; Sacatepequez;
San Marcos; Solola; Suchitepequez; Totonicapan; Zacapa. Southern
Mexico; Honduras; northern Nicaragua.
Plants usually stiffly erect, 1-2 m. tall, rarely as high as 6 m., herbaceous,
suffrutescent, or shrubby, the stems terete, often dark red or purplish, glabrous or
nearly so below but sometimes puberulent to pubescent near the area of
inflorescence; leaves short-petiolate or the uppermost ones subsessile, the blades
mostly elliptic-lanceolate, mostly 4-14 cm. long, 1-4 cm. wide, long-acuminate, acute
or more commonly attenuate to the base, the margins serrate or subentire,
scaberulous above, puberulent or sometimes pubescent beneath, at least on the veins;
inflorescences basically cymose, the cymes disposed in large rounded, open, leafy
panicles, the heads numerous, pedicellate, the pedicels and peduncles more or less
pubescent with spreading hairs; involucres 3-4 mm. high; phyllaries 4-5-seriate, linear,
greenish, pale, the midrib darker, obtuse or acute, sparsely puberulent or glabrous,
somewhat ciliolate; pistillate heads with 25-35 (-55) filiform pistillate flowers and 1-7
hermaphrodite flowers; staminate heads with 22-56 (usually about 30) hermaphrodite
flowers; achenes pubescent, 2-4-costate, about 1 mm. long; pappus sordid, that of the
pistillate flowers 3-4 mm. long, that of the staminate flowers 2-3 mm. long.
Archibaccharis blakeana Standl. & Steyerm. Field Mus. Bot.
22: 296. 1940.
Damp or dry thickets or rocky forest, frequently in pine forest,
1,500-2,100 m.; Chimaltenango; Guatemala; Quezaltenango; Sacate-
pequez (type collected near Antigua, Standley 58597). Mexico
(Chiapas).
Arching or clambering shrubs, 1-3 m. tall, or sometimes herbaceous almost
throughout, usually densely branched, the branches flexuous, sometimes sarmentose,
obtusely angulate, villosulous or almost wholly glabrous; leaves on broadly winged
petioles with dilated, auriculate, clasping bases (except the uppermost leaves
sometimes sessile or subsessile), the blades usually broadly ovate, rounded-ovate, or
lanceolate, mostly 5-12 (-15) cm. long and 2-8 cm. wide (the uppermost ones smaller
134 FIELDIANA: BOTANY, VOLUME 24
and sometimes oblong-ovate), acuminate or long-acuminate, abruptly contracted
toward the base and obtuse to subtruncate, narrowing into the broad wing of the
petiole, scabrous above, sparsely short-villosulous on the costae and veins beneath or
essentially glabrous, the margins undulate-dentate; inflorescence cymose, the small,
rounded cymes usually disposed in large, leafy panicles; heads pedicellate, the
pedicels mostly 3-6 mm. long, villosulous; involucres 3.5-4.5 mm. high; phyllaries 4-6-
seriate, pale greenish white with a darker midrib, linear, acute or obtuse, ciliolate,
sparsely appressed-pilosulous or almost glabrous; pistillate heads with 30-50 filiform
pistillate flowers and 1-3 tubular hermaphrodite flowers, the staminate heads with 17-
25 hermaphrodite flowers; achenes shining, pale, about 1 mm. long, sparsely
appressed-pilosulous; pappus white or sordid, that of the pistillate flowers about 3
mm. long, that of the staminate flowers 2-3 mm. long.
Archibaccharis corymbosa (Donn.-Sm.) Blake, Journ. Wash.
Acad. Sci. 17: 60. 1927. Diplostephium corymbosum Donn.-Sm. Bot.
Gaz. 23: 8. 1897. Hemibaccharis corymbosa Blake, Contr. U. S. Nat.
Herb. 20: 553. 1924.
Damp or dry mountain forest or thickets, frequently in
coniferous forest, 2,100-3,600 m.; Chimaltenango; Huehuetenango
(type from Todos Santos, Nelson 3639); Quezaltenango; Sacatepe-
quez; San Marcos; Totonicapan.
Erect or arching shrubs, 1-3 m. tall, or sometimes almost wholly herbaceous,
branched, the stems densely pubescent or tomentulose with short, pale hairs; leaves
short-petiolate, the blades elliptic-oblong, mostly 5-10 (-12) cm. long and 1-4 cm.
wide, acute or acuminate, rounded to subacute at the base, the margins more or less
serrate, puberulent above, velutinous beneath; inflorescences cymose, the cymes
forming broad, usually dense, corymbiform panicles; heads numerous; involucres of
the pistillate heads about 4 mm. high, those of the staminate heads about 3 mm.
high; phyllaries 4-5-seriate, purplish, linear-lanceolate, acute, densely puberulent or
pilosulous; flowers deep purple to pale pink, the rays about 4 mm. long; pistillate
heads with 27-40 filiform ray flowers and 1-4 hermaphrodite flowers, the staminate
heads with 30-38 hermaphrodite flowers and 9-12 (-23) sterile ray flowers; achenes
minutely glandular and hispidulous, about 1 mm. long; pappus white or sordid,
sometimes the apices tinged with pink, that of the pistillate flowers 3-4 mm. long,
that of the staminate flowers about 2 mm. long.
Archibaccharis flexilis Blake, Journ. Wash. Acad. Sci. 17: 60.
1927. Hemibaccharis flexilis Blake, Contr. U. S. Nat. Herb. 20: 549.
1924.
Damp forest or thickets, 500-2,900 m.; Alta Verapaz (type from
Coban, von Tuerckheim II. 1636); Chimaltenango; Quezaltenango;
Sacatepequez; San Marcos; Solola; Suchitepequez; Totonicapan.
Mexico; El Salvador; Nicaragua; Costa Rica.
Arching shrubs or woody vines, often climbing over medium-sized trees, the
slender branches then greatly elongated, densely sordid-pilose with multiseptate,
spreading hairs, in age glabrate; leaves short-petiolate, the blades elliptic or lance-
NASH: FLORA OF GUATEMALA 135
elliptic to oblanceolate, mostly 4-8 (-13) cm. long, 1.5-4 cm. wide acuminate, usually
attenuate to the base, sometimes narrowing to a cuneate base, the margins remotely
serrate to subentire, puberulent above on the costa and sometimes on the veins,
sordid-pilosulous beneath, often densely so; inflorescences axillary and terminal,
cymose-paniculate, the panicles small, numerous, the pedicels mostly 2-5 mm. long,
sordid-pilosulous; involucres 3-4 mm. high; phyllaries whitish with dark green
midribs, the outer ones ovate-lanceolate, the inner ones linear, acute, ciliolate, the
outermost puberulent; pistillate heads commonly with 16-20 (-25) filiform, pistillate
flowers and 1-4 tubular hermaphrodite flowers; staminate heads with 10-20
hermaphrodite flowers, rarely with a few filiform pistillate flowers; achenes
hispidulous, about 1 mm. long; pappus sordid, that of the hermaphrodite flowers 2.7-3
mm. long, that of the pistillate flowers 3-4.3 mm. long.
Archibaccharis hirtella (A. DC.) Heering, Jahrb. Hanb. Wiss.
Anst. 21: Beih. 3: 41. 1904. Baccharis hirtella A. DC. in DC. Prodr.
5: 418. 1836. Hemibaccharis hirtella Blake, Contr. U. S. Nat. Herb.
20: 549. 1924.
Slender, woody vines, sometimes climbing over medium-sized trees, the branches
densely hirsutulous or puberulent with short, spreading, brownish, gland-tipped hairs
or with longer, eglandular hairs; stems more or less fractiflex; leaves short-petiolate,
the blades mostly ovate-oblong or oblanceolate, mostly 3-6 (-10) cm. long, 1-2.5 (-4)
cm. wide, acute or acuminate, rounded or obtuse at the base, the margins commonly
rather coarsely serrate, sometimes almost entire, scabrous on both surfaces or softly
pubescent beneath, or the costa beneath sometimes glandular-pubescent or pilose
with eglandular hairs; cymes small, the heads short-pedicellate; involucres 2-3 mm.
high; phyllaries 4-5-seriate, purplish, linear-lanceolate to linear, acute, more or less
puberulent, often recurving in age; pistillate heads with 10-18 filiform pistillate
flowers and 1-2 tubular hermaphrodite flowers; staminate heads with 15-25
hermaphrodite flowers; achenes about 1 mm. long, hispidulous; pappus of the
pistillate flowers about 3 mm. long, that of the staminate flowers 1.5-2.5 mm. long.
The typical variety with short, glandular hairs has not as yet
been found in Guatemala, but the following variety is well
distributed there.
Archibaccharis hirtella var. taeniotricha Blake, Journ.
Wash. Acad. Sci. 24: 433. 1934.
Damp or wet thickets or forest, 900-3,000 m.; Chimaltenango
(type from Santa Elena, Skutch 276); Chiquimula; Guatemala;
Jalapa; El Progreso; Quezaltenango; Sacatepequez; San Marcos;
Solola. Mexico; El Salvador.
Differs from the typical variety only in the indument of longer,
eglandular hairs.
Archibaccharis lineariloba J. D. Jackson, Phytologia 28: 300.
1974.
136 FIELDIANA: BOTANY, VOLUME 24
Known only from the type collection, steep, rocky slopes in
montane forest, Sierra Cuchumatanes, 3,700 m., Huehuetenango,
Molina, Burger, & Wallenta 16446.
Slender, erect or arching, suffrutescent plants about 1.5 m. high, the branches
densely covered with short, spreading, multiseptate hairs; uppermost leaves sessile or
subsessile, the lower ones on petioles only 1-4 mm. long, the blades rather thick and
firm, obovate to oblong-ovate or elliptical, mostly 2-6 cm. long, 1.5-3 cm. wide, short-
acuminate, rounded or more or less cordate at the base, the margins serrate,
scaberulous and glandular-pubescent above, pubescent below with mostly gland-
tipped hairs and a few longer, eglandular hairs, the venation commonly elevated and
reticulate; inflorescences cymose-paniculate, the rounded panicles with numerous
heads, the slender pedicels mostly 3-10 mm. long, densely glandular- puberulent;
involucres 2-3 mm. high; phyllaries lanceolate to linear, acuminate or acute,
glandular-puberulent, the midrib and tip dark green to purplish; pistillate heads
commonly with 30-40 white ray flowers about 3 mm. long and 2-4 hermaphrodite
flowers, the corolla tubes puberulent; staminate heads about 7 mm. high, the filiform
pistillate flowers about 9, sterile, the hermaphrodite disc flowers about 36; achenes 1-
1.5 mm. long, shining, glandular and hirsutulous; pappus yellowish white, the bristles
3-4 mm. long.
Archibaccharis salmeoides Blake, Journ. Wash. Acad. Sci.
17: 61. 1927. Hemibaccharis salmeoides Blake, Contr. U. S. Nat.
Herb. 20: 548, t. 50. 1924 (type from Coban, Alta Verapaz, von
Tuerckheim II. 1641).
Known only from the type locality, about 1,350 m., Alta
Verapaz.
Scandent, woody plants, the stems terete, more or less fractiflex, sordid-
puberulent or glabrate; leaves short-petiolate, the blades coriaceous, ovate to
oblanceolate, mostly 4-9 cm. long, 2-5 cm. broad, acute or acuminate, almost rounded
or cuneate at the base, the margins entire or mucronate-dentate, shining and glabrous
above or somewhat puberulent along the costa, obscurely puberulent beneath, the
indument usually confined to costae and veins, the lateral veins conspicuously
curved-anastomosing; inflorescences axillary and terminal, cymose-paniculate,
rounded, composed of few-many pedicellate flower heads; involucres 3-5 (-6) mm.
high; phyllaries more or less hispidulous, the midrib dark gree.i, the margins lacerate-
ciliate, the outermost phyllaries ovate, the inner ones lanceolate to linear-oblong,
obtuse or subacute; pistillate heads with 8-12 pistillate flowers and 1-4 (commonly 2)
hermaphrodite flowers; staminate heads with 15-16 hermaphrodite flowers, these with
tubular corollas with short, broad, almost ovate lobes; achenes about 1.2 mm. long,
hispidulous; pappus brownish, that of both pistillate and staminate flowers 3-4 mm.
long.
Archibaccharis schiedeana (Benth.) J. D. Jackson,
Phytologia 28: 297. 1974. Baccharis schiedeana Benth. in Oerst,
Nat. For. Kjoeb. Vid. Medd. 83. 1852. B. elegans var. seemannii Sch.
Bip. Bot. Voy. Herald: 303. 1856. B. thomasii Klatt, Abh. Naturf.
Ges. Halle 15: 326. 1881. Hemibaccharis torquis Blake, Contr. U. S.
NASH: FLORA OF GUATEMALA 137
Nat. Herb. 20: 550, t. 51. 1924. Archibaccharis torquis Blake in
Standl. op. cit. 23: 1508. 1926. Cahacillo (Chiquimula); culebrina
(Guatemala, fide Aguilar); te silvestre (Retalhuleu).
Damp or wet thickets or forest, 600-2,400 m.; Alta Verapaz;
Baja Verapaz; Chimaltenango; Chiquimula; Guatemala; Huehue-
tenango; Jalapa; Jutiapa; Quezaltenango; El Quiche; Retalhuleu;
Sacatepequez; San Marcos; Santa Rosa; Suchitepequez. Southern
Mexico; El Salvador; Costa Rica; Panama.
Weak shrubs, 1-3 m. tall, or often becoming scandent, scrambling over other
vegetation, the arching branches tortuous or fractiflex, puberulent or pilosulous, soon
glabrate; leaves short-petiolate or the uppermost ones subsessile, the blades
chartaceous, broadly ovate to lance-ovate, mostly 3-8 cm. long, 1.5-5 cm. wide, acute
or short-acuminate, rounded at the base or rounded and rather abruptly cuneate, the
margins mucronate-dentate or denticulate or almost entire, thinly puberulent or
pilosulous above, soon glabrate, sordid-pilosulous beneath, chiefly on the costae and
veins; inflorescences axillary and terminal, cymose, the cymes disposed in small,
rounded panicles, the heads pedicellate; pedicels mostly 2-9 mm. long, puberulous;
involucres 3-4 mm. high; phyllaries 5-6-seriate, linear, acute or acuminate, pale with
green midrib, ciliolate, sparsely puberulent; pistillate heads commonly with 28-30 (17-
50) filiform, pistillate flowers and 1-3 hermaphrodite flowers; staminate heads
commonly with 7-23 hermaphrodite flowers; achenes hispidulous, about 0.8 mm. long;
pappus sordid, that of the pistillate flowers 2-2.5 mm. long, that of the staminate
flowers about 3 mm. long.
Archibaccharis serratifolia (HBK.) Blake, Contr. U. S. Nat.
Herb. 26: 236. 1930. Baccharis serratifolia HBK. Nov. Gen. & Sp. 4:
59. 1820. B. mucronata HBK. torn, cit.: 60. B. micrantha HBK. loc.
cit. Diplostephium paniculatum Donn.-Sm. Bot. Gaz. 23: 8. 1897
(type from Huehuetenango, Nelson 3629). Hemibaccharis mucr-
onata Blake, Contr. U. S. Nat. Herb. 20: 550. 1924. H. mucronata
var. paniculata Blake, torn, cit.: 551. Archibaccharis mucronata
Blake in Standl. op. cit. 23: 1508. 1926. A. mucronata var.
paniculata Blake, torn, cit: 1509. A. serratifolia var. paniculata
Blake, Journ. Wash. Acad. Sci. 31: 328. 1931 (Type from
Huehuetenango, Nelson 3629).
Damp or dry thickets or forest, often in pine-oak forest, (384)
1,200-2,600 m.; Chimaltenango; Escuintla; Guatemala; Huehue-
tenango; Jalapa; Quezaltenango; El Quiche; Sacatepequez; Santa
Rosa. Mexico.
Erect or rarely arching shrubs, 1-3 m. tall, or sometimes suffrutescent plants, the
branches terete, commonly white-tomentose; leaves short-petiolate, the blades
lanceolate, lance-ovate, or oblong-lanceolate (rarely broadly ovate), mostly 3-12 cm.
long and 1.5-6 cm. wide, acuminate, rather abruptly cuneate at the base, the margins
conspicuously dentate or almost entire, more or less scabrous above, usually densely
138 FIELDIANA: BOTANY, VOLUME 24
white-tomentose beneath; inflorescences cymose-paniculate, the pyramidal panicles
usually large and broad, heads numerous, short-pedicellate; involucres 2-3 mm. high;
phyllaries 4-5-seriate, stramineous with dark green midribs and sometimes purple tips,
acuminate, obtuse or acute, almost glabrous, ciliolate; pistillate heads with 18-40
(commonly about 30) filiform pistillate flowers and 1-6 hermaphrodite flowers;
staminate heads with 16-45 (commonly about 20) hermaphrodite flowers; achenes
sparsely hispidulous or glabrous, slightly more than 1 mm. long; pappus sordid, that
of the staminate heads about 2 mm. long, that of the pistillate heads about 3 mm.
long.
Archibaccharis subsessilis Blake, Brittonia 2: 339. 1937.
Wet or dry thickets or forest, often in pine-oak forest, 1,300-
3,000 m.; Alta Verapaz; Huehuetenango; El Quiche (type from
Nebaj, Skutch 1736); San Marcos. Mexico (Chiapas).
Slender, erect or subscandent shrubs, mostly 1-3 m. high, the branches densely
covered with short, spreading, multiseptate hairs; upper leaves sessile, the lower ones
on petioles only 1-3 mm. long, the blades usually rather thick and firm, obovate or
oblong-ovate, mostly 2-8 cm. long, 1-4 cm. wide, short-acuminate, rounded or
narrowly subcordate to subacute at the base, the margins serrulate, scaberulous
above, glandular -punctate beneath and minutely hispidulous at least on the veins
(these commonly elevated and reticulate, especially beneath); inflorescences cymose-
paniculate, the panicles with numerous heads, the slender pedicels mostly 2-10 mm.
long, densely puberulent and sometimes with scattered glands; heads always
heterogamous; involucres 2-3 mm. high; phyllaries lance-oblong to linear, acute or
acuminate, whitish with dark green midribs, often purplish near the apex, puberulent
and glandular; pistillate heads commonly with 17-29 pistillate flowers and 1-7
hermaphrodite flowers; staminate heads with 17-38 hermaphrodite flowers and with
(4-) 12-14 pistillate flowers; achenes mostly trigonous, 1-2.5 mm. long, glandular and
hirsutulous; pappus sordid, that of the pistillate and staminate flowers 3-4 mm. long.
ASTER Linnaeus
Perennial or sometimes annual or biennial herbs, pubescent or glabrous; leaves
alternate, narrow or broad, the margins entire to dentate or rarely pinnatifid;
inflorescences usually racemose or paniculate; heads heterogamous, radiate, the rays
blue, violet, or white, sometimes purple, the disc flowers yellow or sometimes
purplish; ray flowers 1-2-seriate, hermaphrodite, fertile or rarely sterile; disc flowers
hermaphrodite, all or mostly fertile; involucres campanula te or hemispheric;
phyllaries several-seriate, rigid, often scarious-marginate, frequently with herbaceous
tips; receptacles flat or convex, commonly foveolate; ligules of ray flowers usually
spreading, entire or obscurely 2-3-dentate; disc flowers regular, tubular, the long limb
little ampliate, shortly 5-cleft; anthers obtuse and entire at the base; style branches
of the disc flowers complanate, the tips elongated, slender, subulate, lanceolate, or
acutely ovate, papillose or hirtellous; achenes more or less compressed, often with
nervelike margins, the sides often 1-2-costate; pappus bristles slender, scabrous,
numerous, irregularly 2-3-seriate, subequal or the outer ones shorter.
Probably 200 or more species, widely distributed, mostly in
temperate regions, very numerous in temperate North America.
NASH: FLORA OF GUATEMALA 139
Four are native in Guatemala, but five are treated here as A. laevis
is widely planted at almost all elevations. There is also an
unidentified, cultivated species in Totonicapan that has escaped
into fields about 6 miles northeast of Quezaltenango (King 3210);
the place of origin is not known and the specimen has not been
matched in Field Museum.
Plants armed with stout spines; leaves mostly reduced to scales A. spinosus.
Plants unarmed; leaves well developed.
Principal cauline leaves small, mostly 0.8-2.5 cm. long, usually pilose or hirsute,
sometimes merely puberulent A. moranensis.
Principal cauline leaves larger, mostly 3-15 cm. long, glabrous or nearly so.
Leaves, at least the upper ones, cordate-clasping at the base; achenes essentially
glabrous. (Introduced garden ornamentals) .A. laevis.
Leaves narrowed at the base, or if somewhat clasping, always first long-
attenuate and then dilated; achenes pubescent or pilose. (Native plants,
usually of damp or wet places).
Area of inflorescence with remote, much reduced leaves; phyllaries acuminate
or long-acuminate, the margins entire or nearly so, eciliolate.
A. subulatus.
Area of inflorescence usually leafy, dense, often crowded; phyllaries acute, the
margins ciliolate or minutely lacerate .A. bullatus.
Aster bullatus Klatt, Ann. Hofmus. Wien. 9: 359. 1894. Aster
jalapensis Fernald, Proc. Amer. Acad. 35: 572. 1900. Margarita
silvestre (Huehuetenango).
Usually on wet banks along swift streams, sometimes in
crevices of rocks in rivers, rarely in marshes or wet meadows, mostly
at 1,200-1,800 m. (rarely as low as 200 m.); Alta Verapaz; Baja
Verapaz; Chimaltenango; Chiquimula; Huehuetenango; Jalapa;
Peten; El Quiche; Zacapa. Southern Mexico; British Honduras;
Honduras.
Stiff, erect perennials, 10-50 cm. tall, the stems striate, pubescent or essentially
glabrous, simple and usually naked below, sparsely branched and densely leafy above,
the short rootstocks producing numerous, coarse, fleshy-fibrous roots; leaves sessile or
nearly so or the lower ones sometimes narrowing into a short, winged petiole, the
principal leaves commonly 4-11 cm. long and 0.3-1 cm. wide (rarely, in dwarf forms
from lower elevations, the leaves mostly 1-3 cm. long), oblanceolate to nearly elliptic
or linear, sometimes almost spathulate, acute or acuminate, often rather crowded,
long-attenuate to the base and then sometimes more or less dilated and clasping,
often conspicuously ciliate near the base, the margins coarsely serrate or entire,
glabrous or nearly so, reticulate-veined; heads few or numerous, on short or
elongated, sparsely leafy-bracteate peduncles; involucres hemispheric, about 6 mm.
high; phyllaries 3-4-seriate, linear, acute, green above but pale-marginate, more or
less ciliolate or minutely lacerate, glabrous; ray flowers white, about 8 mm. long, the
ligules about 5 mm. long, recurving; achenes pale, pilose, about 1.5 mm. long; pappus
sordid, 4-5 mm. long.
140 FIELDIANA: BOTANY, VOLUME 24
These plants often form dense colonies in the edges of swift
streams.
Aster laevis L. Sp. PL 876. 1753. Li/a; margarita; margari-
tilla.
Native of the United States, where it is widely distributed;
planted commonly for ornament in Guatemalan gardens, at almost
all elevations, and in many other parts of Central America.
Stout, glabrous, erect perennials, commonly 25-60 cm. high, simple below,
branched above; leaves mostly sessile and amplexicaul, lanceolate to oblong-
lanceolate, 3-15 cm. long, acute, the margins mostly entire, sometimes more or less
serrate, scabrous on the margins, the uppermost leaves usually much reduced, the
basal and lowest cauline ones narrowed into winged petioles; inflorescence varying
from thyrsoid-paniculate to open-paniculate; involucres campanulate, 5-8 mm. high;
phyllaries several-seriate, rigid, acute, appressed, green-tipped; ray flowers 15-30, the
ligules 6-10 mm. long, usually blue or violet, rarely white; achenes glabrous or nearly
so; pappus fulvous, 4-6 mm. long.
Called "ramillete" in Yucatan.
Aster moranensis HBK. Nov. Gen. & Sp. 4: 93. 1820.
Diplostephium moranense Nees, Gen. & Sp. Aster. 198. 1832; DC.
Prodr. 5: 273. 1836. A. lima Lindl. in DC. torn. cit. 230. A. bimater
Standl. & Steyerm. Field Mus. Bot. 23: 141. 1944. Margarita de
monte, margarita silvestre (Huehuetenango).
Open, dry, oak or pine-oak forest, 1,400-2,100 m.; Huehue-
tenango; El Quiche; Solola. Mexico.
Erect perennials from thick, woody roots, the stems 1-several, slender, simple
below, racemose-paniculate above, often somewhat glandular, especially in the area
of inflorescence (the glands not always evident due to the often dense indument),
pilose or hirsute with spreading hairs or sometimes strigose or pubescent to
puberulent with appressed hairs, rarely the hairs very short or vestigial and the
glands then conspicuous; leaves sessile, linear or oblong-linear, mostly 8-25 mm. long,
1-3 mm. wide, acute or obtuse and mucronulate, the margins entire, obtuse at the
base, usually densely hirsutulous on both surfaces with spreading, ascending, or
appressed hairs; inflorescence laxly racemose-paniculate, the branches slender,
ascending, each with a single head, sparsely or densely leafy, the peduncles short or
elongated, glandular-puberulent; involucres hemispheric, 5-7 mm. high, 8-10 mm.
broad; phyllaries few-seriate, linear to narrowly lanceolate, subobtuse, commonly
mucronulate, glandular-puberulent or rarely almost glabrous, more or less ciliate,
whitish below, green above, the tips somewhat spreading; ray flowers 15-20, white,
the ligules 6-8 mm. long, 1-1.5 mm. wide; disc flowers 30-40, yellow, 5-6 mm. long;
achenes commonly 1.5-2.5 mm. long (rarely to 4 mm.), obscurely or prominently
costate, sparsely or densely hispidulous; principal pappus bristles 3-6 mm. long.
A variable complex, the several forms differing principally in
indument and intergrading, the differences not explained by any
NASH: FLORA OF GUATEMALA 141
geographical or ecological variations. Several specimens from
Mexico (including Chiapas) have strongly appressed or ascending
hairs on both stems and leaves. Many specimens from Mexico and
Guatemala have only spreading hairs. Some Guatemalan specimens
have spreading stem hairs but ascending or somewhat appressed
leaf hairs. A few specimens from Chiapas and from Huehuetenango
have only vestigial hairs but are strongly glandular, especially on
the peduncles. Two of these (Breedlove 6011 and 7294 from
Chiapas) are notable for their very long (4 mm.) achenes that are
very prominently costate.
Aster spinosus Benth. PL Hartweg. 20. 1839. Espinaza, espina
de agua (Santa Rosa, fide Aguilar). Figure 30.
Among rocks or in sand or gravel along, near, or in streams,
sometimes on large rocks in streams and even partly submerged,
225-1,350 m.; Baja Verapaz; Chiquimula; Jalapa; Jutiapa; El
Progreso; El Quiche; Santa Rosa; Zacapa. Southwestern United
States; Mexico; Honduras; Costa Rica; Nicaragua.
Erect, stout, densely branched, glabrous, perennial herbs or subshrubs to 1 m.
tall, usually appearing leafless or nearly so, the stems pale green, striate, armed with
numerous stout, usually somewhat compressed, sharp spines 0.5-4 (-7) cm. long;
leaves few, linear and bractlike or reduced to scales; heads numerous, solitary on
slender or stout, short or elongated, inconspicuously bracteate peduncles; involucres
hemispheric, 4-5 mm. high; phyllaries 4-5-seriate, stramineous or greenish with pale,
erose margins, glabrous, acute or obtuse, the outer ones short; ray flowers white, 5-6
mm. long, the ligules about 3 mm. long, spreading or often revolute, 2-3-dentate;
achenes pale, glabrous, about 2.5 mm. long; pappus sordid, about 3 mm. long.
Common plants along streambeds at low elevations and chiefly
in the drier regions, often forming large, dense colonies which from
a distance look like beds of coarse grass.
Aster subulatus Michx. Fl. Bor. Am. 2: 111. 1803. A. exilis Ell.
Bot. S. Carol. & Ga. 2: 344. 1823. Conyza squamata Spreng. Syst. 3:
515. 1826. A. exilis var. australis A. Gray, Syn. Fl. N. Am. 1(2): 203.
1884. A. squamatus Hieron. Engl. Bot. Jahrb. 29: 19. 1900. A.
subulatus var. australis Shinners, Field & Lab. 21: 158. 1953.
Often on sandbars along streams, in open marshes, damp or wet
thickets or fields, sometimes on rocks in streams, in coniferous or
mixed forest, occasionally a weed in waste or cultivated ground,
200-2,800 m.; Alta Verapaz; Baja Verapaz; Chimaltenango; Chi-
quimula; Escuintla; Guatemala; Huehuetenango; Jalapa; Jutiapa;
Peten; Quezaltenango; Sacatepequez; San Marcos; Solola; Zacapa.
142 FIELDIANA: BOTANY, VOLUME 24
Southern United States; Mexico; British Honduras; Honduras;
Costa Rica; Nicaragua; West Indies; northern South America.
Erect annuals, mostly 25-75 cm. tall, simple below, openly branched above, the
stems and branches glabrous or nearly so; principal cauline leaves usually scattered
or remote, sessile, or the lower ones narrowed into slender petioles, the blades
narrowly linear to linear-oblanceolate, narrowly elliptic, or spathulate, mostly 3-15
cm. long, 0.2-1.5 cm. wide, those in the area of the inflorescence usually much
reduced, acute or long-acuminate, the margins entire or the lower ones sometimes
with serrate margins, glabrous or nearly so; inflorescences racemose to open-
paniculate, the heads on slender, bracteate pedicels; involucres hemispheric-
campanulate, 4-6 (-8) mm. high; phyllaries about 3-seriate, linear, acuminate or long-
acuminate, entire or nearly so, essentially glabrous, pale greenish, at least the upper
part, the margins stramineous, hyaline, the tips sometimes somewhat purplish; ray
flowers 5-6 mm. long, commonly white (in ours), sometimes bluish or lavender, the
ligules about 3 mm. long, often revolute; disc corollas yellow, about 3.5 mm. long;
achenes pubescent, about 1.5 mm. long; pappus fine, white, about 3 mm. long.
ASTRANTHIUM Nuttall
Reference: D.C.D. De Jong, A systematic study of the genus
Astranthium (Compositae, Astereae), Publ. Mich. State Univ. Biol.
2: 429-528. 1965.
Erect, ascending, or decumbent perennials or biennials usually forming basal
rosettes, sometimes annuals, the stems solitary or many, striate, more or less strigose;
cauline leaves alternate, sessile, spathulate to linear, reduced upward, the margins
usually entire, rarely dentate, more or less ciliate; heads often drooping, commonly
long-pedunculate; involucres campanulate or hemispheric; phyllaries 2-3-seriate,
subequal, ovate to linear-lanceolate, the margins scarious; receptacle conical,
alveolate; ray flowers uniseriate, pistillate, fertile, 10-35, white or lavender, the tube
more or less puberulent, the ligules conspicuous, 2-4 -denticulate; disc flowers
hermaphrodite, 40-260, yellow, the corollas usually abruptly contracted at the throat;
anthers obtuse at the base, apically appendaged; tips of the style branches of the disc
flowers ovate to ovate-lanceolate; achenes laterally compressed, obovate to
oblanceolate, glabrous or glochidiate-pubescent, obscurely 2-nerved to conspicuously
2-3-costate; pappus a minute ring or crown of setae less than 0.1 mm. high, or absent.
Ten species, ranging from Kentucky, in the United States, to
the mountains of Oaxaca and Chiapas, in Mexico. Although none
have been found in Guatemala, the species that occurs in Chiapas is
treated here.
Astranthium purpurascens (Robins.) Larsen, Ann. Mo. Bot.
Gard. 23: 33. 1933. Bellis purpurascens Robins. Proc. Am. Acad. 27:
172. 1892. Figure 31.
Not reported from Guatemala but may be expected there, as
several collections have been made in adjacent Chiapas, Mexico,
1,700-2,300 m.
NASH: FLORA OF GUATEMALA 143
Erect, ascending, or rarely decumbent perennials mostly 10-60 cm. high, the
stems 1-many, simple or sparsely branched, glabrous to densely strigose; basal leaves
narrowly or broadly spathulate, obtuse or mucronate, 2-10 cm. long, usually
narrowing into a winged petiole, essentially glabrous to pubescent, lower cauline
leaves often very similar but the middle and upper leaves progressively reduced in
size, 0.5-6 cm. long, sessile, often becoming obovate to lance-elliptic; heads solitary on
peduncles to 20 cm. long, these more or less pubescent, more densely so just below
the heads; involucres campanulate, 3-7 mm. high; phyllaries 2-3-seriate, broadly
ovate to linear-lanceolate or oblanceolate, greenish with conspicuous to narrow, often
purplish, more or less lacerate-ciliate margins, glabrous or sparsely pubescent; ray
flowers 10-30, white, the ligules 9-14 mm. long, 2-3 mm. wide, 2-3-dentate or
denticulate; disc flowers 40-155, mostly 3-4.5 mm. long; achenes brownish, obovate,
more or less compressed, glabrous to sparsely glochidiate-pubescent, 1.5-2.3 mm. long,
2-nerved; pappus absent or reduced to a minute, pale crown, with a few teeth less
than 0.1 mm. high.
BACCHARIS Linnaeus
Reference: Jose Cuatrecasas, Revision de las especies
Colombianas del genero Baccharis, Rev. Acad. Colomb. Cienc. 13:
5-102. 1967; Notas adicionales, taxonomicas y corologicas sobre
Baccharis, op. cit. 201-226. 1968.
Shrubs or small trees, rarely perennial herbs, dioecious, rarely polygamous;
leaves alternate, often glutinous, the margins of the blades entire, dentate, or serrate;
inflorescences cymose or paniculate; heads discoid; involucres campanulate to ovoid
or subglobse; phyllaries multiseriate, imbricate, dry, sometimes with a green or
purplish costa, scarious- margined, often glutinous; receptacle flat, naked or rarely
paleaceous; corollas of the pistillate heads tubular-filiform, the limb truncate; flowers
of the staminate heads hermaphrodite but sterile, the corollas tubular, the limb 5-
dentate; anthers obtuse and entire at the base; style branches narrow or subulate;
achenes 5-10-costate, glabrous or pubescent; pappus uniseriate or biseriate (rarely
more), whitish or brownish, that of the pistillate flowers soft, persistent, that of the
staminate flowers stiffer, the bristles sometimes a little dilated above.
One of the largest genera of Compositae, with about 400
species, all American and most abundant in South America. Seven
species are found in Guatemala.
Margins of leaf blades entire or occasionally with 1-5 large teeth near the apex.
Leaves mostly 5-12 cm. long, conspicuously triplinerved.
Petioles 1-2.5 cm. long; pedicels glabrous B. pedunculata.
Petioles less than 1 cm. long pedicels pilosulous B. trinervis.
Leaves mostly 1-5 cm. long, the lateral nerves often obscure.
Leaf blades oblanceolate to spathulate, 2-5 cm. long, more than twice as long as
broad, often with 3-4 large teeth near the apex B. heterophylla.
Leaf blades obovate to elliptic, 1-2.5 cm. long, usually less than twice as long as
broad, entire, except on vigorous sterile branches rarely coarsely dentate
near the apex B. vaccinioides.
Margins of leaf blades serrate.
144 FIELDIANA: BOTANY, VOLUME 24
Leaf blades linear to narrowly linear-oblanceolate, mostly 5-12 cm. long; pedicels
essentially glabrous B. sahcifolia.
Leaf blades broadly oblanceolate to rather narrowly elliptic or oblong-elliptic,
mostly 2-6 cm. long; pedicels pilosulous.
Involucres of pistillate heads 4-5 mm. high; pistillate flowers commonly 18-26
(-32) per head B. serraefolia.
Involucres of pistillate heads 5-6 mm. high; pistillate flowers commonly 36-39 per
head B. prorepens.
Baccharis heterophylla HBK. Nov. Gen. & Sp. 4: 62. 1820. B.
cuneata A. DC. in DC. Prodr. 5: 408. 1836. B. spathulata Schauer,
Linnaea 19: 724. 1845. Arrayan (Huehuetenango).
Damp, brushy hillsides, often in oak and/or pine forest,
frequently on sandbars along stream beds, 1,500-2,100 m.; Huehue-
tenango; Jalapa. Southern Mexico.
Erect, densely branched shrubs, glutinous throughout, 1-3 (-4) m. tall, the
branches stiff, brownish, striate, densely leafy, glabrous or nearly so; leaves sessile,
subsessile, or very short-pedicellate, the blades oblanceolate, oblong-oblanceolate, or
spathulate, mostly 2-5 cm. long, 0.5-2 cm. wide, obtuse or rounded at the apex or
rarely subacute, attenuate to the base, coriaceous, lustrous and glutinous, the
margins entire or with a few coarse, irregular teeth toward the apex, or the leaves of
juvenile shoots often conspicuously and coarsely dentate, 1-nerved or obscurely
triplinerved; heads small, numerous, subsessile, disposed in dense, rounded clusters at
the ends of the branches; involucres about 3 mm. high; phyllaries pale greenish-white
to stramineous, with a purplish midrib and/or tip, glabrous or nearly so, ovate to
lanceolate, erose and ciliate; achenes glabrous, about 10-costate; pappus of the
staminate flowers stramineous, about 3 mm. long, that of the pistillate flowers 4-6
mm. long.
This species is quite similar in appearance and in habitat to the
Mexican B. conferta HBK. (which may prove to be synonymous
with the South American B. tricuneata (L. f. ) Persoon). The
Mexican and South American specimens I have seen appear to have
consistently smaller leaves than B. heterophylla, 1-2 cm. long, 0.4-1
cm. wide, commonly with more consistently dentate margins.
Baccharis pedunculata (Mill.) Cabrera, Bol. Soc. Arg. Bot. 7:
240. 1959; Cuatrecasas, Rev. Acad. Colomb. Cienc. 13: 48. 1967.
Conyza pedunculata Miller, Gard. Diet. ed. 8: 15. 1768. B.
cinnamomifolia HBK. Nov. Gen. & Sp. 5: 65. 1820. B. speciosa A.
DC. in DC. Prodr. 5: 399. 1836. Eupatorium braunii Polak, Linnaea
41: 576. 1877. B. splendens Heering, Schrift. Natur. W. Vereins.
Scheswig. Hoist. 13: 48. 1906. B. braunii Standley, Field Mus. Bot.
18: 1433. 1938.
In rocky, brushy streambeds, wet thickets, or on roadside banks
or brushy hillsides, 500-1,650 m.; Alta Verapaz; Escuintla; Guate-
NASH: FLORA OF GUATEMALA 145
mala; Quezaltenango; El Quiche. Honduras to Panama; West
Indies; South America.
Erect shrubs or small trees, 1.5-2.5 (-4) m. tall, essentially glabrous throughout,
very glutinous, the branches striate, rather densely leafy; leaves on petioles 1-2.5 cm.
long, the blades lance-elliptic or oblong-elliptic, mostly 5-12 cm. long, 2-5 cm. wide,
subacute and abruptly cuspidate, cuneate at the base, conspicuously triplinerved,
very lustrous on the upper surface, the margins entire; inflorescences terminal,
appearing pedunculate, the heads slender-pedicellate, disposed in corymbose panicles;
involucres hemispheric to broadly campanulate, 4-6 mm. high; phyllaries 5-6-seriate,
greenish white, usually tinged or tipped with purple, ovate to lanceolate, acute or
subacute, the margins erose, scarious, ciliate; achenes glabrous, 5-costate, about 2
mm. long; pappus dirty white to pale stramineous, uniseriate, 6-8 mm. long in the
pistillate flowers, about 4 mm. long in the staminate flowers.
Baccharis prorepens (Blake) Jackson, Taxon 19: 263. 1970.
Archibaccharis prorepens Blake, Journ. Wash. Acad. Sci. 24: 432.
1934. Te de montaha.
Dry banks, open, often damp forest, 2,400-2,700 m., Chimal-
tenango (type from Santa Elena, Skutch 190).
Erect, suffruticose, branching plants 1-1.5 m. high, the stems angled, striate, more
or less pilosulous with crisped hairs; leaves on petioles 1-5 mm. long, the blades
oblong-elliptic, lanceolate, or obovate, 2.5-7.5 cm. long, acute, cuneate at the base,
the margins sharply serrate, essentially glabrous above except pilosulous along the
costa, more or less puberulent below, obscurely or conspicuously triplinerved;
inflorescences cymose-paniculate; involucres 5-6 mm. high; phyllaries 3-4-seriate,
oblong to linear, acute to subobtuse, glabrous or nearly so, sometimes sparsely
glandular, the costae green, the margins subscarious, more or less ciliate, especially
near the apex; filiform flowers of the pistillate heads commonly 36-39 per head,
greenish white or purple near the apex; hermaphrodite disc flowers of the staminate
heads commonly 15-23 per head; achenes ovate to oblong, 5-8-costate, brown, dull,
glabrous, those of the pistillate flowers 1.2-1.5 mm. long; pappus white, 3-4 mm. long.
Baccharis salicifolia (R. & P.) Persoon, Syn. PI. 2: 425. 1807;
Cuatrecasas, Rev. Acad. Colomb. Cienc. 13: 204. 1968. Molina
salicifolia Ruiz & Pavon, Syst. Veg. 210. 1798. B. glutinosa Persoon,
Syn. PI. 2: 425. 1807. B. coerulescens A. DC. in DC. Prodr. 5: 402.
1836. B. longifolia A. DC. I.e. B. linifolia A. DC. torn. cit. 420. B.
salicifolia var. longifolia Cuatr. Rev. Acad. Colomb. Cienc. 13: 212.
1968. Chilca.
Usually confined to sandy or rocky thickets along stream beds,
sometimes on damp plains or in fields, near sea level to 2,800 m.;
Alta Verapaz; Baja Verapaz; Chimaltenango; Chiquimula; Es-
cuintla; Guatemala; Huehuetenango; Izabal; Jalapa; Jutiapa; El
Progreso; Quezaltenango; Sacatepequez; San Marcos; Santa Rosa;
Solola; Zacapa. Southwestern United States; Mexico; El Salvador;
Honduras; South America.
146 FIELDIANA: BOTANY. VOLUME 24
Erect shrubs or small trees, rarely as much as 6 m. tall, glutinous throughout,
glabrous or nearly so; the young stems and twigs striate to somewhat alate and
sometimes appearing angulate; leaves short-petiolate or subsessile, the blades linear
or narrowly linear-oblanceolate, mostly 5-12 cm. long, 0.3-1.5 cm. wide, long-
attenuate to each end, triplinerved but usually not conspicuously so, the margins
rather remotely but evenly serrate or serrulate; inflorescences terminal, corymbose-
paniculate, the numerous heads pedicellate, many-flowered; involucres stramineous,
3-5 mm. high; phyllaries appressed, pale with a darker midrib, obtuse or subacute,
erose and scarious-margined; achenes oblong, narrowing at the base, glabrous, 5-
costate, scarcely more than 1 mm. long; pappus uniseriate, dirty white to yellowish,
that of the pistillate flowers 4-5 mm. long, that of the hermaphrodite flowers about 3
mm. long.
One of the most common and characteristic shrubs of sandy or
rocky river bottoms, frequently forming wide thickets, sometimes in
association with willows (Salix).
Baccharis serraefolia A. DC. in DC. Prodr. 5: 403. 1836. B.
parviflora Less, in Schlecht. & Cham. Linnaea 5: 146. 1830, not B.
parviflora Pers. 1807. B. kellermanii Greenm. Field Mus. Bot. 2:
267. 1907 (type from Volcan de Atitlan, Solola, Kellerman 5356).
Te; te de monte.
Open or brushy slopes or open forest, in dry to wet situations,
frequently in pine-oak forest or on open, rocky banks, 1,200-3,000
m.; Alta Verapaz; Baja Verapaz; Chimaltenango; Escuintla;
Guatemala; Huehuetenango; Jalapa; Quezaltenango; El Quiche;
Sacatepequez; San Marcos; Solola; Totonicapan. Mexico; El
Salvador; Honduras; Nicaragua.
Erect shrubs or suffruticose plants, 1-2 m. high, simple or sparsely branched, the
stems striate, glutinous, when young pilosulous with crisped hairs, becoming glabrate
below; leaves short-petiolate, the blades thick and rather rigid, broadly oblanceolate
to rather narrowly elliptic or oblong-elliptic, mostly 2-5 cm. long, obtuse or rounded
at the apex and frequently mucronulate, cuneate or attenuate to the base, the
margins coarsely and evely serrate, somewhat triplinerved, villosulous or glabrate
beneath, the veins prominent; inflorescences cymose, usually forming small or large,
rounded, terminal panicles, the heads on slender, pilosulous pedicels; involucres 3-5
mm. high; phyllaries pale with a darker midrib, linear-oblong, obtuse or subobtuse,
glabrous or more or less puberulent, ciliate; pistillate heads commonly with 18-26
(-32) flowers; hermaphrodite ones commonly with 15-20 flowers but rarely only 6-10;
achenes oblong, 5-costate, glabrous, 1-2 mm. long; pappus uniseriate, brownish white,
that of the pistillate flowers about 4 mm. long, of the hermaphrodite flowers about 3
mm. long.
This species is well known in Guatemala as the leaves and
stems are commonly employed in a decoction as a substitute for
coffee.
Baccharis trinervis (Lam.) Persoon, Syn. PI. 2: 423. 1807;
NASH: FLORA OF GUATEMALA 147
Cuatrecasas, Rev. Acad. Colomb. Cienc. 13: 49. 1967. Conyza
trinervis Lam. Encycl. 2: 85. 1786. B. rhexioides HBK. Nov. Gen. &
Sp. 4: 66. 1820. B. trichoclada A. DC. in DC. Prodr. 5: 400. 1836. B.
trinervis var. rhexioides Baker, Fl. Bras. 6 (3): 73. 1844. Arnica;
chilca (Escuintla); barba fina (Izabal); crucito (Jalapa): bisib,
bisik'am, t'isib (quecchi, Alta Verapaz); Santo Domingo (Huehue-
tenango).
Damp or wet thickets, frequent in pine-oak forest, sea level to
2,200 m.; Alta Verapaz; Baja Verapaz; Chimaltenango; Escuintla;
Guatemala; Huehuetenango; Izabal; Jalapa; Jutiapa; Peten; Que-
zaltenango; El Quiche; Sacatepequez; Santa Rosa; Solola; Suchite-
pequez; Zacapa. Southern Mexico; British Honduras; El Salvador
to Panama; tropical South America.
Usually dense shrubs to about 3 m. tall (rarely to 6 m.) erect or often with
arching branches sometimes 5 m. long and supported on other plants, the stems
striate and angulate, glutinous and usually glabrate, sometimes sordid-villosulous or
tomentulose, especially when young; leaves subsessile or on very short petioles less
than 1 cm. long, the blades mostly lance-oblong to elliptic, 5-10 cm. long, 0.5-3.5 cm.
wide, commonly acuminate to long-acuminate but sometimes acute, cuneate at the
base, conspicuously triplinerved, usually lustrous above, the margins entire; area of
inflorescence broad, the heads sessile or on short, pilosulous pedicels, disposed in
short, dense panicles at the ends of the branches; involucres 3.5-5.5 mm. high (those
of the staminate flowers usually not more than 4 mm. high); phyllaries 4-5-seriate,
ovate to oblong-lanceolate, acute or obtuse, erose, pale yellowish to light greenish,
with a darker midrib, glandular-puberulent to nearly glabrous, the margins scarious,
ciliate; achenes pilose, about 1 mm. long; pappus uniseriate, 3-4 mm. long, whitish to
pale yellowish.
Baccharis vaccinioides HBK. Nov. Gen. & Sp. 4: 50. 1820. B.
land folia Less. Linnaea 9: 266. 1834. Array an; limoncillo; raijan;
raijan cachump. Figure 32.
Damp or dry hillsides, brushy pastures, open broadleaf and
coniferous forest, 1,500-3,900 m.; Chimaltenango; Guatemala;
Huehuetenango; Jalapa; El Progreso; Quezaltenango; El Quiche;
Sacatepequez; San Marcos; Solola; Totonicapan; Zacapa. Moun-
tains of southern Mexico; El Salvador, Honduras.
Shrubs or small trees, usually less than 6 m. tall, with short trunks and very
dense, rounded crowns, essentially glabrous throughout, very glutinous; leaves sessile
or short-petiolate, the blades obovate or elliptic, mostly 1-2.5 cm. long, 0.6-1.5 cm.
wide, obtuse or subacute, sometimes apiculate, acute at the base, often lustrous
above, the margins entire, except the leaves of vigorous sterile branches rarely
coarsely dentate near the apex; heads subsessile, disposed in small, dense clusters at
the ends of the branches; involucres 3-4 mm. high; phyllaries about 5-seriate,
greenish white or cream, ovate to lanceolate or the innermost ones linear-lanceolate
or linear-oblong, tomentulose or glabrous, ciliolate; achenes pale, glabrous, 5-costate,
148 FIELDIANA: BOTANY, VOLUME 24
about 1 mm. long; pappus yellowish white, that of the pistillate flowers about 5 mm.
long, of the hermaphrodite flowers about 3 mm. long.
This species is one of the most typical and tiresomely abundant
shrubs or small trees of the Guatemalan mountains, often forming
exceedingly dense stands of great extent. These stands constitute
distinct plant formations on the higher slopes of the volcanoes and
in the Sierra de los Cuchumatanes. They are often so dense that
little else grows in them. On the Volcan de Santa Maria B.
vaccinioides forms a distinct belt just at the base of the cone
proper. Little else but bunchgrass is found in this broad belt. The
small thick leaves adhere persistently to the dried branches, which
often are used in bunches as rough brooms.
CONYZA Linnaeus
References: Arthur Cronquist, The separation of Erigeron from
Conyza, Bull. Torr. Bot. Club 70: 629-632. 1943. Jose Cuatrecasas,
Notes on neotropical Compositae, I, Phytologia 9: 1-7. 1963.
Annual or biennial (rarely perennial) herbs, usually more or less pubescent;
leaves alternate, dentate, crenate, dissected, or entire; heads heterogamous, usually
numerous and disposed in corymbose or racemose panicles, sometimes forming dense
spikes, rarely solitary or few at the ends of the branches; involucres more or less
campanulate; phyllaries 2-many-seriate, unequal, imbricate, linear or linear-
lanceolate, the outer ones shorter; receptacles flat or convex, naked or the margins of
the foveae fimbriate, produced as very short pales; pistillate flowers numerous, the
corollas filiform, the ligules minute and inconspicuous or none, shorter than the tubes
and scarcely if at all exceeding the pappus; central disc flowers few, hermaphrodite,
tubular, little ampliate, the limb 5-dentate, all or most of the flowers fertile; anthers
obtuse at the base; style branches of the hermaphrodite flowers complanate, the
appendages lanceolate, short or elongated; achenes small, compressed, the sides
commonly one-nerved; pappus uniseriate or rarely biseriate, the outer bristles shorter,
unequal.
Probably less than 50 species, in the tropics of both
hemispheres, a very few extending into temperate regions. There are
eight in Guatemala, but 10 are treated here, as two occurring in
nearby Chiapas, Mexico may also be expected in Guatemala.
Stems conspicuously lanate with white, more or less matted hairs.
Inflorescence commonly densely spikelike (sometimes densely pyramidal-
paniculate); heads very numerous.
Involucres more or less lanate, eglandular; pistillate flowers 40-50 per head; rays
none; achenes densely sericeous-pilose; pappus biseriate C. filaginoides.
Involucres villous-pilose, glandular; pistillate flowers 200-300 per head; rays
present but minute; achenes pubescent or glabrous; pappus uniseriate.
C. schiediana.
Inflorescence commonly laxly racemose or paniculate (rarely spikelike but then not
densely so); heads relatively few C. confusa.
NASH: FLORA OF GUATEMALA 149
Stems not lanate, the indument chiefly of straight hairs.
Leaves (at least the lower ones) pinnatifid or bipinnatifid.
Principal cauline leaves commonly pinnate-lobate with few lobes (rarely merely
coarsely crenate); involucres 4-5 mm. high; denuded receptacles mostly 4-6
mm. wide; pistillate flowers more than 200 per head C. coronopifolia.
Principal cauline leaves commonly bipinnatifid with numerous linear segments;
involucres 2-3 mm. high; denuded receptacles only 1.5-2 mm. wide; pistillate
flowers 75-110 per head C. sophiifolia.
Leaves never pinnatifid or bipinnatifid.
Plants perennial or biennial, the large basal leaves persistent; inflorescence
composed of few heads clustered at ends of sterns; involucres 6-8 mm. high;
pistillate flowers about 380 per head C. chilensis.
Plants annual, lacking basal leaves; inflorescence composed of usually numerous
heads, forming racemose or open panicles; involucres 2.5-5 mm. high;
pistillate flowers 40-240 per head.
Heads very small, the involucres mostly 2.5-3.5 mm. high, glabrous; denuded
receptacles only 1.2-2.5 mm. wide; pistillate flowers 40-50 per head;
achenes essentially glabrous C. canadensis.
Heads larger, the involucres 3.5-6 mm. high, hirsute or hispidulous or
sometimes glabrate; denuded receptacles commonly 3-5 mm. wide;
pistillate flowers 60-240 per head; achenes more or less villous.
Lower leaves oblong-ovate or spathulate, acute or obtuse, rather abruptly
narrowed to a winged petiole; involucres about 3.5 mm. high; pistillate
flowers 200-240 per head; pappus about 2 mm. long C. apurensis.
Lower leaves oblanceolate, acuminate, attenuate to a sessile base; involucres
mostly 4-5 mm. high; pistillate flowers commonly 60-180 per head;
pappus 3-4 mm. long C. bonariensis.
Conyza apurensis HBK. Nov. Gen. & Sp. 4: 73. 1820.
Erigeron spathulatus Vahl in West, Bidr. Beskr. St. Croix 303. 1793,
not C. spathulata Hornem. 1807, nor E. spathulatum Wedd. 1857.
E, chinensis Jacq. PI. Hort. Schoenbr. 3: 30, t. 303. 1798. E.
apurensis Griseb. Fl. Brit. W. Ind. 365. 1861. C. subspathulata
Cronquist, Bull. Torr. Bot. Club 70: 632. 1943.
Wet to dry thickets or open fields, sandbars along streams,
sometimes in open, oak-pine or pine forest, often a weed in
cultivated or waste ground, near sea level to 2,800 m.; Alta Verapaz;
Guatemala; Izabal; Jalapa; Jutiapa; Peten; El Progreso; Quezal-
tenango; El Quiche; Retalhuleu; Sacatepequez; Santa Rosa;
Totonicapan; Zacapa. Mexico and British Honduras to El Salvador
and Panama; West Indies; South America.
Erect annuals, commonly 0.3-1.5 m. tall, often much branched, usually more or
less hirsute throughout, especially on stems and leaves, sometimes short-pilose, rarely
glabrate; lower leaves oblong-ovate or spathulate, acute or obtuse, mostly 4-10 cm.
long including the petiole, the blades 2-3 cm. broad, rather abruptly narrowed to a
winged petiole, the margins coarsely dentate or crenate, middle and upper leaves
smaller, the upper ones 2-4 cm. long, sessile or nearly so, oblong, linear-oblong, or
150 FIELDIANA: BOTANY, VOLUME 24
narrowly elliptic, acute or obtuse, the margins entire or dentate; inflorescence laxly
corymbose-paniculate, the heads on slender pedicels mostly 1-4 cm. long; involucres
about 3.5 mm. high; phyllaries stramineous with a green or purplish midrib, subequal,
linear, acuminate, hispidulous or glabrate; denuded receptacles commonly 3-5 mm.
wide; pistillate flowers 200-240 per head, the ligules small and inconspicuous or
obsolete; achenes scarcely 1 mm. long, pale, sparsely appressed-villous or glabrate;
pappus uniseriate, sordid, about 2 mm. long.
Conyza bonariensis (L.) Cronquist, Bull. Torr. Bot. Club 70:
632. 1943. Erigeron bonariensis L. Sp. PL 863. 1753. E. linifolius
Willd. Sp. PL 3: 1955. 1804. C. floribunda HBK. Nov. Gen. & Sp. 4:
73. 1820. E. bonariensis var. leiothecus Blake, Contr. Gray Herb. 52:
28. 1917. Marsea bonariensis Badillo, Bol. Soc. Venez. Cienc. Nat.
10: 256. 1946. M. bonariensis var. leiotheca Badillo, I.e. C.
bonariensis var. leiotheca Cuatr. Phytologia 9: 5. 1963. Figure 33.
Damp thickets, fields, or open forest, often in oak or pine-oak
forest, or a weed in cultivated or waste ground, 500-3,400 m.; Alta
Verapaz; Baja Verapaz; Chimaltenango; Huehuetenango; Izabal;
Jalapa; Jutiapa; Peten; Quezaltenango; El Quiche; Sacatepequez;
Santa Rosa; Suchitepequez; probably also in other departments.
Southern United States; Mexico; Honduras to Panama; West
Indies; South America.
Erect annuals, 0.5-2 m. tall, simple or sparsely branched, rather densely leafy,
the stems striate, sparsely or densely hirsute or hispidulous or sometimes glabrate;
lower leaves rather narrowly lanceolate or oblanceolate, mostly 6-15 cm. long, 0.5-2
cm. wide, acuminate, attenuate to a sessile base, dentate or entire; upper leaves
smaller, mostly linear and entire; inflorescence usually much branched, racemose-
paniculate or broadly paniculate, the heads usually numerous; involucres pale
greenish, 4-5 mm. high; phyllaries linear, acuminate, hirsute, villous, hispidulous, or
sometimes glabrous or glabrate; denuded receptacles 3-5 mm. broad; pistillate flowers
60-180 per head, the ligules very short; achenes pale, about 1.3 mm. long, sparsely
villous; pappus uniseriate, sordid, 3-4 mm. long.
A pantropical weed.
Conyza canadensis (L.) Cronquist, Bull. Torr. Bot. Club 70:
632. 1943. Erigeron canadensis L. Sp. PL 863. 1753. E. pusillus
Nutt. Gen. PL 2: 148. 1818, not C. pusilla HBK. 1820. C. parva
Cronquist, Bull. Torr. Bot. Club 70: 632. 1943. C. canadensis var.
pusilla Cronquist, op. cit. 74: 150. 1947.
Damp fields, thickets, sandbars, along rocky streambeds, in oak
or pine-oak forest, often weeds in cultivated or waste ground, 1,000-
2,700 m.; Alta Verapaz; Baja Verapaz; Chimaltenango; Guatemala;
Huehuetenango; Izabal; Jalapa; Peten; Quezaltenango; Sacatepe-
quez; San Marcos; Santa Rosa; Solola. Widely distributed in North
NASH: FLORA OF GUATEMALA 151
America, southward through Mexico; British Honduras and
Honduras to Costa Rica; South America; introduced in the Old
World tropics.
Erect annuals, mostly 0.3-2 m. tall, simple below the area of inflorescence, the
stems hispid, hirsute or glabrous; lowermost leaves petiolate, narrowly oblanceolate,
remotely incised-dentate, the principal and upper leaves sessile, linear and entire,
commonly ciliate and hispidulous, sometimes glabrous; inflorescence racemose or
paniculate, the heads short-pedicellate; involucres mostly 2.5-3.5 mm. high, 7-8 mm.
in diameter (expanded); phyllaries pale greenish white, unequal, linear, acute,
glabrous or nearly so; denuded receptacles 1.2-2.5 mm. in diameter; pistillate flowers
40-50 per head, the ligules minute, shorter than the pappus; achenes pale, about 1
mm. long, essentially glabrous; pappus uniseriate, sordid, 2-2.5 mm. long.
An abundant weed in temperate regions of North America but
seldom plentiful in Central America, although widely distributed.
Conyza chilensis Spreng. Nov. Prov. Hal. 1: 14. 1819.
Erigeron chilensis D. Don, Hort. Brit.: 343. 1830. Marsea chilensis
Badillo, Bol. Soc. Venez. Cienc. Nat. 10: 257. 1946.
Open, grassy slopes, damp thickets or fields, 750-1,800 m.; Alta
Verapaz; Baja Verapaz; Chiquimula; Guatemala; Huehuetenango;
Jalapa; Santa Rosa; Zacapa. Southern Mexico; El Salvador to
Panama; South America.
Erect biennial or perennial herbs, mostly 25-75 cm. tall, eglandular, the stems
stout, simple below, sometimes with a few erect branches above, usually densely
short-pilose with mostly appressed or erect hairs; basal leaves several and usually
persistent, narrowing to winged petioles, oblanceolate, obtuse, usually apiculate, 5-20
cm. long, thinly scaberulous or appressed-pilose, the margins undulate-crenate,
shallowly undulate-lobate, or essentially entire; cauline leaves few, sessile, the lower
ones oblanceolate or linear-oblanceolate, the upper ones linear, mostly 1-6 cm. long,
remotely crenate to entire; heads few, usually clustered at the end of the stem or its
branches, short-pedicellate or subsessile, mostly 10-16 mm. broad; involucres 6-8 mm.
high; phyllaries very unequal, greenish, often with purplish tips, linear, acuminate,
appressed-pilosulous or scabrous; denuded receptacles commonly 5-6 mm. broad;
pistillate flowers about 380 per head, 3-4 mm. long; achenes pale brown, glabrous, 1.5-
2 mm. long; pappus uniseriate, sordid, about 5 mm. long.
Conyza confusa Cronquist, Bull Torr. Bot. Club 70: 632. 1943.
Erigeron gnaphalioides HBK. Nov. Gen. & Sp. 4: 88, t. 331. 1820,
not C. gnaphalioides HBK. 1820.
Not yet reported from Guatemala but included here as there
are several collections from Chiapas, Mexico, 1,500-2,400 m. North
and south Mexico.
Erect plants mostly 20-60 cm. tall, the stems usually 1-3, white-lanate; leaves
sessile, white-lanate on both surfaces, more densely so beneath, or sometimes
essentially glabrous above, the basal leaves spathulate, obtuse and commonly
152 FIELDIANA: BOTANY, VOLUME 24
apiculate, mostly 2.5-8 cm. long, 0.4-1 cm. wide, the margins entire or sometimes
coarsely dentate near the apex; cauline leaves mostly 1-3 cm. long, clasping, linear to
linear-obovate, obtuse; area of inflorescence commonly appearing laxly racemose or
paniculate with relatively few heads but occasionally the heads more numerous and
on shorter pedicels and the inflorescence then spikelike; involucres 4-6 mm. high,
lanate at the base; phyllaries unequal, linear and acuminate, often tipped with
purple, the inner ones essentially glabrous; heads with 90-100 pistillate flowers, the
ligules white, 0.7-2 mm. long, and with about 10 tubular, hermaphrodite disc flowers;
achenes pale, pubescent, about 1.5 mm. long; pappus obscurely biseriate, white, the
inner bristles about 3 mm. long, the outer ones minute, unequal and inconspicuous.
Conyza coronopifolia HBK. Nov. Gen. & Sp. 4: 70. 1820.
Erigeron variifolius Blake, Contr. Gray Herb. 52: 32. 1917.
Usually in damp or wet meadows, pastures, or thickets,
sometimes weeds in cultivated ground, rarely in dry oak forest and
on open, limestone hillsides, 1,400-3,750 m.; Alta Verapaz; Chimal-
tenango; Guatemala; Huehuetenango; Quezaltenango; El Quiche;
Sacatepequez; San Marcos; Solola. Southern Mexico; Costa Rica.
Erect or ascending annuals or perennials, the stems sometimes solitary but more
often several from a single root, mostly 10-75 cm. tall, simple or abundantly
branched, more or less canescent-hirsute; basal leaves narrowing to winged petioles,
the blades usually pinnatifid, with few or numerous, linear or oblong segments, the
cauline leaves sessile, mostly 3-8 cm. long, linear to oblong or oblanceolate, pinnate-
lobate with few, obtuse lobes, sometimes only coarsely crenate, hirsute with
spreading, white hairs; heads usually few, pedicellate, clustered at the ends of the
stems or branches; involucres 4-5 mm. high, usually densely pilose and glandular;
phyllaries often purplish or tipped with purple; pistillate flowers more than 200,
white, the ligules inconspicuous; hermaphrodite flowers about 30, yellow; denuded
receptacles commonly 4-6 mm. in diameter but occasionally in dwarf forms only
about 2 mm.; achenes pale brownish, about 1 mm. long, glabrous; pappus uniseriate,
white, about 3 mm. long.
Conyza filaginoides (A. DC.) Hieron. Bot. Jahrb. 28: 588.
1901. Laenecia filaginoides A. DC. in DC. Prodr. 5: 376. 1836.
Open or brushy slopes or fields, or frequently in pine-oak forest,
sometimes a weed in cornfields, 1,500-3,300 m.; Chimaltenango;
Guatemala; Huehuetenango; Quezaltenango; El Quiche; Sacatepe-
quez; San Marcos; Solola. Mexico; South America (Colombia and
Ecuador).
Erect annuals, 15-50 cm. tall, simple or sparsely branched, the stems lanate or
arachnoid-tomentose, usually very leafy; leaves cauline, sessile, linear to oblanceo-
late-linear, mostly 10-30 mm. long, 1-3 mm. wide, the margins dentate but often
revolute and appearing crenate, more or less arachnoid-lanate on both surfaces or
sometimes glabrate in age; heads mostly sessile or short-petiolate in the leaf axils and
forming a long, narrow, spikelike area of inflorescence; involucres 4-5 mm. high, more
or less lanate; phyllaries linear-lanceolate, acuminate, greenish or often with purple
NASH: FLORA OF GUATEMALA 153
tips; pistillate flowers 40-50 per head, the filiform corollas about half as long as the
styles, the ligules wanting; achenes about 1 mm. long, densely sericeous-pilose with
rather long, white hairs; pappus biseriate, the inner bristles white or yellowish, 2-3
mm. long, the outer bristles less than 1 mm. long.
Conyza microcephala Hemsley, Biol. Centr. Am. Bot. 2: 126.
1882.
Not reported from Guatemala, but to be expected there as it
has been collected several times in nearby Chiapas, Mexico.
Erect, slender annuals, the stems striate, sparsely pilose; leaves sessile, suberect,
linear to linear-lanceolate, 1.5-3.5 cm. long, acute or obtuse, at least the lower ones
with 2-5 shallow lobes or teeth near the apex, the middle and upper ones entire or 2-
3-dentate, more or less pilose along the costae, ciliate or pilose on the margins,
usually glabrous elsewhere; inflorescences terminal, the short, corymbiform panicles
composed of numerous, densely clustered heads on short pedicels 2-5 mm. long; heads
many-flowered, each 3-4 mm. broad; involucres 2-3 mm. high; phyllaries linear-
subulate; receptacles concave; pistillate flowers 75-100, the corollas glabrous; achenes
compressed, oblong, sparsely appressed-hirsute; pappus white, uniseriate.
Conyza schiedeana (Less.) Cronquist, Bull. Torr. Bot. Club
70: 632. 1943. Erigeron schiedeanus Less. Linnaea 5: 145. 1830. C.
subdecurrens A. DC. in DC. Prodr. 5: 379. 1836. E. subspicatus
Benth. in Oerst. Vid. Medd. Kjoebenhavn 1853: 82. 1853, not C.
subspicata phil. 1856. E. subdecurrens Sch. Bip. ex A. Gray, Syn. Fl.
N. Am. 1(2): 220. 1884, not E. subdecurrens A. Gray, 1856. E.
subspicatus var. leiocarpus Blake, Brittonia 2: 337. 1937 (type from
Huehuetenango, Sierra de los Cuchumatanes, Skutch 1214). C.
prolialba Cuatr. Phytologia 9: 3. 1963. C. prolialba var. subspicata
Cuatr. torn, cit.: 5.
Damp thickets or alpine meadows, 2,500-3,500 m.; Huehue-
tenango; Quezaltenango. Texas; Mexico; Costa Rica; South Amer-
ica.
Erect annuals or perennials, 20-60 cm. tall, rather densely white-lanate on all
younger parts, usually simple but occasionally with some very erect or ascending
branches; cauline leaves numerous, often crowded, sessile, mostly 1-3 cm. long, linear-
oblong or sometimes linear-lanceolate, obtuse or acute, the upper ones dilated and
amplexicaul at the base, the margins coarsely dentate, crenate-lobate, or sometimes
essentially entire, commonly revolute, usually more or less lanate, and with shorter,
glandular hairs often hidden by the longer hairs, sometimes glabrate in age, at least
the upper lobes or teeth mucronulate; inflorescence spikelike, the heads subsessile
and often glomerate in leaf axils, the "spikes" sometimes forming pyramidal panicles;
involucres 4-5 mm. high; phyllaries about 3-seriate, unequal, pale green to purple,
lance-linear, acuminate, villous-pilose, more or less glandular and with longer hairs
along the costae, the margins scarious and ciliate, at least near the apex; pistillate
flowers commonly 200-300 per head, the corollas very minutely ligulate; hermaphro-
154 FIELDIANA: BOTANY, VOLUME 24
dite flowers 10-20, the tubular corollas yellow; achenes about 1 mm. long, pubescent
or glabrous; pappus uniseriate, about 3 mm. long.
Specimens from Guatemala and Mexico tend to have glabrous
achenes, although a few plants have achenes that are sparsely
pubescent; specimens from Costa Rica usually have pubescent
achenes, but those of South American plants are essentially
glabrous.
Conyza sophiifolia HBK. Nov. Gen. & Sp. 4: 72, t. 326. 1820.
C. coulteri Gray var. tenuisecta Gray, Syn. Fl. 1(2): 221. 1884.
Eschenbachia tenuisecta Wooten & Standley, Contr. U. S. Nat.
Herb. 16: 186. 1913. Marsea sophiaefolia Badillo, Bol. Soc. Venez.
Cienc. Nat. 10: 256. 1946. Rastrojera (Guatemala, fide Aguilar).
Wet to dry fields and thickets, often an abundant weed of
abandoned fields and waste ground, sometimes on sandbars along
streams, occasionally in open pine or oak-pine forest, 1,200-3,000 m.;
Chimaltenango; Chiquimula; Guatemala; Huehuetenango; Jalapa;
El Progreso; Quezaltenango; Sacatepequez; San Marcos; Santa
Rosa; Solola; Totonicapan. Mexico; El Salvador; South America
(Ecuador, Colombia, Bolivia).
Erect annuals to about 1 m. high, often much branched, the stems striate, more
or less hirsute with fairly long, multiseptate hairs and hispidulous with short, usually
glandular hairs; upper leaves sessile, the lower ones petiolate or sessile, the blades
mostly 2-9 cm. long, mostly bipinnatifid, the segments numerous, linear to linear-
oblong, obtuse, densely hispidulous or puberulent; inflorescences racemiform or
thyrsiform, often becoming paniculate, the branches erect or ascending; heads very
numerous, small, the involucres 2-3 mm. high; phyllaries rather few, greenish with
pale margins, subequal, lance-linear, acuminate, villous-pilosulous; pistillate flowers
75-110 per head; achenes pale, scarcely 1 mm. long; pappus uniseriate, yellowish
white, about 2.5 mm. long.
EGLETES Cassini
Reference: L. H. Shinners, Revision of the genus Egletes
Cassini north of South America, Lloydia 12: 239-250. 1949; Two
additions to the genus Egletes Cassini from northern South
America, torn, cit.: 248-250.
Annuals, usually very viscid, branched, erect; leaves alternate, sessile or
petiolate, the margins dentate, lobate, pinnatifid or bipinnatifid; heads heterogamous,
radiate, in fruit subglobose, pedunculate, solitary, few or numerous, the lax cymes
sometimes appearing corymbose; involucres campanulate; phyllaries 2-3-seriate,
subequal, acute or acuminate, the outer ones herbaceous, the inner membranaceous;
receptacles conic or in age ovoid, naked; ray flowers pistillate, 1 -several-seriate, the
ligules white; disc flowers hermaphrodite, fertile, the corollas tubular, yellow, 3-5-
cleft at the apex; anthers obtuse and entire at the base; styles of the disc flowers
NASH: FLORA OF GUATEMALA 155
short, flattened, their appendages very short, obtuse; achenes oblong, somewhat
compressed, ecostate, constricted at the base, producing at the apex a cartilaginous
crown, ring, or cupule.
Shinners recognized nine species, with the following two in
Guatemala.
Ray flowers in 1-2 series; ligules 0.4-1.6 mm. wide E. viscosa.
Ray flowers in 3-4 series; ligules less than 0.25 mm. wide.
E. liebmannii var. yucatana.
Egletes liebmannii Sch. Bip. Leopoldina 23: 88. 1887. E.
pringlei Greenman, Field Mus. Bot. 2: 265. 1907.
Erect, usually widely branching herbs, mostly 20-40 cm. tall, the stems and
branches glandular-pubescent; leaves on narrowly winged petioles, these more or less
clasping at the base, the blades oblong-oval to oblong, mostly 3-7 cm. long, 2-4.5 cm.
wide, lyrate-lobate or lobate-dentate, commonly sparsely glandular-pubescent, the
uppermost leaves smaller and sometimes cuneate to a sessile base; heads few or
numerous, on peduncles 0.3-1.5 cm. long; involucres 3.5-4.5 mm. high; phyllaries 2-3-
seriate, subequal, lanceolate, glandular- pubescent; ray flowers in 3-4 series, the ligules
white, about 1 mm. long, very narrowly linear, less than 0.25 mm. wide; disc corollas
tubular, yellow; achenes about 1 mm. long, glandular-pubescent, the crown or ring
forming a shallow cup.
The typical variety has not been found in Guatemala, and
according to Shinners, is restricted to eastern Mexico; only the
following variety is known in Guatemala.
Egletes liebmannii var. yucatana Shinners, Lloydia 12: 246.
1949. Figure 34.
Swampy thickets, clearings, and waste ground, 50-75 m.; Peten.
Mexico; British Honduras; Honduras.
Differs from the typical variety in its broadly winged petioles,
pinnatifid leaves, and more conspicuous ray flowers with ligules
about 2 mm. long.
Egletes viscosa (L.) Less. Syn. Comp. 252. 1832. Cotula
viscosa L. Sp. PL 2: 892. 1753. E. viscosa f. bipinnatifida Shinners,
Lloydia 12: 244. 1949.
Grassy fields about the margins of lakes or in waste ground,
near sea level to 800 m.; Escuintla; Jutiapa. Southern Texas
through most of Mexico; El Salvador; Nicaragua; Costa Rica;
Cuba.
Erect or ascending plants, the stems usually much branched, 12-60 cm. tall,
usually viscid-villous throughout; leaves very variable in form, the lower ones long-
petiolate and clasping at the petiolar base, the blades often pinna te-lo bate, the
middle and upper leaves sessile or petiolate, oblong to broadly obovate, obtuse,
156 FIELDIANA: BOTANY, VOLUME 24
mostly 4-11 cm. long, 2-6 cm. wide, shallowly or deeply lobate or pinnatifid or
bipinnatifid, the uppermost leaves smaller, narrower, and commonly less deeply
divided, amplexicaul; heads usually few, sometimes numerous, on short or rarely
elongated peduncles, the lax cymes sometimes appearing corymbose; involucres
mostly 3-4.5 mm. high; phyllaries biseriate, subequal, lanceolate to ovate-lanceolate,
acute, more or less hispid and and glandular- pubescent; ray flowers in 1-2-series, the
ligules white, only slightly exceeding the phyllaries, 0.4-1.6 mm. wide; disc corollas
tubular, yellow; achenes 1.3-1.5 mm. long, slightly compressed, more or less
glandular-puberulent, the crown or ring narrow and uneven.
ERIGERON Linnaeus
References: Arthur Cronquist, The separation of Erigeron from
Conyza, Bull. Torr. Bot. Club 70: 629-632. 1943; Otto T. Solbrig, the
South American species of Erigeron, Contr. Gray Herb. 191: 3-79.
1962.
Usually perennial herbs, sometimes biennial, rarely shrubby, commonly
pubescent; leaves alternate or rosulate, the margins dentate, crenate, entire, or
sometimes incised or dissected; heads heterogamous, radiate, solitary and peduncu-
late (in ours); involucres hemispheric or campanulate; phyllaries usually biseriate,
narrow, subequal; ray flowers numerous, usually arranged in one series, the ligules
larger than the tubular flowers and equalling or exceeding the pappus, white, pink,
violet, or purple; disc flowers commonly yellow, hermaphrodite, tubular, mostly 5-
dentate, usually fertile; receptacles flat or nearly so, naked or rarely the foveae with
very small pales or fimbrillae; anthers obtuse and entire at the base; style branches
of the disc flowers very short with triangular-tipped stigmas, acute or obtuse; achenes
very small, more or less compressed, the margins usually nervelike, the faces without
nerves, 1-2-nerved, or rarely as much as 14-nerved; pappus 1-2-seriate, the inner
bristles slender and soft, the outer ones irregular, usually much shorter.
Perhaps 200 species, widely distributed in both hemispheres,
most numerous in temperate and cold montane regions, with four in
Guatemala.
Stems scapiform, the principal leaves mostly confined to the lower half of the stem or
mostly basal and forming a rosette.
Lower leaves usually numerous, with or without a basal rosette; heads about 15
mm. broad, involucres about 7 mm. high; ray flowers conspicuous, the ligules
5-7 mm. long E. scaposus.
Lower leaves few, basal, forming a rosette; heads only 5-7 mm. broad, involucres 3-
4 mm. high; ray flowers inconspicuous, the ligules 1-2 mm. long.
E. jamaicensis .
Stems leafy throughout.
Plants suberect or decumbent; indument of stems and peduncles composed of
spreading or retro rse hairs; leaves oblong- lanceolate, elliptic-oblong, or linear -
oblanceolate, the margins entire or remotely crenate E. aquarius.
Plants erect or ascending and usually forming dense clumps; indument of stems
and peduncles composed of ascending or appressed hairs; leaves varying in
form from narrowly linear and entire to obovate and entire or crenate; or
deeply 3-5-lobate, to elliptic and entire, dentate, or shallowly lobate.
E. karvinskianus.
NASH: FLORA OF GUATEMALA 157
Erigeron aquarius Standl. & Steyerm. Field Mus. Bot. 22:
301. 1940.
Known only from the original locality, damp forest, Volcan de
Agua, above Santa Maria de Jesus, Sacatepequez, about 2,500 m.,
the type being Standley 65053.
Suberect or decumbent herbs, 30-50 cm. high, probably perennial, branched, leafy
throughout, the stems pilose with lax, spreading or retrorse hairs; cauline leaves
sessile, narrowly oblong-lanceolate, elliptic-oblong or linear-oblanceolate, 2.5-6.5 cm.
long, 0.6-1 cm. wide, acute or subobtuse and apiculate, gradually narrowed to the
acute or broadly cuneate base, the margins entire or the larger leaves remotely
crenate, rather densely pilose on both surfaces with spreading hairs; heads solitary at
the ends of the numerous branches, on slender, elongated peduncles, these pilose with
spreading hairs; involucres broadly campanulate, about 8 mm. high and 12 mm.
broad; phyllaries 2-3-seriate, unequal, linear, attenuate, pale-marginate, sparsely
appressed-pilose; ray flowers very numerous, white, narrowly linear, the ligules about
3 mm. long, erect or spreading; receptacle 3 mm. broad; disc corollas 4.5 mm. long,
glabrous, yellow; achenes about 1 mm. long, sparsely appressed-pilose, the pappus
sordid, 3-4 mm. long.
Erigeron jamaicensis L. Syst. ed. 10. 1213. 1759. Aster
jamaicensis Less. Linnaea 5: 144. 1830.
Damp or wet fields or open banks, near sea level to 900 m.;
Peten; Santa Rosa. Mexico; Honduras; Costa Rica; Panama;
Greater Antilles.
Plants perennial from short or elongated rootstocks, the stems very slender,
scapiform, simple or sparsely branched, erect or ascending, mostly 10-30 cm. long,
pubescent or glabrate; basal leaves usually few, forming a small rosette, spathulate or
obovate, mostly 3-10 cm. long, 0.5-2 cm. broad, subentire or coarsely dentate or
crenate, obtuse, cuneately narrowed to the sessile base or forming a short, winged
petiole, the cauline leaves sessile, very small, narrow, often bractlike, linear or oblong,
mostly entire; heads 1-few at the ends of the stems, small, mostly 5-7 mm. broad;
involucres 3-4 mm. high; phyllaries linear, acuminate, narrowly scarious-marginate,
sparsely hispidulous or glabrous; ray flowers white, the ligules short and
inconspicuous, 1-2 mm. long, spreading; achenes 1.5 mm. long, pubescent; pappus
sordid, about 3 mm. long.
Erigeron karvinskianus DC. Prodr. 5: 285. 1836. E. mucr-
onatus DC. I.e. E. pacayensis Greenm. Field Mus. Bot. 2: 266. 1907
(type from Crater of Volcan de Pacaya, Guatemala, Kellerman
6111). E. deamii Robins. Proc. Am. Acad. 45: 410. 1910 (type from
Fiscal, Guatemala, Deam 6159). E. tripartitus Blake, Brittonia 2:
337. 1937 (type from southern slope of Volcan de Atitlan,
Suchitepequez, Skutch 2124). Cambray menudo (fide Morales),
mansanilla de monte, margarita, sombrita (Guatemala); margarita
silvestre (Huehuetenango). Figure 35.
158 FIELDIANA: BOTANY, VOLUME 24
Most common on steep, open banks or hillsides, often on cliffs
or in rock crevices, abundant in damp or wet thickets or open forest,
often in pine-oak forest, 900-3,700 m., most abundant at 1,300-2,800
m.; Alta Verapaz; Chimaltenango; Chiquimula; Guatemala;
Huehuetenango; Jalapa; Quezaltenango; El Quiche; Retalhuleu;
Sacatepequez; San Marcos; Santa Rosa; Solola; Suchitepequez;
Totonicapan. Mexico; Honduras; El Salvador; Nicaragua; Costa
Rica; South America.
Perennial herbs with numerous, slender, branching stems, usually forming dense
clumps, erect, ascending, or the clump sometimes pendent from banks, mostly 10-30
cm. tall, pilose or hispidulous with short, ascending or appressed hairs or glabrate,
leafy throughout; leaves small, mostly 1-4 cm. long, more or less strigose, especially
on the veins, rarely almost glabrous, varying in form from narrowly linear (1-3 mm.
wide) and entire, or the lower ones with 3 linear lobes to obovate (3-10 mm. wide)
and entire, crenate, or deeply 3-5-lobate, to elliptic (3-10 mm. wide) and entire or
dentate or shallowly lobate; heads usually numerous, solitary on 1-3 slender
peduncles at the ends of stems and branches, these commonly 2-8 cm. long, the heads
hemispheric, variable in size, the denuded discs mostly 3-8 mm. broad; involucres 3-6
mm. high; phyllaries subequal, linear, acuminate, often purplish, strigose or glabrate;
rays 75-100 or more, white, pink or rarely purplish, the ligules minutely 2-3-dentate,
varying from 3-4 mm. long to 9-10 mm. long, disc corollas numerous, yellow; achenes
pale, sparsely pilosulous or almost glabrous, about 1 mm. long; pappus white or
yellowish white, 2-3 mm. long.
A pantropical, weedy complex; one of the most common small
plants of the mountains of Guatemala. The plants are highly
variable in foliage and several not very extreme forms have been
described from Guatemala. These descriptions probably would have
been delayed if the authors had seen such a quantity of material as
has since been collected in Guatemala. E. pacayensis is a dwarf
form, not otherwise noteworthy, of the exposed, windswept slopes of
the summit of Volcan de Pacaya. E. deamii is a stunted, reduced
form of dry areas with narrow leaves and rather small heads,
although no smaller than may be frequently found in other areas.
The most extreme of the local forms is E. tripartitus, in which some
or most of the leaves are deeply 3-5-lobate, but the leaves vary in
this respect upon a single plant (as in many other species of
Erigeron). A count of 110 specimens from all departments revealed
that 42 had mostly linear, entire leaves 1-3 mm. wide, but 20 of
these specimens also had a few deeply lobate leaves; 39 specimens
had leaves 3-10 mm. wide, mostly elliptic, but about half of these
also displayed a few obovate, lobate leaves and had linear upper
leaves; 29 specimens (900-2,800 m.) had obovate, mostly lobate
leaves 4-10 mm. wide, but in many of these the upper leaves were
often more or less elliptic and the uppermost ones linear. Plants
NASH: FLORA OF GUATEMALA 159
with the broadest leaves, whether elliptic or obovate and deeply
lobate, tend to have the largest heads.
Erigeron scaposus DC. Prodr. 5: 287. 1836. E. scaposus var.
latifolius DC. I.e. E. affine DC. torn, cit: 289, not E. affinis Badillo,
1946. Margarita de rosa (Guatemala, fide Morales).
Wet to dry meadows or open forest, sometimes a weed in
cultivated ground, occasionally in open pine forest (perhaps the
original habitat?), 800-2,500 m.; Alta Verapaz; Baja Verapaz;
Chimaltenango; Chiquimula; Escuintla; Guatemala; Huehue-
tenango; San Marcos; Santa Rosa; Solola; Totonicapan; Zacapa.
Mexico; Honduras; Nicaragua.
Perennials from slender, elongating rootstocks, the stems simple, scapiform,
rarely with a few branches, slender, erect or somewhat decumbent, mostly 10-30 cm.
long, striate, usually densely pubescent with short, mostly spreading or reflexed hairs,
sometimes glabrate, leafy below the middle and especially so near the base, with or
without a basal rosette, occasionally with some well developed leaves above the
middle but most often the upper ones much reduced or lacking or inconspicuous;
leaves usually numerous, sessile and mostly amplexicaul or the lowermost ones
sometimes narrowing into a winged petiole, the blades oblong to obovate-oblong or
spathulate-oblong, the principal ones mostly 3-6 cm. long, obtuse or rounded at the
apex, often mucronulate, the margins crenate or coarsely serrate, pilose on both
surfaces, most often with short hairs but sometimes the hairs rather long,
subappressed or somewhat spreading; heads solitary (or very rarely 2) at the ends of
the stems, on long, slender, naked peduncles, these often much longer than the leafy
portion of the stem, usually densely pubescent; heads about 1.5 cm. broad,
hemispheric; involucres about 7 mm. high; phyllaries numerous, subequal, linear,
acuminate, more or less pilosulous, pale-marginate; ray flowers numerous, white or
rarely pinkish or in age purplish, the ligules very narrow but rather conspicuous, 5-7
mm. long, spreading; disc flowers yellow; achenes scarcely 1 mm. long, thinly pilose;
pappus sordid, about 3.5 mm. long.
GYMNOSPERMA Lessing
Reference: Otto T. Solbrig, The status of the genera Amphi-
pappus, Amphiachyris, Greenella, Gutierrezia, Gymnos-
perma and Xanthocephalum (Compositae), Rhodora 62: 43-54. 1960.
Erect, usually densely branched perennials, herbaceous or suffrutescent, very
glutinous; leaves alternate, linear to linear- lanceolate, the margins entire, both
surfaces punctate; inflorescences corymbose, the heads heterogamous, radiate,
disposed in small cymes; involucres narrowly ovoid-campanulate; phyllaries 4-5-
seriate, with hyaline margins and dark green tips, obtuse or subacute, the outer ones
smaller; receptacles foveolate; ray flowers few (less than 10), uniseriate, pistillate,
fertile, the ligules short and inconspicuous, not surpassing the tubular corollas; disc
flowers hermaphrodite, fertile or some sterile, tubular, the limb narrowly camp-
anulate, 5-cleft; anthers obtuse and entire at the base; style branches of the disc
160 FIELDIANA: BOTANY, VOLUME 24
flowers elliptic; achenes oblong, somewhat compressed, pubescent, 4-5-costate;
pappus reduced to an inconspicuous crown of minute scales.
The genus consists of a single species.
Gymnosperma glutinosa (Spreng.) Less. Syn. Comp. 194. 1832.
Selloa glutinosa Spreng. Nov. Prov. Hal. 36. 1819. G. corymbosum
DC. Prodr. 5: 312. 1836. G. multiflorum DC., I.e. G. scoparium DC.
I.e. S. multiflora O. Ktze. Rev. Gen. 1: 362. 1891. S. scoparia O.
Ktze. I.e. Xanthocephalum glutinosum Shinners, Field & Lab. 18:
25-32. 1950. Figure 36.
Dry, rocky, brushy, mountain slopes, often on serpentine,
sometimes in gravelly river beds, oak or oak-pine forest, 1,200-2,800
m.; Huehuetenango. Texas to Arizona; Mexico.
Stiffly erect perennials to about 1 m. tall, glabrous or nearly so, very glutinous,
the stems striate, often much branched, the branches erect or nearly so; leaves sessile,
often with fascicles of leaves in their axils, narrowly or broadly linear to linear-
lanceolate, acute or or acuminate, mostly 2-7 cm. long, 0.2-0.7 cm. wide, triplinerved,
conspicuously punctate; inflorescences corymbose, usually dense, the sessile or short-
pedicellate heads usually 2-7 on the bracteate peduncles; involucres narrowly ovoid-
campanulate, 4-5 mm. high; phyllaries 4-5-seriate, pale yellowish with narrow,
scarious margins, green-tipped; ray flowers commonly 6, yellow, very short; disc
flowers about 6, yellow; achenes pubescent, about 1.5 mm. long.
HAPLOPAPPUS Cassini ex Endlicher
Reference: H. M. Hall, The genus Haplopappus, Carn. Inst.
Wash. Publ. 389. 1928.
Herbs or shrubs from taproots and/or rhizomes, usually more or less resinous or
glandular; leaves all or mostly alternate, sometimes all basal, the margins entire,
dentate, or lobate; heads solitary on long peduncles (in ours) or variously disposed,
heterogamous, usually radiate; involucres hemispheric to cylindrical; phyllaries
several-seriate, numerous, subequal or unequal and imbricate, herbaceous to
chartaceous, often with greenish tips; receptacles flat or slightly convex, naked,
foveolate; ray flowers uniseriate, few to moderately numerous or rarely wanting,
pistillate or neutral, fertile or sterile, the ligules yellow (in ours) or rarely white; disc
flowers few-many, hermaphrodite, yellow, white, or purplish, fertile or rarely sterile,
the corollas funnelform to subcylindric, the limb 5-dentate or 5-lobate; anthers
essentially entire at the base; style branches flattened, the appendages ovate and
acute or more often acuminate; achenes commonly 4-5-costate, striate or smooth,
pubescent or glabrous; pappus uniseriate, the bristles numerous, unequal, often
sordid.
About 150 species, all American, numerous in the western
United States, Mexico, and in the cooler regions of South America,
with only one reaching Central America.
NASH: FLORA OF GUATEMALA 161
Haplopappus stoloniferus DC. Prodr. 5: 349. 1836. Aster
stolonifer Ktze. Rev. Gen. 1: 318. 1891. Aplopappus stoloniferus var.
glabratus Coulter, Bot. Gaz. 16: 98. 1891 (type from crater of
Volcan de Agua, Sacatepequez, J. D. Smith 2328). Osbertia
stolonifera Greene, Erythea 3: 14. 1895. O. heleniastrum var.
glabrata Greene, I.e. O. heleniastrum var. scabrella Greene, I.e.
(type from Sacatepequez, J. D. Smith 3697). Aplopappus stoloni-
ferus var. genuinus Blake, Contr. Gray Herb. 52: 25. 1917. A.
stoloniferus var. heleniastrum Blake, I.e. Figure 37.
In alpine situations on the highest mountains, usually growing
in open, sandy soil or among rocks, but also in open pine forest and
in damp meadows, 2,500-4,600 m.; Chimaltenango (Volcan de
Acatenango); Huehuetenango; Quezaltenango (Volcan de Santa
Maria); San Marcos (volcanoes of Tajumulco and Tacana);
Sacatepequez (Volcan de Agua); Solola (Volcan de Atitlan);
Totonicapan. Mountains of southern Mexico.
Perennials from thick rootstocks, usually with numerous basal leaves, sometimes
producing elongated, leafy stolons; stems 1-several, erect, 4-30 (-50) cm. tall,
bracteate or bearing several-numerous, reduced leaves, glandular and variously
pubescent; basal leaves on short or long petioles, the blades membranaceous to rather
thick, narrowly or broadly elliptic to broadly obovate, oblanceolate, or spathulate,
mostly 2-10 cm. long, obtuse, often mucronate, the margins entire or remotely
dentate or denticulate, short-pilose to densely hirsute or even lanate, sometimes
puberulent or glabrate, minutely glandular, the cauline leaves few or numerous, often
much reduced, narrow, bractlike; heads solitary; involucres hemispheric, 10-15 mm.
high; phyllaries numerous, subequal, linear, long-acuminate, usually densely white-
villous and glandular; ray flowers 50-120, yellow, mostly 12-15 mm. long, the ligules
7-9 mm. long, disc flowers very numerous, usually deep yellow or orange-yellow;
achenes subcylindrical, about 2 mm. long, pubescent to densely villous; pappus
uniseriate, 4-5 mm. long, sordid or brownish.
Alpine plants, more characteristic of exposed, rocky places than
of alpine meadows. Guatemalan material, like that from Mexico, is
fairly ample and highly variable, chiefly in the quantity of stem
foliage and in indument. The extreme forms are so connected by
intergrading forms that it scarcely seems desirable to recognize
them by name.
HETEROTHECA Cassini
Perennial or annual herbs, the pubescence sericeous or hispid-pubescent, the
stems branching above; leaves alternate, sessile, commonly narrow, net-veined, the
margins mostly entire, the basal leaves often conspicuous; heads heterogamous,
radiate or rarely discoid, usually solitary at the ends of the branches but forming
laxly corymbose inflorescences; involucres campanulate or hemispheric; phyllaries
imbricate in several series, narrow, scarious to subherbaceous; receptacles flat or
162 FIELDIANA: BOTANY, VOLUME 24
slightly convex, naked, foveolate; ray flowers yellow, pistillate and fertile; disc
flowers yellow, hermaphrodite, fertile; anthers essentially entire at the base; style
branches somewhat flattened, their appendages linear or subulate, puberulent
exteriorly; achenes more or less compressed, pubescent, oblong-linear or obovate;
pappus of disc flowers biseriate, the inner series of numerous, rough, capillary bristles,
the outer of small or minute scales or bristles; pappus absent in ray flowers or
vestigial.
About 20 species, widely dispersed in the United States and
Mexico, extending southward into Guatemala. Only the following
one is known from Central America.
Heterotheca graminifolia (Michx.) Shinners, Field & Lab. 19:
71. 1951. Inula graminifolia Michx. Fl. Bor. Am. 2: 122. 1803.
Chrysopsis graminifolia Ell. Bot. S. Car. & Ga. 2: 334. 1824.
Diplopappus graminifolius Less. Linnaea 5: 310. 1830. Aplopappus
gramineus Benth. PI. Hartweg. 65. 1840. Pityopsis graminifolia
Nutt. Trans. Am. Phil. Soc. 7: 317. 1841. Figure 38.
In open, grassy, pine forest, 1,000-2,500 m.; Chiquimula;
Huehuetenango; Zacapa. Southeastern United States; Mexico;
British Honduras; Honduras; Bahamas.
Perennials from thick rootstocks and sometimes stoloniferous, the stems stiffly
erect, slender, commonly 30-50 cm. tall, usually densely silvery-sericeous almost
throughout, branched above; leaves linear or lance-linear, often appearing grasslike,
3-5-nerved, the basal ones mostly 10-20 cm. long, 0.4-1 (-2) cm. wide, the middle ones
reduced, usually amplexicaul, the upper ones bractlike; inflorescences branching,
laxly corymbiform, the heads several or numerous, solitary at the ends of the
bracteate peduncles; involucres campanula te, mostly 7-12 mm. high; phyllaries linear,
very unequal, densely or laxly sericeous, acute or acuminate, the outer ones often
reflexing in age; rays conspicuous but short, bright yellow; achenes linear-fusiform,
about 4 mm. long, sparsely sericeous; pappus about 7 mm. long, the inner bristles
sordid, minutely barbellate.
LAGENOPHORA Cassini
Small, rhizomatous, perennial herbs forming basal rosettes, the stems simple, 1-
several; basal leaves usually petiolate, the cauline leaves alternate, usually few,
sessile or short-petiolate, reduced upward, the margins entire, dentate, serrate, or
crenate; heads heterogamous, usually solitary on short or elongated peduncles;
involucres campanulate; phyllaries 2-3-seriate, with scarious margins; receptacles
naked, convex or somewhat concave; ray flowers uniseriate, pistillate, white or
purplish, often scarcely exceeding the disc flowers, the ligules often reflexed or
revolute, the apex 2-3-denticulate; disc flowers hermaphrodite, tubular, yellow, 5-
cleft, their style branches somewhat flattened, with triangular or ovate tips; stamens
with minute apical appendages, the anthers obtuse at the base; achenes usually more
or less compressed, the margins thickened, the apex with a short, viscid beak, the base
more or less contracted; pappus none.
NASH: FLORA OF GUATEMALA 163
About 20 species, distributed from Malaysia and the southern
coast of Asia to Australia, Tasmania, New Zealand, Hawaii, South
America, Panama, and one in Guatemala.
Lagenophora cuchumatanica Beaman & De Jong, Rhodora
67: 36-41. 1965. Figure 39.
In pine forest, 3,200-3,365 m., Huehuetenango (Beaman 3756,
type); Totonicapan.
Decumbent or rarely ascending perennials with 1-4 simple stems, mostly 5-20 cm.
long, greenish or reddish, striate, flattened or subterete, pilose with spreading to
appressed hairs, leafy throughout or with naked peduncles sometimes elongating to 6
or 7 cm., a little enlarged and densely pubescent just below the heads; basal leaves
few, on pilose petioles 0.7-3.3 cm. long, the blades ovate-elliptic to orbicular, mostly 1-
2 cm. long, the margins crenate-dentate, more or less pubescent with appressed hairs
and glandular-punctate on both surfaces, or the upper surfaces sometimes glabrate or
nearly so, the lower cauline leaves similar, middle and upper leaves sessile,
oblanceolate to spathulate and serrate, sometimes punctate only near the apex;
heads terminal, usually solitary, often subtended by a single bract; involucres
campanula te, 4-5 mm. high; phyllaries about 3-seriate, lance-oblong to narrowly
lanceolate, glabrate and shining or sparsely pilose, the margins narrowly scarious and
more or less ciliate; receptacles somewhat concave, naked; ray flowers uniseriate, 10-
20, pistillate, 2-3 mm. long, the ligules greenish white, becoming purplish in age,
reflexed, the apex 2-3-denticulate; disc flowers 8-14, hermaphrodite, greenish yellow,
mostly 2-2.5 mm. long, campanulate, 5-lobate, the style branches pubescent outside,
the tips ovate; achenes 3-4 mm. long, brown, glabrous, shining obovate, compressed,
the margins thick-nerved, the short beak viscid.
SOLIDAGO Linneaus
Erect perennial herbs, the stems slender, glabrous or pubescent; leaves alternate,
the cauline ones commonly sessile or nearly so, the basal ones often petiolate, the
margins entire, serrate, or dentate; inflorescences spicate, racemose, or paniculate,
rarely corymbose; heads radiate (in ours), few-many-flowered; involucres camp-
anulate or hemispheric; phyllaries several-seriate, imbricate; receptacles flat or
slightly convex, naked; ray flowers pistillate and fertile, the ligules yellow (in ours);
disc flowers hermaphrodite, fertile, yellow; anthers essentially entire at the base;
style branches flattened, the appendages mostly lanceolate; achenes glabrous or
pubescent, costate; pappus uniseriate, white, the bristles capillary, equal.
Perhaps 100 species, chiefly of North America, a few extending
into South America and a few in Eurasia. A very complex genus
with many species hybridizing freely and difficult to define.
Although none have been reported from Guatemala, the following
species is treated here, as several collections have been made in
Chiapas. Mexico.
Solidago stricta Ait. Hort. Kew. 3: 216. 1789; McVaugh,
Contr. Univ. Mich. Herb. 9: 365-367. 1972. Figure 40.
164 FIELDIANA: BOTANY, VOLUME 24
Damp slopes and bogs, Chiapas, Mexico, 1,000-2,100 m., and
perhaps lower; to be expected in Guatemala. Eastern United States
to Mexico.
Erect perennials with slender stems mostly 0.5-1.5 m. tall, essentially glabrous
with the exception of some inconspicuous pubescence of the inflorescence; basal
leaves more or less spathulate, obtuse, the margins obscurely callose-dentate or
almost entire, the cauline leaves numerous, oblanceolate to lanceolate or elliptic,
rapidly reduced upward, the uppermost bractlike and appressed; inflorescence
narrowly cylindrical or spiciform or sometimes branching and becoming somewhat
paniculate; peduncles usually minutely hispidulous; heads numerous; involucres
about 5 mm. high; phyllaries 3-4-seriate, linear or linear-lanceolate; ray flowers
commonly 5-9 (-11), the ligules yellow, 2.5-3 mm. long; disc flowers 12-14; achenes
essentially glabrous to rather densely pubescent, 1.5-2.2 mm. long; pappus uniseriate,
white, about 3 mm. long.
TRIBE IV. INULEAE
By DOROTHY L. NASH
Plants herbaceous or suffrutescent or rarely shrubs (Pluchea); leaves alternate,
often densely tomentose; inflorescences commonly cymose, sometimes corymbose or
paniculate, rarely spicate (Gnaphalium); heads heterogamous, discoid; involucres
ovoid to hemispheric; phyllaries few-multiseriate, often scarious; receptacles naked or
rarely with a few short pales (Achyrocline), usually flat, sometimes convex or
subconic; corollas pink, purplish, white, rarely yellow (Gnaphalium); anthers
sagittate at the base, the auricles caudate, the apices not appendaged; style branches
truncate or more or less subulate; achenes various; pappus wanting or composed of
bristles, these usually uniseriate.
Represented in Guatemala by six genera (23 species). The
largest and most difficult genus is the cosmopolitan Gnaphalium,
with 16 species in Guatemala, although none are conspicuously
abundant in the flora. Most are plants of the high country, several
found in alpine meadows, 3,000-4,500 m.
Leaves, at least the lower ones, more or less lyrate-pinnatifid.
Achenes about 5.5 mm. long, stipitate-glandular; pappus wanting Adenocaulon.
Achenes about 1 mm. long, hispid, not stipitate-glandular; pappus present.
Pseudoconyza
Leaves not lyrate-pinnatifid.
Heads 4-5-flowered, the hermaphrodite flowers solitary or two Achyrocline.
Heads many-flowered, the hermaphrodite flowers several or numerous.
Pappus present.
Stems and at least the lower leaf surfaces commonly more or less lanate;
phyllaries scarious, at least at the apex Gnaphalium.
Stems and lower leaf surfaces not lanate; phyllaries not scarious Pluchea.
Pappus wanting Epaltes.
NASH: FLORA OF GUATEMALA 165
ACHYROCLINE Lessing
Herbaceous or suffrutescent perennials, pale-tomentose or rarely glabrate; leaves
alternate, petiolate or sessile; inflorescences cymose, often becoming paniculate;
heads small, heterogamous, discoid, with 3-6 outer pistillate flowers and 1-2
hermaphrodite disc flowers, these all fertile; involucres fuscous to yellowish or white,
oblong; phyllaries multiseriate, imbricate, scarious, obtuse or subacute, the outer ones
gradually shorter, the outermost barely lanate dorsally; receptacles small, naked or
with a few short pales; pistillate corollas filiform, minutely dentate or shallowly 2-4-
cleft; hermaphrodite corollas regular, tubular, dilated at the base, the limb little
ampliate, shallowly 5-lobate; anthers sagittate at the base, the auricles usually short-
caudate; style branches slender, truncate; achenes oblong, subcompressed; pappus
bristles scabrous, caducous.
About 15 species, in tropical America and Africa, with only one
known in Central America.
Achyrocline deflexa Robins. & Greenm. Amer. Journ. Sci. 50:
153. 1895. A. yunckeri Blake in Yuncker, Field Mus. Bot. 17: 399.pl.
16. 1938 (type collected near Siguatepeque, Honduras). Figure 41.
Usually in open pine or oak forest, sometimes in mixed forest or
on brushy cliffs, 1,500-3,000 m.; Chimaltenango; Guatemala;
Huehuetenango; Jalapa; El Progreso; Quezaltenango; El Quiche;
Zacapa. Southern Mexico; Honduras; Nicaragua.
Erect perennials to about 1 m. high, sometimes woody below, the stems simple or
branched, densely white-lanate; leaves short-petiolate, the blades thick, lanceolate or
oblong-lanceolate, mostly 3.5-7.5 cm. long, acute at each end, mucronate, the margins
entire, thinly arachnoid above, densely white-lanate beneath; heads about 5-f lowered,
glomerate, disposed in large, broad, rounded panicles; phyllaries oblong, subacute,
white; flowers all fertile; achenes ovate, glabrous.
ADENOCAULON Hooker
Erect or ascending, annual or perennial herbs, bearing numerous stipitate, viscid
glands, especially in the inflorescence; leaves alternate, the lower ones long-petiolate,
the blades white-tomentose beneath; heads small, heterogamous, discoid, disposed in
large, irregular panicles; outer flowers 4-7, pistillate, fertile; disc flowers of the same
number, sterile; involucres broadly campanulate; phyllaries 5-8, subequal, herba-
ceous; receptacle flat, naked; corollas regular or bilabiate, tubular, the tube in the
pistillate flowers very short, in the hermaphrodite flowers slender; anthers short-
caudate at the base; achenes obovoid, somewhat compressed, obscurely costate,
stipitate-glandular; pappus wanting.
About six species, in western North America, Chile, and eastern
Asia. Only one is known in North America south of the United
States.
166 FIELDIANA: BOTANY, VOLUME 24
Adenocaulon lyratum Blake, Journ. Wash. Acad. Sci. 24: 435.
/. /. 1934. Figure 42.
Usually in damp oak forest, sometimes in thickets, 1,900-2,700
m.; Chimaltenango (type from Chichavac, Skutch 622}; Huehue-
tenango. Mexico (Chiapas).
Plants erect or nearly so, from short rootstocks, the stems rather stout, mostly
50-75 cm. tall, simple or sparsely branched, thinly arachnoid-tomentose, narrowly
winged by the decurrent leaf bases; larger leaves crowded near the base of the stem,
obovate in outline, lyrate-pinnatifid, mostly 15-25 cm. long, green and glabrate on the
upper surface, densely and closely arachnoid-tomentose beneath, the terminal lobe
broadly deltoid and subcordate, acute or obtuse, minutely repand-dentate, the lateral
lobes short, acute or very obtuse, the broadly winged base of the leaf petioliform;
upper leaves much smaller and less lobate; heads solitary or two in the axils of the
small bracts, the peduncles mostly 2-4 cm. long; heads 10- 16-f lowered, at anthesis 2-3
mm. broad; phyllaries 6-8, subequal, ovate, acute, 1.5-2 mm. long, thinly arachnoid;
pistillate flowers 5-8, the corollas white, about 1 mm. long; hermaphrodite flowers 5-
8, the corollas about 2.2 mm. long; achenes cuneate-obovoid, commonly 3-nerved on
each side, stipitate-glandular, about 5.5 mm. long.
A strangely isolated species, its nearest congeners being found
in northwestern United States and Chile. The plants seem to be of
very local occurrence in Guatemala but are often plentiful where
found.
EPALTES Cassini
Erect or decumbent, branching herbs or suffrutescent plants, the stems winged,
puberulent or glabrate; leaves alternate, sessile, clasping and long-decurrent on the
stem, the blades oblong to obovate or spathulate, the margins entire or serrate-
dentate; inflorescences cymose-corymbose; heads small, heterogamous, discoid;
involucres campanulate or hemispheric; phyllaries multiseriate, imbricate, dry;
receptacle flat or convex and somewhat concave in the middle, naked; outer pistillate
flowers multiseriate, fertile; inner flowers usually few, hermaphrodite, sterile; corollas
pink or purplish (in ours), the pistillate ones filiform, the limb bifid or trifid, the
others tubular, 3-5-cleft; anthers sagittate at the base, the auricles acuminate-
caudate; styles of the hermaphrodite flowers subulate, simple or bifid; achenes 5-10-
costate; pappus wanting.
Perhaps 10 species, all tropical, in both hemispheres. Although
none are known to occur in Guatemala, one species is treated here,
as it has been collected in Chiapas, Mexico and may be expected in
Guatemala.
Epaltes mexicana Less, in Schlecht., Linnaea 5: 147. 1830.
Figure 43.
Not reported from Guatemala, but to be expected there. Mexico
(Chiapas, Oaxaca, Tabasco, Veracruz), near sea level to 900 m..
NASH: FLORA OF GUATEMALA 167
Suffrutescent plants, the stems striate, puberulent to pilosulous or glabrate,
winged; leaves sessile, clasping, long-decurrent on the stems, the blades linear-oblong
to spathulate or ovate-oblong, mostly 3-12 cm. long, 0.5-4 cm. wide, acute, more or
less pilose on both surfaces or glabrate, the margins irregularly serrulate;
inflorescences cymose-corymbose, the corymbs commonly 3-7 cm. broad, sometimes
larger; heads usually few, 5-6 mm. high and broad; involucres broadly campanulate
to nearly hemispheric, pilose to arachnoid-tomentose; phyllaries 2-3-seriate, ovate to
lanceolate, acute; pistillate corollas filiform, trifid, the others tubular, 4-5-cleft; styles
of the sterile hermaphrodite flowers bifid; achenes 5-costate, glabrous, less than 1
mm. long.
GNAPHALIUM Linneaus
References: A. L. Cabrera, Observaciones sobre las Inuleae-
Gnaphalineae (Compositae) de America del Sur, Bol. Soc. Arg. Bot.
9: 359-386. 1961; Flora de la provincia de Buenos Aires, 4(6): 443 pp.
1963. D. G. Drury, The American spicate cudweeds adventive to
New Zealand: (Gnaphalium Section Gamochaeta — Compositae),
New Zealand Journ. Bot. 9(1): 157-185. 1971.
Annual or perennial herbs, rarely suffrutescent, usually white-tomentose; leaves
alternate, mostly sessile, the blades entire, usually narrow; inflorescences composed
of cymes or glomerules of heads sessile at the ends of branches or in the upper leaf
axils, sometimes spikelike and sometimes forming large panicles; heads discoid;
involucres ovoid or campanulate; phyllaries several-seriate, numerous, imbricate,
thickened at the base, the larger upper portion usually hyaline, shining, rarely all of
the scarious tissue opaque, or sometimes the body of the phyllary membranous with
the tip thickened and opaque, the outermost ones often lanate, at least at the base;
receptacle flat to subconic, naked; outer pistillate corollas numerous, filiform,
minutely 3-4-cleft; inner hermaphrodite flowers few, the corollas yellow, white, or
purplish, tubular, 5-dentate; anthers sagittate at the base, the auricles caudate; style
branches of the disc flowers flattened, truncate at the apex; achenes very small, 0.4-
1.5 mm. long, oblong, subterete, or somewhat compressed, ecostate; pappus bristles
uniseriate, slender or sometimes dilated at the apex, falling separately from the
achene or sometimes connate at the base and then deciduous in a ring.
More than 100 often closely related and perplexing species of
cosmopolitan distribution, with 16 in Guatemala. The genus has
apparently been avoided by many students of the Compositae and,
at least in tropical America, the nomenclature is still in a rather
chaotic state; the interpretation and validity of many specific
names may well be questioned.
Pappus bristles united at the base into a ring, deciduous together from the achene.
Plants dwarf, 2-4 cm. high G. standleyi.
Plants much larger, the stems mostly 7-30 cm. long.
Plants developing long, slender stolons; middle and inner phyllaries white-tipped.
G. stolonatum.
Plants not stoloniferous; phyllaries not white-tipped.
168 FIELDIANA: BOTANY, VOLUME 24
Basal leaves in rosettes, conspicuous, persistent at flowering time; only the
lower leaf surfaces densely lanate, the upper ones thinly tomentose or
glabrate; heads 4-5 mm. high; involucres glabrate G. americanum.
Basal leaves not in rosettes, inconspicuous or withered at flowering time; both
leaf surfaces more or less lanate; heads about 3 mm. high; involucres
densely lanate G. pensylvanicum.
Pappus bristles always distinct, deciduous separately from the achene.
Phyllaries opaque, dull, papery, white, or the tips thickened, opaque and white.
Phyllaries completely opaque, dull, papery, white G. leucocephalum.
Phyllaries white-tipped, the tips thickened and opaque, the lower body of the
phyllary shining, sometimes rose-pink at the base of the white tip.
G. salicifolium.
Phyllaries shining, translucent (if white, neither opaque nor opaque- tipped).
Leaves mostly elliptic (rarely linear to linear-oblanceolate), attenuate to the
base, not dilated at the base nor decurrent on the stem G. attenuatum.
Leaves not elliptic, not attenuate to the base, more or less dilated and clasping
at the base (this not always evident in plants with linear leaves), sometimes
decurrent on the stem.
Leaf length mostly 0.6-1.5 (-2) cm., densely crowded on the stems; heads small,
the involucres about 3 mm. high G. brachyphyllum.
Leaf length 2-8 cm., often numerous but not conspicuously crowded on the
stem; heads larger, the involucres mostly 4-6 mm. high.
Leaves very narrowly linear, mostly 1-3 (-4) mm. broad (in ours).
Phyllaries mostly obtuse, or only the outermost ones acute; flowers 50-175
or more per head.
Leaf bases long-decurrent on the stem, forming a wing reaching to the
next node; upper leaf surfaces glabrate or thinly tomentose, only
the lower surface densely lanate; flowers 50-60 per head.
G. alatocaule.
Leaf bases clasping but not decurrent on the stem, both surfaces often
densely lanate; flowers 175 or more per head G. stramineum.
Phyllaries all acute or subacute; flowers 30-50 per head.
Leaves 15-20 times longer than broad; phyllaries rose or pink.
G. attenuatum var. silvicola.
Leaves 30-50 times longer than broad; phyllaries creamy white to
stramineous G. greenmanii.
Leaves linear, oblanceolate, linear-oblong, or sagittate, mostly 3-15 mm.
broad.
Phyllaries commonly pale yellow to deep reddish brown or rarely pinkish
when young (in G. liebmannii) but then soon becoming brown.
Phyllaries pale cream to yellow or stramineous; flowers commonly 200
or more per head (1,100-2,800 m.) G. viscosum.
Phyllaries light brown to deep reddish brown (rarely pinkish when
young); flowers commonly 40-80 per head (1,600-4,400 m., but
usually above 2,800 m.).
Plants mostly 10-30 cm. high; leaf apices acute; heads 6-8 mm. high;
hermaphrodite flowers commonly 5-7 (-10) per head.
G. liebmannii.
Plants often 50-100 cm. high; leaf apices acuminate; heads 4.5-6 mm.
high; hermaphrodite flowers 10-20 per head.
G. liebmannii var. monticola.
NASH: FLORA OF GUATEMALA 169
Phyllaries white or pink when fresh (sometimes turning yellow in treated
herbarium material).
Leaves concolorous or not conspicuously bicolored, both surfaces
commonly densely covered with white, feltlike tomentum; phyl-
laries obtuse or subacute G. roseum.
Leaves usually conspicuously bicolored, the upper surfaces commonly
green, only the lower surface densely white-tomentose; phyllaries
acute.
Heads small, the involucres 4-5 mm. high; hermaphrodite flowers
commonly 4-7 (-9) G. semiamplexicaule.
Heads larger, the involucres (5-) 6-7 mm. high; hermaphrodite flowers
commonly (7-) 10-14 G. brachypterum.
Gnaphalium alatocaule D. Nash, Fieldiana: Botany 36 (9):
73. 1974.
Known only from the type locality, 1,200-1,500 m., Carrizal,
Casillas, Santa Rosa, Heyde & Lux 4217 (type).
Coarse herbs, probably at least 1 m. tall, usually much branched in the area of
inflorescence, the stems terete, more or less arachnoid-tomentose; leaves sessile,
linear, acuminate, mostly 2-8 cm. long, 0.2-0.4 cm. wide, the base amplexicaul and
long-decurrent on the stem, forming a conspicuous wing reaching beyond the next
node, the margins often revolute, the upper surfaces green, glabrate or thinly
tomentose, eglandular, the lower surfaces densely covered with white, feltlike,
arachnoid tomentum; inflorescences broad and open, composed of numerous, small,
rounded, cymose glomerules of sessile heads at the ends of the stems and branches;
involucres campanulate, about 4 mm. long; phyllaries 4-5-seriate, glabrous, shining,
white or cream-colored, the outer ones ovate, acute, the middle ones lanceolate,
obtuse and often somewhat erose, the innermost ones linear obtuse or subacute and
often erose; heads each with 50-60 filiform, pistillate flowers and 7-10 tubular
hermaphrodite flowers; achenes brown, glabrous, about 0.5 mm. long; pappus white,
the bristles falling separately from the achenes.
Gnaphalium americanum Mill. Gard. Diet. ed. 8, no. 17. 1768.
Gamochaeta americana Weddell, Chlor. And. 1: 151. 1856. Gn.
purpureum L. var. americanum Klatt, Linnaea 42: 140. 1878. Gn.
guatemalense Gandoger, Bull. Soc. Bot. France 65: 42. 1918. G.
guatemalensis Cabrera, Bol. Soc. Arg. Bot. 9: 371. 1961. Chu-
chulken (Alta Verapaz, Quecchi); sacamal (Quezaltenango). Figure
44.
Dry or damp, open, often rocky places, pastures, sandy banks,
roadsides, cornfields, often in thickets, frequently in pine, pine-oak,
or Alnus forest, 400-3,800 m.; Alta Verapaz; Baja Verapaz;
Chimaltenango; Chiquimula; Guatemala; Huehuetenango; Jalapa;
El Progreso; Quezaltenango; El Quiche; Retalhuleu; Sacatepequez;
San Marcos; Solola; Totonicapan; probably in most of the other
departments. British Columbia through southern and eastern
170 FIELDIANA: BOTANY, VOLUME 24
United States; Mexico; El Salvador to Panama; West Indies; South
America; New Zealand.
Usually erect, rarely procumbent, usually biennial plants, mostly 15-20 cm. tall,
the stems solitary or several from each root, usually simple below the inflorescence,
white-tomentose; basal leaves often present and conspicuous, oblanceolate or
narrowly spathulate, to about 7 cm. long and 1 cm. wide, obtuse, mucronulate,
attenuate to the base, white-tomentose beneath, thinly tomentose or glabrate on the
upper surface, the stem leaves smaller, linear-oblong to spathulate, rather distant,
ascending, not decurrent at the base; heads numerous, sessile, usually forming an
elongated, spikelike inflorescence, at least the lower half leafy; involucres 4-5 mm.
high, glabrate, phyllaries 3-5-seriate, shining, pale or dark golden brown, the outer
ones ovate, acute, sometimes cuspidate, often reflexed, the inner ones oblong to
linear, acute; pistillate flowers numerous, the hermaphrodite ones 3-5; achenes
minute, less than 1 mm. long, the pappus about 3 mm. long, white, the bristles united
at the base and falling together from the achene.
Common, variable, weedy plants, often confused with G.
spicatum Lam., which has smaller heads, the involucres about 2.5
mm. high, and with G. purpureum L., which may be distinguished
(fide Drury) by its pink or purple inner phyllary tips and by its
wooly involucres. The relative size of these species is too variable for
this character to be very useful in diagnoses, at least in Central
American plants, although it is sometimes employed by other
workers.
Gnaphalium attenuatum DC. Prodr. 6: 228. 1837. Sanalatodo
(Escuintla); yucul Q'en (Quecchi, Alta Verapaz).
Damp or dry thickets or in forest, frequently in pine or pine-
oak forest, sometimes on brushy, rocky hillsides, grassy, open slopes,
clay banks, or in cornfields, 385-2,800 m., more common above 1,200
m.; Alta Verapaz; Baja Verapaz; Chimaltenango; Escuintla;
Guatemala; Huehuetenango; Jalapa; Jutiapa; Quezaltenango; El
Quiche; Sacatepequez; Santa Rosa; Solola; Suchitepequez; Zacapa.
Mexico; British Honduras; Honduras and El Salvador to Panama.
Erect perennials, mostly 0.5-2 m. tall, the stems simple or branched, densely
white-tomentose; leaves mostly elliptic, sometimes linear to linear-oblanceolate,
sometimes slightly falcate, mostly 3-7 (-8) cm. long, 0.4-1.5 cm. wide, attenuate to
each end, neither auriculate at the base nor decurrent on the stem, tomentose or
glabrate and green above, usually glandular-puberulent or short-villosulous under-
neath the tomentum, this short pubescence exposed in age, the lower surface densely
white-lanate; inflorescences commonly broad and open, paniculate, rarely a few
heads crowded into a small, rounded inflorescence; heads usually numerous, sessile or
nearly so, the involucres 4-5 mm. high; phyllaries shining, acute, stramineous to pale
yellow or creamy white, the innermost series 13-21; flowers 30-40, with hermaphrodite
flowers 3-7; pappus copious, white or brownish white, the bristles falling separately
from the achenes.
NASH: FLORA OF GUATEMALA 171
Gnaphalium attenuatum var. silvicola McVaugh, Contr.
Univ. Mich. Herb. 9: 465. 1972.
Usually in oak or pine-oak forest and damp thickets, sometimes
on grassy slopes, 1,200-2,500 m.; Baja Verapaz; Chimaltenango;
Chiquimula; Escuintla; Guatemala; Huehuetenango; Quezal-
tenango. Mexico.
Differs from the typical variety in its linear leaves (mostly 2-5
(-8) mm. wide, the margins revolute and the leaves therefore
appearing to be only 1-2 mm. wide, 15-20 times longer than broad)
and in its rose or pink phyllaries, the innermost series only 12-13.
Gnaphalium brachyphyllum Greenm. Field Mus. Bot. 2: 267.
1907. Figure 45.
Damp or dry, pine-oak or Cupressus forest, most common on
cliffs or steep, clay banks, sometimes along dusty roadsides, 2,000-
3,800 m.; Chimaltenango; Quezaltenango (type from Cerro Que-
mado, Kellerman 5301); El Quiche; San Marcos; Solola; Totonica-
pan.
Perennial, often suffrutescent plants, erect or pendent from banks or cliffs, the
stems usually very numerous and forming a dense tangled mass, slender, leafy, simple
or branched, mostly 10-35 cm. long, densely tomentose; leaves very numerous,
crowded on the stems, spreading, sessile, linear-oblong to oblong, mostly 0.6-1.5 (-2)
cm. long, 0.2-0.5 cm. wide, obtuse or acute, obtuse and subamplexicaul at the base,
densely tomentose; inflorescences terminal, dense, small, rounded, usually 3-4 cm.
broad; heads numerous, sessile; involucres about 3 mm. high; phyllaries 2-4-seriate,
glabrous except the outer ones tomentose at the base, pale-stramineous or yellowish,
shining, acute, the outer ones ovate, the inner ones lanceolate to linear; pistillate
flowers numerous; achenes glabrous, about 1 mm. long, reddish; pappus bristles
white, deciduous separately from the achene.
Gnaphalium brachypterum DC. Prodr. 6: 226. 1837.
Damp or dry, sometimes rocky, open banks or slopes, in
thickets, or in oak or pine-oak forest, 1,100-3,300 m. (most common
below 2,800 m.); Alta Verapaz; Baja Verapaz; Chimaltenango;
Guatemala; Huehuetenango; Jalapa; Quezaltenango; El Quiche;
Sacatepequez; San Marcos; Totonicapan. Mexico; Honduras; Nica-
ragua; Costa Rica; Panama.
Erect, robust plants, mostly 0.5-1 m. tall, the stems simple or branched, densely
covered with white, arachnoid tomentum; leaves sessile, oblanceolate to linear-
oblong, mostly 1.5-6 cm. long, 0.5-1 cm. broad, acute, dilated at the base and
amplexicaul, conspicuously bicolored, the upper surfaces commonly green and often
glandular-pubescent and scabrous, sometimes sparsely tomentose, the lower surfaces
172 FIELDIANA: BOTANY, VOLUME 24
densely white-tomentose; inflorescences large and ouen or dense and crowded,
composed of glomerules of few-many heads; involucres (5-) 6-7 mm. high; phyllaries
4-5-seriate, shining, white or cream-colored when fresh (some treated specimens
turning yellow in herbarium material), acute; flowers commonly 50-60 per head, with
(7-) 10-14 hermaphrodite flowers; pappus sordid, the bristles falling separately from
the glabrous achenes.
Gnaphalium greenmanii Blake, Journ. Wash. Acad. Sci. 21:
329. 1931. G. linearifolium Greenm. Proc. Am. Acad. 32: 308. 1897,
not G. linearifolium (Wedd.) French. 1892.
Usually in oak or pine-oak forest, sometimes on open banks,
1,150-2,400 m.; Baja Verapaz; Chimaltenango; Escuintla; Huehue-
tenango; Jalapa; Zacapa. Mexico.
Erect annuals, the stems simple or branched, terete, white-lanate, mostly 50-100
cm. tall; leaves sessile, narrowly linear, mostly 4-10 cm. long, 1-2 (-3) mm. wide,
acuminate, somewhat dilated and clasping at the base, the margins revolute, green
above but also more or less arachnoid-pubescent, the lower surfaces densely white-
tomentose; inflorescences corymbose; heads sessile or nearly so in small, dense
glomerules; involucres 4-5 mm. high; phyllaries about 4-seriate, creamy white to
stramineous, the outermost ones ovate and acute, the innermost ones oblong and
subacute; flowers commonly 30-50 per head, the hermaphrodite flowers 5-10 per
head; pappus bristles falling separately from the glabrous achenes.
Gnaphalium leucocephalum A. Gray, Smiths. Contr. 5(6): 99.
1853.
Open, dry banks or damp ravines, mixed or oak forest, 1,800-
1,900 m.; Huehuetenango. Southwestern United States; Mexico.
Erect or ascending perennials, the stems solitary or several, simple or branching,
mostly 20-60 cm. tall, densely lanate and glandular-puberulent beneath the heavy
indument, leafy to the top; leaves sessile, narrowly linear (in ours) or sometimes
narrowly linear-lanceolate, mostly 2-4 (-6) cm. long, acute or acuminate, the bases
shortly decurrent on the stem, the margins often somewhat revolute, the upper
surfaces green and more or less glandular or sparsely or densely lanate; inflorescences
terminal, short, glomerate-corymbiform; heads with 60-80 pistillate flowers and 10-15
hermaphrodite flowers; involucres 4-6 mm. high; phyllaries multiseriate, pearly white,
opaque, the outermost ones ovate and acute, the others obovate to linear and obtuse;
corollas yellowish; achenes minute; pappus white, the bristles falling separately from
the achene.
Gnaphalium liebmannii Sch. Bip. ex Klatt, Leopoldina 23: 89.
1887. G. vulcanicum I. M. Johnston, Contr. Gray Herb. 68: 100.
1923.
Open, rocky slopes or sandy soil near summits of volcanoes,
sometimes in grassy meadows or damp thickets, or in Juniperus
forest, alpine or subalpine locations, 2,900-4,400 m.; Chimaltenango;
NASH: FLORA OF GUATEMALA 173
Huehuetenango; Quezaltenango; Sacatepequez; San Marcos;
Solola; Totonicapan. High mountains of southern Mexico and Costa
Rica.
Stout herbs, forming basal rosettes, the stems solitary or several, simple or
branched, mostly 10-30 cm. high (rarely the dwarf form 2-3 cm. high where sheep
have regularly grazed), densely and laxly lanate-tomentose; leaves lanate-tomentose
and glandular at least on the upper surfaces (the tomentum often obscuring the
glandular condition), the basal ones spathulate or oblanceolate, mostly 5-7 cm. long,
obtuse, the cauline leaves oblanceolate to linear-oblong, mostly 3-6 cm. long, acute,
more or less dilated and auriculate at the base, sometimes short-decurrent on one
side; inflorescences globose-glomerate at the ends of the stems or sometimes several
of these forming a small, more or less corymbiform inflorescence; heads numerous, 6-
8 mm. high; phyllaries 3-4-seriate, shining, pale brown or reddish brown, sometimes
roseate at first but in age usually dark golden brown or dark reddish brown, acute,
the outer ones ovate, somewhat lanate at the base, elsewhere glabrous; flowers
commonly 40-80 per head, with (3-) 5-7 (-10) hermaphrodite flowers; achenes oblong,
glabrous or nearly so, about 0.8 mm. long; pappus bristles about 4 mm. long, falling
separately from the achene.
Floral counts were made of five heads taken respectively from
the type of G. liebmannii (Liebmann 310), the type of G.
vulcanicum (Purpus 2782), from a specimen cited by Johnston as G.
vulcanicum (Purpus 3033), from a dwarfed specimen (Degener &
Degener 26704), and one from Steyermark 50163. The first count
was 50 (including five hermaphrodite flowers); the second was 80
(including seven hermaphrodite flowers), the third, 40 (including
five hermaphrodite flowers); the fourth 64 (including three
hermaphrodite flowers), and the fifth, 68 (including 10 hermaphro-
dite flowers).
Gnaphalium liebmannii var. monticola (McVaugh) D. Nash,
Fieldiana: Bot. 36(9): 74. 1974. G. vulcanicum I. M. Johnston var.
monticola McVaugh, Contr. Univ. Mich. Herb. 9: 466. 1972.
Sanatodo (Solola).
Open forest or damp thickets, 1,600-3,150 m.; Chimaltenango;
Huehuetenango; El Progreso; Quezaltenango; Sacatepequez; San
Marcos; Solola; Totonicapan. Mexico; Costa Rica.
Differs from the typical variety in its larger size, often 50-100
cm. high, its usually acuminate leaves that are conspicuously
glandular on the upper surfaces due to less indument, its usually
smaller heads, 4.5-6 mm. high, and the greater number of
hermaphrodite flowers, commonly 10-20 per head.
Gnaphalium pensylvanicum Willd., Enum. PL Hort. Berol.:
867. 1809. Gn. spathulatum Lamarck, Encycl. Meth. 2: 758. 1786
(not Burm. f., 1768). Gn. purpureum var. spathulatum Baker in
174 FIELDIANA: BOTANY, VOLUME 24
Mart. Fl. Brasil. 6(3): 125. 1882. Gn. peregrinum Fern. Rhodora 45:
479. t. 795. 1943. Gamochaeta pensylvanica Cabrera, Bol. Soc. Arg.
Bot. 9: 375. 1961.
Damp or dry, often rocky or sandy places, along streams or on
brushy oak slopes, sometimes a weed in streets, 1,350-2,500 m.;
Guatemala; Jalapa; Quezaltenango. Southern United States;
Honduras; Costa Rica; West Indies; South America; New Zealand.
Annual herbs, the stems ascending or suberect, or sometimes more or less
decumbent, often diffusely branching from the base, the stems leafy, laxly
lanuginose; leaves more or less spathulate or obovate, usually narrow, mostly 1-4 cm.
long, acute or obtuse, lanate on both surfaces but more densely so beneath;
inflorescences spikelike, leafy, commonly interrupted; heads numerous in small,
compact glomerules; involucres lanate at the base, about 3 mm. high; phyllaries 2-3-
seriate, at least some of them obtuse or subobtuse (usually the middle and/or inner
ones), the outer ones often acute, pale stramineous to light brownish; achenes oblong,
minutely papillose; pappus bristles white, united at the base into an annulus and all
falling together from the achene.
Variable in form, this species ranges in size from small plants
less than 10 cm. tall to the more common ones about 30 cm. tall.
Gnaphalium roseum HBK. Nov. Gen. & Sp. 4: 81. 1820.
Oak or pine-oak forest, 1,500-2,100 m.; Chimaltenango; Chi-
quimula; Escuintla; Huehuetenango; Jalapa; El Quiche; Sacatepe-
quez; Zacapa. Mexico.
Erect, usually rather stout perennials, mostly 30-70 cm. high, the stems usually
simple, branching only in the area of inflorescence, densely white-tomentose; leaves
sessile, oblanceolate or linear-oblanceolate, acute, somewhat dilated at the base and
semiamplexicaul, densely white-tomentose to white-lanate on both surfaces;
inflorescences composed of numerous, rounded glomerules of heads, the heads
numerous, crowded, sessile or nearly so; involucres 4-5 mm. high; phyllaries several-
seriate, shining, pink or white (sometimes both colors on the same plant), commonly
obtuse but sometimes acute or subobtuse or the outermost ones acute and the others
obtuse; flowers commonly less than 50 per head with 3-7 hermaphrodite flowers;
pappus white, the bristles falling separately from the achene.
According to the original description, the phyllaries of this
species are "acute." However, a note by S. F. Blake on U. S.
National Herbarium sheet No. 460996, Pringle 11533, reads, "Fair
match for type of G. roseum at Paris, 1925." The phyllaries of this
collection are all obtuse with the exception of the outermost ones,
some of which are acute.
Although reported from Costa Rica by Standley, the Costa
Rican specimens I have seen that are so annotated by him, all from
volcanic summits, above 3,200 m., are referable to G. liebmannii
NASH: FLORA OF GUATEMALA 175
Sch. Bip. The several varieties described by Bentham (Oerst. Vid.
Medd. Kjoebenhavn 1852: 105. 1853) appear not to be this species.
Gnaphalium salicifolium (Bertol.) Sch. Bip. Bot. Zeit. 3: 172.
1845. Helichrysum salicifolium Bertol. Fl. Guat. 433. 1840. G.
rhodanthum Sch. Bip. in Seem. Bot. Voy. Herald 310. 1856.
Damp or wet meadows or banks, often on sandy soil and
exposed, rocky places, sometimes in thickets, open, mixed forest,
coniferous, scrub, or oak-pine forest, 2,300-4,500 m.; Chimaltenango;
Guatemala; Huehuetenango; El Progreso; Quezaltenango; El
Quiche; Sacatepequez (type from Volcan de Agua, Velasquez}; San
Marcos; Solola; Totonicapan. Mountains of Southern Mexico and
Costa Rica.
Erect perennials, often suffrutescent, simple or branched, the stems densely
tomentose to lanate, mostly 10-40 cm. tall; leaves very numerous, sessile, spreading or
ascending, linear or lance-linear, mostly 2-8 cm. long, 0.1-0.5 cm. wide, acute or
obtuse, attenuate to the base but then the base slightly dilated, clasping, and more or
less decurrent on the stem, the margins often revolute, white beneath with dense,
often sericeous, long tomentum or sometimes lanate, laxly tomentose or glabrate on
the upper surface; heads numerous, short-pedicellate or subsessile, glomerate and
forming small, dense, rounded inflorescences; involucres about 5 mm. high; phyllaries
strongly graduate, the outer ones short, laxly tomentose, the inner ones linear, much
longer, erect, the body shining and translucent, the tip thick, opaque, pure white,
sometimes rose-pink just below the white tip; pappus bristles falling separately from
the achene.
Gnaphalium semiamplexicaule DC. in DC. Prodr. 6: 228.
1837.
Wet to dry thickets or forest, sometimes in pine or pine-oak
forest, 500-2,850 m. (most common above 1,500 m.); Baja Verapaz;
Chimaltenango; Chiquimula; Escuintla; Guatemala; Huehue-
tenango; Jalapa; Quezaltenango; El Quiche; Sacatepequez; San
Marcos; Solola; Totonicapan; Zacapa. Mexico; British Honduras;
Honduras; El Salvador; Nicaragua.
Erect, often robust perennials mostly 30-75 cm. high, simple or branched, the
stems densely covered with white, arachnoid tomentum; leaves sessile, the blades
lanceolate, lance-oblong, linear-oblong, or linear-lanceolate (very rarely narrowly
linear), mostly 2-7 cm. long and 0.5-1 cm. wide, acute or ocuminate, a little attenuate
to the somewhat dilated, auriculate and semiamplexicaul base, or not at all
attenuate, commonly arachnoid-tomentose on both surfaces but usually much more
densely so beneath and therefore usually conspicuously bicolored, rarely glabrate and
scabrous above; inflorescences corymbiform or large and paniculate, the glomerules
of heads numerous, rounded, the heads crowded; involucres 4-5 mm. high, glabrous
except at the base, there somewhat lanate; phyllaries 4-5-seriate, shining, white or
cream-colored (sometimes treated herbarium material darker, becoming stramineous
176 FIELDIANA: BOTANY, VOLUME 24
or yellowish), mostly acute, sometimes the middle and/or inner ones obtuse; flowers
more than 50 per head, the hermaphrodite flowers commonly 4-7 (-9); achenes
minute, brown, glabrous; pappus white or yellowish white, the bristles falling
separately from the achene.
According to the original description, the pistillate flowers were
numbered at only 12-15, due probably to the fact that the type
specimens are immature. A head removed from the isotype
(Berlandier 2188) was treated with glycerine, the very young florets
allowed to float free, then counted with the aid of a compound
microscope; more than 50 were clearly seen.
This species is very similar to G. bicolor Bioletti (1893) of the
western United States, which may prove to be a synonym, and to G.
oxyphyllum DC. (1837) which differs in having only 35-50 flowers
per head. G. semiamplexicaule has also been confused in herbaria
with G. brachypterum DC. which may be distinguished by its larger
involucres, mostly 6-7 mm. high, and the greater number of
hermaphrodite flowers, commonly 10-14.
Gnaphalium standleyi Steyerm. Field Mus. Bot. 23: 99. 1944.
Sacahuax (Huehuetenango).
Damp or wet alpine meadows or slopes, sometimes in Juniperus
forest, 2,700-4,300 m.; Huehuetenango (type from region of Chemal,
Sierra de los Cuchumatanes, Standley 81097); San Marcos; Solola;
Totonicapan.
Plants probably annual, 2-4.5 cm. high, densely white-tomentose throughout, the
tomentum close and appressed; leaves mostly clustered in a basal rosette, the blades
elliptic-obovate to oblanceolate or spathulate, mostly 6-18 mm. long, 2-6 mm. wide,
rounded and mucronulate at the apex, attenuate to the base, the cauline leaves 2-3 or
none; heads small, lanate at the base, disposed in terminal, spikelike inflorescences 6-
25 mm. long and 5-13 mm. broad; involucres about 3 mm. high; phyllaries 4-seriate,
lustrous, brownish tipped, the outer ones ovate, acute, the inner oblanceolate,
spathulate, or nearly linear, subacute; flowers 60-70 per head; achenes dark brown,
hispidulous-tuberculate, about 0.8 mm. long; pappus bristles about 2.5 mm. long,
united at the base into an annulus and all falling together.
Gnaphalium stolonatum Blake, Brittonia 2: 341. 1937.
Alpine meadows, often on outcrops of limestone, 3,100-4,030 m.;
Huehuetenango (Sierra de los Cuchumatanes, the type collected
along trail between Huehuetenango and Soloma, probably in the
region of Chemal, 3,150 meters, Skutch 1098); San Marcos.
Perennials, mostly 10-30 cm. tall, the stems thinly arachnoid-lanate, simple or
nearly so, producing at the base and from the lowest leaf axils 1-2 filiform stolons 3-
10 cm. long, these rooting at the apex; leaves basal and cauline, oblanceolate to
linear, the lowest 10-40 mm. long and 2-4 mm. wide, the upper ones usually smaller,
NASH: FLORA OF GUATEMALA 177
thinly arachnoid-lanate and sparsely stipitate-glandular above or glabrate, densely
lanate-tomentose beneath; heads sessile, many-flowered, crowded in a subglobose to
ovoid or short-cylindrical glomerule 1-2 cm. in diameter; involucres 4-6 mm. high;
phyllaries 4-seriate, the outer ones thinly arachnoid at least at the base, ovate, with
dark tips and margins, the middle and inner ones oblong-lanceolate to almost linear,
subobtuse or acute, white-tipped and spreading in age; pistillate corollas about 2.5
mm. long; achenes quadrangular-ellipsoid, brownish, about 1 mm. long, glabrous or
very minutely hispidulous; pappus bristles about 3 mm. long, white, united at the
base into an annulus and all falling together from the achene.
Characteristic alpine plants of the Cuchumatanes, in general
appearance similar to many species of Antennaria, a dioecious
genus.
Gnaphalium stramineum HBK. Nov. Gen. & Sp. 4: 85. 1820.
G. chilense Spreng. Syst. 3: 480. 1826. G. sprengelii Hook. & Arn.
Bot. Voy. Beechey 150. 1833. G. berlandieri DC. in DC. Prodr. 6:
223. 1838. G. chilense var. confertifolium Greene, Fl. Fran. 400. 1897.
Pine or pine-oak forest, 1,800-2,400 m.; Escuintla; Guatemala;
Huehuetenango; Zacapa. Western United States; Mexico.
Usually aromatic herbs from strong taproots, the stems solitary or several, erect
or rarely decumbent, more or less floccose-lanate throughout, usually densely so;
basal and lowermost cauline leaves spathulate to oblong, usually obtuse, the middle
and upper cauline leaves mostly linear, mostly 2-5 cm. long, 0.2-0.4 cm. broad (in
ours), usually obtuse, somewhat dilated and clasping at the base, short-decurrent on
the stem, densely lanate; inflorescences compact, headlike, the heads crowded into
small glomerules; involucres 4-6 mm. high; phyllaries shining, first whitish but soon
becoming stramineous or yellow, ovate to obovate and obtuse or the outermost ones
acute and the innermost ones linear and obtuse; flowers 175 or more per head, with
15-20 or more hermaphrodite flowers; pappus bristles falling separately from the
achenes.
A variable species, in the United States and Mexico found in
various habitats, a form with larger leaves to about 8 mm. broad
often collected there.
Gnaphalium viscosum HBK. Nov. Gen. & Sp. 4: 82. 1820. G.
hirtum HBK. I.e. G. tenue HBK. torn. cit. 83. G. leptophyllum DC. in
DC. Prodr. 6: 226. 1838. Flor de la seda, sac-moquan (Huehue-
tenango); sanatodo (Chiquimula): sanalotodo (Sacatepequez).
Brushy or grassy slopes, open fields, sandy soil of clearings or
thickets, sometimes in pine-oak forest, 1,100-2,800 m.; Chimal-
tenango; Chiquimula; Guatemala; Huehuetenango; Jalapa; Quezal-
tenango; El Quiche; Sacatepequez; San Marcos; Solola; Zacapa.
Western Texas; Mexico; Honduras.
Erect annuals, simple or branched, mostly 30-100 cm. high, more or less viscid
throughout, the stems laxly tomentose and glandular-pubescent, usually densely
178 FIELDIANA: BOTANY, VOLUME 24
leafy; leaves ascending or spreading, narrowly sagittate or narrowly linear-lanceolate,
mo'-My 4-7 cm. long, 0.3-1 cm. broad, acute or acuminate, usually somewhat dilated
at i.j base, amplexicaul, and often long-decurrent on the stems, green above and
strongly glandular-pubescent, densely white-tomentose beneath with close, arachnoid
tomentum; 'nflorescences composed of glomerules of heads; involucres globose-
campanula^, about 5 mm. high; phyllaries pale yellowish or creamy white, ovate to
lanceolate and acute or the innermost ones linear and subacute; flowers commonly
200 or more per head, with 5-15 hermaphrodite flowers; mature receptacles 3-4 mm.
wide; pappus bristles falling separately from the glabrous achenes.
PLUCHEA Cassini
Reference: R. K. Godfrey, Pluchea, section Stylimnus, in North
America, Journ. Elisha Mitchell Soc. 68(1): 238-272. 1952.
Annual or perennial herbs or shrubs, usually tomentose, villous, or floccose, often
viscid; leaves alternate, penninerved, the margins entire, dentate, or serrate;
inflorescences cymose-corymbose, the heads numerous, heterogamous, discoid;
involucres broadly campanulate or hemispheric; phyllaries few-multiseriate, imbri-
cate, ovate or lanceolate, dry, submembranaceous; receptacle flat, naked; outer,
pistillate flowers numerous, multiseriate, fertile, the inner flowers usually few,
hermaphrodite, sterile; corollas mostly pink or purplish but sometimes white, the
pistillate ones filiform, trifid, the hermaphrodite ones tubular, the limb little dilated,
5-dentate; anthers sagittate at base, the auricles acuminate-caudate; styles of the
hermaphrodite flowers simple or bifid, hirtellous or papillose; achenes very small, not
more than 1 mm. long, glabrous or pubescent, cylindrical, 4-5-angulate; pappus
uniseriate, the bristles slender, free or more or less connate at the base.
Probably about 20 species, in temperate and tropical regions of
North and South America, Africa, and Asia to Australia. Although
only two occur in Guatemala, a third species from British Honduras
is also treated here.
Leaves linear-lanceolate, linear, or linear-elliptic, commonly 9-10 times longer than
broad, conspicuously decurrent on the stems P. salicifolia.
Leaves ovate-oblong to lanceolate or oblong-elliptic, commonly only 3-5 times longer
than broad, petiolate and not decurrent on the stems.
Plants woody; leaves mostly 7-15 cm. long, the margins mostly entire or nearly so;
inflorescences large, mostly 6-15 cm. broad P. odorata.
Plants herbaceous; leaves mostly 5-8 cm. long, the margins mostly undulate-serrate
or crenate-serrate; inflorescences small, mostly 3-6 cm. broad.
P. purpurascens.
Pluchea odorata (L.) Cass. Diet. Sci. Nat. 42: 3. 1826;
Godfrey, Journ. Elisha Mitchell Soc. 68(1): 247. 1952. Conyza
odorata L. Syst. ed. 10. 1213. 1760. C. cortesii HBK. Nov. Gen. &
Sp. 4: 75. 1820. Pluchea cortesii DC. in DC. Prodr. 5: 452. 1836.
Siguapate, Santa Maria, chalche (fide Standley); sesc > oh
(Quecchi, Alta Verapaz). Figure 46.
NASH: FLORA OF GUATEMALA 179
Dry to wet thickets, on plains or hillsides, most abundant in
sandy or rocky places along stream beds, sometimes in pine or oak
forest, rarely in hedges, sea level to 2,500 m.; Alta Verapaz; Baja
Verapaz; Chiquimula; Escuintla; Guatemala; Huehuetenango;
Izabal; Jalapa; Peten; El Progreso; Quezaltenango; El Quiche;
Retalhuleu; Sacatepequez; San Marcos; Santa Rosa; Solola;
Zacapa. Southern Florida; Mexico; British Honduras to El Salvador
and Panama; West Indies; northern South America; naturalized in
Hawaii.
Erect, usually much branched shrubs, commonly 1-2.5 m. tall, the branches
stout, densely tomentose; leaves on stout petioles 1-2.5 cm. long, the blades ovate-
oblong to elliptic, mostly 7-15 cm. long, 2.5-6 cm. broad, narrowed to the obtuse,
mucronulate apex, usually attenuate to the base, rarely obtuse, entire or nearly so,
thinly tomentulose to short- pubescent or glabrate above, densely and laxly sordid-
tomentose beneath; inflorescences rather large, broad corymbs, mostly 6-15 (-20) cm.
broad, the pedicels and peduncles stout and densely tomentose; heads campanulate,
about 7 mm. high and 6-9 mm. broad; phyllaries 5-6-seriate, viscid-tomentose and at
least the outer ones ciliate, the outermost ones ovate, obtuse, the inner and middle
ones more or less oblong-lanceolate, and the innermost ones linear, attenuate and
glabrous; corollas purplish; achenes minute, less than 1 mm. long, the pappus bristles
soft, dirty white, 3-4 mm. long.
Aromatic plants, a decoction of the leaves much used as a
remedy for affections of the stomach.
Pluchea purpurascens (Sw.) DC. in DC. Prodr. 5: 452. 1836;
Godfrey, Journ. Elisha Mitchell Soc. 68 (1): 247. 1952. Conyza
purpurascens Swartz, Prodr. Fl. Ind. Occ. 112. 1788. Santa Maria
cimarrona.
Open, damp, weedy places, at or little above sea level; British
Honduras. Southern United States; Mexico; Honduras; Nicaragua;
Panama; West Indies; Colombia; Venezuela; British Guiana.
Coarse, erect, annual herbs to about 1 m. tall, simple or much branched, the
stems thinly puberulent or glabrate below, striate, often purplish, abundantly
puberulent above; leaves commonly short-petiolate, sometimes sessile, the blades
ovate-oblong to lanceolate or elliptic, mostly 5-8 cm. long, obtuse or acute, acute or
obtuse at the base, the margins undulate-serrate or crenate-serrate, commonly more
or less puberulent on both surfaces; inflorescences corymbose, mostly 3-6 cm. broad,
the pedicels slender, puberulent; heads usually few, mostly 5-6 mm. high and about as
broad; involucres campanulate; phyllaries 4-seriate, the outer ones ovate, puberulent,
ciliate, the middle and inner ones oblong-lanceolate, the innermost ones linear-
oblong, acuminate, scarious, viscid; corollas 3-4 mm. long, purplish; achenes about 1
mm. long; pappus soft, white, about 3 mm. long.
Pluchea salicifolia (Mill.) Blake, Contr. U. S. Nat. Herb. 26:
237. 1930; Godfrey, Journ. Elisha Mitchell Soc. 68 (1): 250. 1952.
180 FIELDIANA: BOTANY, VOLUME 24
Conyza salicifolia Miller, Gard. Diet. ed. 8, Conyza No. 6. 1768. P.
subdecurrens Cass. Diet. Sci. Nat. 42: 4. 1826.
Along stream beds and on damp or dry slopes, Baja Verapaz,
about 600 m. Mexico.
Suffrutescent plants 1-2 m. tall, branching, the stems glandular and more or less
pilose, winged with decurrent leaf bases, and with dense tufts of silky hairs in the leaf
axils; leaves sessile, clasping and long-decurrent on the stem, the blades linear, linear-
oblong, or linear-lanceolate, mostly 6-12 cm. long, commonly 9-10 times longer than
broad, acute, the margins unevenly serrate-dentate or sometimes essentially entire,
both surfaces minutely glandular-punctate, sparsely pubescent or glabrate above,
more or less pilosulous beneath, especially on costae and veins; inflorescences
cymose-corymbose, sometimes becoming paniculiform, mostly 5-15 cm. broad, the
pedicels and peduncles long-pilose to arachnoid-tomentose; heads usually numerous,
4-5 mm. high and broad; involucres campanulate; phyllaries about 4-seriate, the
outer and middle ones lanceolate, glandular and pilose, the inner ones linear,
acuminate, glabrous; corollas pink or white; achenes less than 1 mm. long; pappus
dirty white, about 3 mm. long.
PSEUDOCONYZA Cuatrecasas
Reference: Jose Cuatrecasas, Pseudoconyza, in "Notas sobre
Astereas Andinas", Ciencia (Mex.) 21: 30-32. 1961; Supplemental
characterization of the genus Pseudoconyza (Compositae, Inuleae-
Pluchiinae), Phytologia 26: 410-411. 1973. Figure 47.
Erect herbs with alternate, more or less lyrate-pinnatifid leaves; heads
heterogamous, discoid, with numerous (180-280) pistillate flowers and 8-12 sterile,
hermaphrodite flowers in the center; receptacle flat, alveolate; phyllaries about 4-
seriate, narrow, imbricate, unequal, the inner ones with scarious margins, the outer
ones much shorter, subherbaceous; corollas of the hermaphrodite flowers tubular,
white, the limb briefly 5-dentate; anthers caudiculate; style branches linear-subulate,
hispid; corollas of the pistillate flowers filiform, tridentate; styles short, with the
branches minutely papillose; ovary oblong; achenes almost terete, contracted just
below the apex, shining but hispid; pappus uniseriate with 10-12 scabridulous bristles.
Only the following species is recognized at present.
Pseudoconyza viscosa (Mill.) D'Arcy, Phytologia 25: 281.
1973. Conyza viscosa Mill. Gard. Diet. ed. 8. 1768. C. lyrata HBK.
Nov. Gen. & Sp. 4: 70. 1820. Erigeron lyratum Gomez, Ann. Hist.
Nat. Madrid 19: 272. 1890. Eupatorium lyratum Coulter, Bot. Gaz.
16: 96. 1891 (type from Lago de Amatitlan, Guatemala, J. D. Smith
2393). C. lyrata var. pilosa Fern. Proc. Am. Acad. 36: 506. 1901.
Eschenbachia lyrata Britt. & Millsp. Fl. Baham. 444. 1920. C.
chiapensis Brandg. Univ. Calif. Publ. Bot. 10: 419. 1924. Blumea
lyrata Badillo, Bol. Soc. Venez. Cienc. Nat. 10: 257. 1946.
NASH: FLORA OF GUATEMALA 181
Pseudoconyza lyrata Cuatr. Ciencia (Mex.) 21: 31. 1961. Laggera
lyrata Liens, Mitt. hot. Staatssaml. Munchen 6: 107. 1971. Tabaco
cimarrbn (Jutiapa); tabaquillo and vitalino (Santa Rosa).
Wet to dry fields or thickets, sometimes on sandbars or in very
sandy soil, often a weed about houses or in cultivated ground,
sometimes growing in streets, on adobe or rock walls, or on salt
flats, sea level to 1,800 m.; Escuintla; Guatemala; Jutiapa; El
Progreso; Retalhuleu; San Marcos; Santa Rosa; Zacapa. Mexico; El
Salvador to Panama; western South America.
Erect annuals, viscid throughout, ill-scented, mostly 15-60 cm. tall (rarely
attaining more than 1 m.), usually much branched, the branches commonly densely
villous-hirsute with long, spreading hairs and glandular-pilose; leaves short-petiolate
or the upper ones sessile, the blades thin, oblong-ovate or spathulate-oblanceolate, at
least the lower ones more or less lyrate-pinnatifid near the base with few segments,
the upper ones mostly obovate and often only coarsely dentate, obtuse or acute,
commonly attenuate to the base, mostly 2-10 cm. long, 1-4 cm. wide, densely pilose
and glandular-puberulent on both surfaces; inflorescences cymose, the area of
inflorescence sometimes large and appearing paniculate; heads few or numerous,
sometimes rather densely clustered near and at the ends of the branches, slender-
pedicellate; involucres 6-7 mm. high, densely pilose and glandular; phyllaries about 4-
seriate, linear-lanceolate and acute to linear and acuminate, the outer ones much
shorter than the inner ones, the margins scarious; achenes fuscous, about 1 mm. long,
commonly villous, sometimes glabrate, contracted just below the apex; pappus about
4 mm. long.
Common and unpleasant weeds of the Central American
lowlands, very viscid, adhering easily to clothing, and with a most
disagreeable odor.
TRIBE V. HELIANTHEAE
By DOROTHY L. NASH
Annual or perennial herbs, shrubs, or trees, the plants usually dioecious (rarely
monoecious, Ambrosia); leaves usually opposite or at least the lower ones usually so,
rarely alternate, the blades simple, lobate, incised, or dissected; inflorescences of
solitary heads or cymose and often paniculate; heads more commonly heterogamous
and radiate, sometimes the pistillate flowers lacking rays, or sometimes the heads
homogamous and discoid; involucres usually campanulate or hemispheric, sometimes
orbicular or ovoid; phyllaries commonly distinct, in 2-6 graduated series, rarely the
involucres tubular and gamophyllous; receptacles usually paleaceous throughout,
sometimes only on the margin, rarely epaleaceous, the pales varying from filiform or
setaceous and rigid and merely subtending the flowers, to broad, scarious, or
membranous, embracing or enfolding the flowers; ray corollas usually ligulate,
sometimes merely a slender tube, the ligules often yellow or white but may also be
variously colored; disc corollas regular, tubular, the limb 4-5-cleft, commonly yellow,
sometimes orange, white, greenish, or rarely bluish or purplish to almost black;
anthers usually more or less sagittate at the base (rarely entire), with usually
182 FIELDIANA: BOTANY, VOLUME 24
somewhat concave apical appendages; style branches various but usually acute,
sometimes obtuse, rarely truncate, terete or flattened, often hirtellous; achenes
usually thick and hard, sometimes flattened, sometimes winged; pappus commonly of
2-many, more or less rigid owns and/or scales or sometimes absent.
This is the largest and most diverse tribe in the Compositae,
with 67 genera in Guatemala.
1. Plants monoecious; pistillate heads 1 -few-flowered, the staminate heads many-
flowered Ambrosia.
I. Plants not monoecious 2.
2. Heads 1-2-flowered, the involucres tubular or compressed and orbicular 3.
2. Heads several-many-flowered, or if 1-flowered, the involucres not as above 4.
3. Involucres tubular, gamophyllous Lagascea.
3. Involucres compressed, orbicular, the phyllaries distinct Delilea.
4. Ray flowers (when present) pistillate and fertile; hermaphrodite disc flowers
sterile 5.
4. Ray flowers (when present) pistillate and fertile or neutral; hermaphrodite disc
flowers fertile 16.
5. Plants scandent; leaves compound or appearing so, with few large leaflets.
Hidalgoa.
5. Plants not scandent; leaves simple, lobate, or pinnatifid, but not compound 6.
6. Heads discoid 7.
6. Heads radiate 8.
7. Achenes loosely enclosed in sac-shaped phyllaries Desmanthodium.
7. Achenes not enclosed in the phyllaries Clibadium.
8. Fruits echinate Acanthospermum.
8. Fruits not echinate 9.
9. Phyllaries subtending the ray achenes completely or almost completely enclosing
the mature achenes 10.
9. Phyllaries subtending the ray achenes sometimes embracing but never completely
enclosing the mature achenes 12.
10. Ray achenes enclosed by indurate phyllaries, these often beaked or appendaged.
Melampo di um.
10. Ray achenes enclosed by somewhat fleshy phyllaries, these neither beaked nor
appendaged 11.
II. Leaves (in ours) pinnatifid or bipinnatifid; heads with more than one ray flower;
phyllaries enclosing the ray achenes splitting into 3 parts at maturity.
Parthenium.
11. Leaves never pinnatifid; heads with only one ray flower; phyllaries enclosing the
ray achenes not splitting at maturity Milleria.
12. Plants shrubs or small trees; ray achenes obcompressed, conspicuously winged,
the wings prolonged above into 2 awnlike, lacerate teeth Rensonia.
12. Plants commonly herbaceous, rarely shrubby; ray achenes obovoid to spherical
or triquetrous, not winged 13.
13. Ray flowers commonly 3, the ligules broad, more or less fan-shaped (in ours),
deeply trilobate Trigonospermum.
13. Ray flowers commonly 5-20, the ligules relatively narrow, more or less strap-
shaped or triangular, shallowly bifid or tridentate 14.
NASH: FLORA OF GUATEMALA 183
14. Leaves usually more or less angulate, lobate, or pinnate-lobate; petioles
frequently winged and often dilated and clasping at the base (rarely not
winged); ray achenes obovoid to spherical Polymnia.
14. Leaves neither angulate, lobate, nor pinnate-lobate; petioles not winged; ray
achenes triquetrous or dorsally compressed 15.
15. Ray corollas with a definite tube, the ligules strap-shaped; achenes triquetrous.
Baltimora.
15. Ray corollas lacking a definite tube, sessile on the achene, the ligules triangular,
acute; ray achenes dorsally compressed Tragoceras.
16. Ray corollas lacking a distinct tube, sessile on the achenes, usually persistent.
17.
16. Ray corollas (when present) with a distinct tube, deciduous 20.
17. Rays as long as or longer than their mature achenes; ray achenes awnless or with
one awn 18.
17. Rays shorter than their mature achenes; ray achenes with 3 awns Sanuitalia.
18. Achenes of ray flowers awnless (in ours) 19.
18. Achenes of ray flowers with one awn Philactis.
19. Leaves petiolate Heliopsis.
19. Leaves sessile Zinnia.
20. Pappus of plumose awns or squamellae (10-45) Tridax.
20. Pappus (when present) not of plumose awns or squamellae 21.
21. Disc achenes completely enclosed in turgid pales 22.
21. Disc achenes often subtended by or enfolded in or surrounded by pales but not
completely enclosed, the pales not turgid 23.
22. Rays linear-oblong; disc corollas with definite tubes; style short, with flattened
branches Aldama.
22. Rays ovate to orbicular; disc corollas without definite tubes; style long, with
terete branches Sclerocarpus.
23. Mature disc achenes distinctly rostrate 24.
23. Mature disc achenes not distinctly rostrate 25.
24. Ray flowers fertile, the ligules small and inconspicuous Heterosperma.
24. Ray flowers sterile, the ligules conspicuous, often showy Cosmos.
25. Pales conspicuously accrescent in fruit, greatly surpassing and completely
including the disc achenes and sometimes the ray achenes 26.
25. Pales not conspicuously accrescent in fruit, although often surpassing the achenes
and sometimes embracing or enfolding them, but not completely including them.
27.
26. Disc flowers brown to black; pales setulose-pectinate; achenes with pappus of
about 10 upwardly serrate awns Rojasianthe.
26. Disc flowers white or yellow; pales not setulose-pectinate; achenes without
pappus Montanoa.
27. Ray achenes with deeply lobate wings, the lobes ascending, sometimes almost
setaceous Synedrella.
27. Ray achenes not winged, or if wings present, these not lobate 28.
28. Mature disc achenes often conspicuously tuberculate or rugose (in ours) 29.
28. Mature disc achenes usually relatively smooth or sulcate or striate, not usually
conspicuously tuberculate or rugose 31.
29. Heads radiate; pales setaceous, subtending the flowers Eclipta.
29. Heads discoid; pales membranaceous, embracing the flowers 30.
30. Phyllaries hispid, the hairs often more than 1 mm. long; pappus cyathiform, on
neck or short beak of the achene .... Eleutheranthera.
184 FIELDIANA: BOTANY, VOLUME 24
30. Phyllaries strigillose with very short hairs; pappus cyathiform, annular, the
achenes truncate at the apex Garcilassa.
31. Outer achenes cylindrical, 8-10-sulcate, crowned by an epigynous disc.
ChrysantheUum.
31. Outer achenes not as above 32.
32. Peduncles and phyllaries conspicuously stipitate-glandular; phyllaries linear-
oblanceolate to spathulate; achenes often incurved Sigesbeckia.
32. Peduncles and phyllaries not stipitate-glandular, or if so, then the phyllaries
ovate to broadly elliptic or oblanceolate; achenes not incurved 33.
33. Disc achenes with only one wing, or if 2 wings, one always much larger than the
other and adnate to or confluent with the longer awn 34.
33. Disc achenes not as above 35.
34. Squamellae entirely absent or very inconspicuous Notoptera.
34. Squamellae present, united into a more or less lacerate, paleaceous corona
adnate to the pappus awns Otopappus.
35. Leaf blades conspicuously asymmetric, oblique at the base (rounded on one side,
acute on the other) Goldmanella.
35. Leaf blades not conspicuously asymmetric 36.
36. Pappus awns or squamellae inserted on a neck or beak of the achene (this
distinct on mature achenes, but awns are sometimes soon deciduous) 37.
36. Pappus awns or squamellae not inserted on a neck or beak of the achene
(pappus sometimes wanting) 39.
37. Awns commonly 15-30 Perymenium.
37. Awns commonly 1-3 38.
38. Leaves never lobate nor hastate; plants often large, climbing shrubs with
ferruginous pubescence, or the leaves canescent-pilose beneath, or the leaves
decurrent on the petiole and clasping the stem, or the involucres distinctly
biseriate, the outer phyllaries foliaceous and longer than the inner,
membranous ones, or the heads in clusters of (l-)3-5, the involucres 2-3-seriate,
the outer foliaceous phyllaries longer than the inner ones Zexmenia.
38. Leaves sometimes lobate; plants not exhibiting any of the conditions listed
above Wedelia.
39. Plants minute, inconspicuous, 1-2 cm. high (in ours) 40.
39. Plants neither minute nor inconspicuous, at least 5 cm. high (most much higher).
41.
40. Heads discoid; phyllaries only 2 Cuchumatanea.
40. Heads radiate; phyllaries 5 or more Aphanactis.
41. Heads discoid 42.
41. Heads radiate 49.
42. Plants prostrate; disc flowers in anthesis bluish or purplish Trichospira.
42. Plants not prostrate, or if so, the disc flowers in anthesis yellow, orange, white,
or greenish, never bluish or purplish 43.
43. Disc achenes rather thick, subterete or 4-5-angulate 44.
43. Disc achenes more or less compressed 47.
44. Disc flowers yellow 45.
44. Disc flowers white or greenish white Melanthera.
45. Leaves often incised or dissected; phyllaries biseriate; pappus of 1-4 awns, these
usually retrorsely barbed Bidens.
45. Leaves simple or trilobate (but not otherwise incised or dissected); phyllaries
usually 3-4-seriate; pappus of 4-70 awns or squamellae 46.
NASH: FLORA OF GUATEMALA 185
46. Pappus of 4-20 broad-based, lacerate awns or squamellae; achenes 4-5-angulate,
usually pubescent or ciliate on the angles Calea.
46. Pappus commonly of 20-70 setiform awns; achenes not angulate but sometimes
obscurely costate, not pubescent or ciliate on the costae Neurolaena.
47. Leaves often lobate or pinnate, at the base often decurrent as wings on petioles
and/or stems; achenes winged on each margin or rarely on only one Verbesina.
47. Leaves neither lobate nor pinnate, seldom if ever decurrent at the base as wings
on petioles or stems; achenes not winged but usually ciliate 48.
48. Plants herbs; heads usually long-pedunculate Spilanthes.
48. Plants shrubs; heads pedicellate or sessile, disposed in cymose, often paniculate
inflorescences Salmea.
49. Plants scapose; ray flowers neutral lostephane.
49. Plants not scapose (sometimes plants may appear subscapose, the flowering heads
solitary on elongated peduncles, but then the ray flowers fertile 50.
50. Lower leaves broadly ovate to suborbicular and shallowly angulate-lobate, very
large, frequently as much as 30 cm. long and broad, tomentose on both
surfaces; plants trees or large shrubs, with conspicuous white ray flowers 8-15
mm. long Podachaenium.
50. Lower leaves not as above; plants herbs or shrubs, rarely trees but then the ray
flowers yellow, conspicuous or not, or if white, then small, often inconspicuous,
1-6 mm. long (some Calea and Verbesina species) 51.
51. Plants confined to sea beaches and dune areas; leaves carnose, oblanceolate to
spathulate, sessile or with a petioliform base; achenes crowned with a short,
dentate cup less than 0.5 mm. high Borrichia.
51. Plants not usually found on sea beaches or dune areas; leaves not usually carnose,
variously shaped; achenes not crowned with a dentate cup less than 0.5 mm. high
(sometimes a lacerate crown of squamellae present but this at least 0.5 mm.
high) 52.
52. Heads usually large, the rays mostly 2-6 cm. long 53.
52. Heads smaller, the rays mostly less than 2 cm. long 54.
53. Phyllaries biseriate, the outer ones spreading or strongly reflexed at anthesis,
somewhat carnose and appearing ceraceous; pales scarious; leaves penninerved;
rays variously colored; pappus of 2 minute awns or 2 slender, filamentous awns,
or wanting Dahlia.
53. Phyllaries commonly about 4-seriate (rarely 2-5-seriate), the outer ones spreading
or not, neither carnose nor ceraceous; pales rigid; leaves triplinerved; rays always
yellow or orange-yellow; pappus of 1-2 awns and 4-12 squamellae, or wanting.
Tithonia.
54. Ray flowers commonly neutral (rarely pistillate but then usually sterile) 55.
54. Ray flowers commonly pistillate and fertile (rarely sterile) 59.
55. Disc achenes usually conspicuously marginate or winged when mature.
Coreopsis.
55. Disc achenes neither conspicuously marginate nor winged 56.
56. Pappus composed of both awns and squamellae Viguiera.
56. Pappus wanting or composed of 1-3 awns, or of squamellae, but not of awns
and squamellae together 57.
57. Leaves often incised or dissected; pappus awns usually retrorsely barbed (rarely
smooth) Bidens.
57. Leaves neither incised nor dissected but sometimes trilobate or hastate (SimsiaY,
pappus awns (when present) never retrorsely barbed 58.
186 FIELDIANA: BOTANY, VOLUME 24
58. Achenes epappose or with a few short, inconspicuous, fimbriate squamellae.
Hymenostephium.
58. Achenes with 2 or more awns Simsia.
59. Plants decumbent, procumbent, or prostrate (rarely ascending) 60.
59. Plants commonly erect (rarely procumbent or decumbent in some species of
Spilanthes and Wedelia) 63.
60. Pappus or 2 stout, smooth, spinelike awns Calyptocarpus.
60. Pappus none or reduced to a minute, inconspicuous annulus, or composed of
few-many, fimbriate, awnlike squamellae 61.
61. Ray flowers inconspicuous, the ligules yellow to orange, but often becoming
whitish when dried, about 1.5 mm. long; pappus none or reduced to a minute
annulus Jaegeria.
61. Ray flowers small but usually conspicuous, the ligules white above, often pinkish
or purplish beneath, 4-10 mm. long; pappus none or composed of few-many,
fimbriate, awnlike squamellae 62.
62. Principal leaves sessile, the blades spathulate; ray flowers 4-5 mm. long; pappus
none Selloa.
62. Principal leaves short-petiolate, the blades ovate or rhombic-lanceolate to
linear-lanceolate or linear-elliptic; ray flowers 5-10 mm. long; pappus or few-
many, fimbriate, awnlike squamellae (rarely absent) Sabazia.
63. Leaves frequently but not always lobate or pinnate, their petioles sometimes
winged 64.
63. Leaves neither lobate nor pinnate, their petioles not usually winged, but if so,
tapering from the leaf base to a narrow margin, not a broad wing to the base of
the petiole 65.
64. Achenes usually broadly winged, with 1-2 awns, these appearing more or less
confluent with margins of the achenes (sterile ray achenes sometimes with 3
awns) Verbesina.
64. Achenes not winged, with numerous awns, these not confluent with margins of
the achenes Neurolaena.
65. Heads usually solitary on elongated peduncles Spilanthes.
65. Heads not usually solitary on elongated peduncles; inflorescences usually cymose
and frequently paniculate 66.
66. Pappus of numerous, slender awns, often easily deciduous 67.
66. Pappus not as above, sometimes consisting of awns or awnlike scales but these
broad-based and persistent, or pappus sometimes absent 69.
67. Disc paleaceous only on margin Alepidocline.
67. Disc paleaceous throughout 68.
68. Ligules of pistillate flowers 0.3-9 mm. long; pales mostly trilobate or irregularly
3-cleft Schistocarpha.
68. Ligules of pistillate flowers 10-15 mm. long (in ours); pales essentially entire,
unlobed (in ours) Oteiza.
69. Achenes usually winged, or if not winged, the plants procumbent or decumbent,
the leaves trilobate or hastate (some species of Wedelia) 70.
69. Achenes not winged, the plants commonly erect, the leaves neither trilobate nor
hastate 71.
70. Leaves never lobate nor hastate; plants often large, climbing shrubs with
ferrugineous pubescence, or the leaves canescent-pilose beneath, or the leaves
decurrent on petioles and clasping the stems, or involucres distinctly biseriate,
the outer phyllaries foliaceous and longer than the inner, membranous ones, or
NASH: FLORA OF GUATEMALA 187
heads in clusters of (l-)3-5, the involucres 2-3-seriate, the outer foliaceous
phyllaries longer than the inner ones Zexmenia.
70. Leaves sometimes lobate; plants not exhibiting any of the conditions listed
above Wedelia.
71. Disc achenes with pappus of several lacerate or fimbriate, awnlike scales but no
true awns 72.
71. Disc achenes with true awns or epappose 73.
72. Plants shrubs, trees, or perennial herbs; pales concave, rigid or thin, at least
partially enfolding the achenes Calea.
72. Plants annual herbs; pales narrow, thin, subtending but not enfolding the
achenes Galinsoga.
73. Outer phyllaries spreading; achenes epappose Rumfordia.
73. Outer phyllaries erect; achenes with 2 awns and small squamellae Lasianthaea.
ACANTHOSPERMUM Schrank
Reference: S. F. Blake, Revision of the genus Acanthospermum,
Contr. U. S. Nat. Herb. 20: 383-392, t. 23. 1921.
Pubescent annuals, dichotomously branched; leaves opposite, the margins
subentire to pinnatifid; heads radiate, small, sessile or short-pedunculate, solitary in
the leaf axils and in the forks of the branches; phyllaries in two series, those of the
outer, involucral series herbaceous, elliptic to ovate, those of the inner series as many
as the achenes and closely enveloping them, greatly enlarged in fruit; receptacle
small, convex; pales membranous, subtending the disc flowers, somewhat persistent;
ray flowers 5-8, fertile, uniseriate, conspicuous or not, the ligules pale yellow, elliptic
to ovate, emarginate or tridenticulate; disc flowers 5-30, hermaphrodite, sterile, the
corollas yellow; anthers sagittate at the base, the apical appendages ovate, obtuse;
styles of hermaphrodite flowers clavate, obtuse, hispidulous; fruits obtriangular or
deltoid to oblong-fusiform, rarely trigonous-turbinate, weakly or strongly compressed
laterally, echinate on the entire surface, the angles or rarely only the apex with
straight or uncinate spinules, those at the apex of the fruit usually elongated; pappus
none.
Eight species, in tropical America, with only one in Guatemala.
Acanthospermum hispidum DC. in DC. Prodr. 5: 522. 1836.
Figure 48.
In weedy fields or waste ground around dwellings, about 850
m.; Jutiapa. Introduced as a weed in southern United States;
Honduras; El Salvador; Costa Rica; Nicaragua; West Indies; South
America; naturalized in Africa and the Hawaiian Islands.
Usually erect, bushy herbs about 60 cm. high, the stems densely hispid-pilose
with spreading hairs; leaves sessile, the blades elliptic to ovate or triangular-ovate,
mostly 2-12 cm. long, acute or obtuse, mucronate, gradually cuneate below the
middle to the sessile base, the margins serrulate to entire, hispid- pilose on both
surfaces, gland-dotted beneath; heads in an thesis 4-5 mm. broad, in fruit 13-18 mm.
broad, on peduncles 3-15 mm. long; outer phyllaries 5, ovate, subacute, 3.5-4 mm.
long; ray flowers 5-8, their corollas pale yellow, about 1.5 mm. long; disc flowers
188 FIELDIANA: BOTANY, VOLUME 24
about 7, the corollas shortly hispid-pilose, about 1.7 mm. long; pales stipitate-
glandular dorsally, lacerate- cilia te at the truncate apex; fruits deltoid, strongly
compressed, gland-dotted, rather densely uncinate- hispid all over the body, 4-5 mm.
long, the 2 terminal spinules straight or curved, strongly divergent, 3-4 mm. long.
Plants of this genus are seldom abundant in the few localities
in Central America where they have been found. It is quite possible
that they are not native, but introduced from South America.
ALDAMA La Llave & Lexarza
Reference: Charles Feddema, Re-establishment of the genus
Aldama (Compositae-Heliantheae), Phytologia 21: 308-314. 1971.
Erect, branching annuals, the stems strigose-hispidulous or glabrate; lower leaves
opposite, the upper ones alternate, short-petiolate, the blades lanceolate or linear-
lanceolate, the margins entire or minutely denticulate; heads numerous, radiate,
often long-pedunculate; receptacles convex to conical, paleaceous; involucres
campanulate; phyllaries biseriate, subfoliaceous, subequal or sometimes the outer
ones considerably shorter, appressed; pales enclosing the achenes and deciduous with
them; ray flowers neutral, sterile, with short tubes and linear-oblong to broadly
elliptical ligules; disc flowers perfect, fertile, their short tubes 10-nerved, the limb 5-
lobate; style short, the stigmatic branches flattened; anthers exserted at anthesis,
sagittate at the base, their apical appendages lanceolate; achenes black, each
enclosed in a wrinkled or pitted pale; pappus a low crown of setaceous squamellae or
sometimes reduced to a low ridge, or wanting.
A single species is known.
Aldama dent at a La Llave & Lex. Nov. Veg. Descr. 1: 14. 1824.
Gymnopsis dentata DC. Prodr. 5: 561. 1836. G. schiedena DC. I.e.
Sclerocarpus dentatus Benth. & Hook, ex Hemsl. Biol. Centr. Am.
Bot. 2: 164. 1881. S. schiedeanus Benth. & Hook, ex Hemsl. I.e. S.
kerberi Fourn. Bull. Bot. Soc. Fr. 20: 183. 1883. S. schiedeanus var.
elongatus Greenm. Proc. Amer. Acad. 32: 309. 1897. G. acuminata
Blake ex Robinson, op. cit.: 49. 505. 1913. S. elongatus Greenm. &
Thompson, Ann. Mo. Bot. Gard. 1: 412. 1915. Figure 49.
Damp or dry thickets, on slopes or plains, 200-1,360 m.;
Chiquimula; Jalapa; Jutiapa; Santa Rosa; Zacapa. Mexico; British
Honduras; Honduras; northern Venezuela.
Erect, slender, branching annuals, mostly 0.5-1.5 m. high, the stems subterete,
more or less strigose-hispidulous or glabrate, often tinged with red or purple; leaves
short-petiolate, the uppermost alternate, the middle and lower ones opposite, the
blades linear-lanceolate to ovate-lanceolate, mostly 3-12 cm. long, commonly
acuminate, rarely obtuse, cuneate to almost rounded at the base, hispid or substrigose
above, sometimes scabrous, strigose-hispidulous to hirsute beneath, the margins entire
or remotely denticulate; heads numerous, long-pedunculate, disposed in very lax
panicles, the peduncles mostly 5-12 cm. long; phyllaries biseriate, 5-9 mm. long, the
NASH: FLORA OF GUATEMALA 189
outer ones ovate to elliptic-lanceolate, strigose or hispidulous, ciliate, acute to obtuse,
the inner ones usually longer and broader; ray flowers 5-11, the ligules bright yellow,
linear-oblong to broadly elliptic, mostly 8-13 mm. long; disc flowers 20-70, the
corollas yellow, 3-4 mm. long, sparsely puberulent or glabrate; pales 5-8 mm. long,
tubular, more or less compressed and wrinkled at the base, enclosing the black
achenes; achenes 2-3 mm. long, glabrous, more or less narrowly obovoid; pappus of
minute scales or absent.
ALEPIDOCLINE Blake
Reference'. H. Robinson and R. D. Brettell, Tribal revisions in
the Asteraceae. IV, The relationships of Neurolaena, Schistocarpha
and Alepidocline, Phytologia 25: 439-445. 1973.
Erect, branched annuals, pubescent and sparsely glandular; leaves opposite,
petiolate, the blades thin, ovate, triplinerved, the margins serrate; inflorescences
cymose; heads pedicellate, heterogamous, radiate; involucres ovoid or hemispheric;
phyllaries multiseriate, broadly oblong or oval or the innermost ones lance-oblong,
greenish, obtuse or the inner ones acute; receptacle convex to conic, hirsutulous, with
a few linear pales near the margin; ray flowers uniseriate, pistillate, the corolla tube
slender, the ligule small, spreading, tridentate, first white, then pink or reddish purple
in age; disc flowers numerous, hermaphrodite, fertile, the corollas tubular, the limb
shorter than the tube, shortly 5-dentate, yellow but sometimes in age the limb
becoming purplish; anthers blackish, obtusely sagittate at the base, with an obtuse
apical appendage; style branches linear, minutely hispidulous at the apex; achenes
obovoid, obcompressed, plano-convex, black, lustrous, glabrous; pappus caducous, the
8-10 bristles uniseriate, subequal, hispidulous.
The genus consists of a single species.
Alepidocline annua Blake, Journ. Wash. Acad. Sci. 24: 441, f.
2. 1934. Figure 50.
Most abundant as a weed in cornfields and along roadsides,
also in dense or open, damp or wet thickets and in oak or coniferous
or mixed forest, frequently on limestone, 1,800-3,800 m.; Chimal-
tenango (type from Chichavac, Skutch 722); Huehuetenango;
Jalapa; Quezaltenango; Sacatepequez; San Marcos; Solola;
Totonicapan.
Erect annuals (10) 20-60 cm. high, often much branched, the stems sparsely
pilose with spreading hairs, glandular-pilose in the inflorescences; leaves petiolate, the
upper ones short-petiolate, the blades broadly ovate to oblong-lanceolate, the
principal ones mostly 3-7.5 cm. long, acuminate, cuneate at the base or almost
rounded and then rather abruptly cuneate, the margins serrate, sparsely pilose on
both surfaces, usually more densely so beneath, the hairs multiseptate; heads on
pedicels mostly 1-5 cm. long; involucres 5-6 mm. high, 5-8 mm. broad; phyllaries 5-6-
seriate, the outermost ones short, elliptic or oblong, about 2.5 mm. long, the others
oval to oblong, stria te, glabrous; ray flowers 10-17, the ligules first white, then pink,
purplish red or dull red in age, about 1.5 mm. long; disc flowers numerous, the
190 FIELDIANA: BOTANY, VOLUME 24
corollas yellow or sometimes purplish in age, 3-3.5 mm. long; achenes about 1.5 mm.
long, finely and obscurely striate; pappus bristles 1.5-2 mm. long.
Although one of the most common cornfield weeds in the
mountains of some regions of Guatemala (where the plants often
remain green until killed by severe frost), these plants have not
been found in any other country.
AMBROSIA Linneaus Ragweed.
References: P. A. Rydberg, N. Amer. Fl. 33: 15-22. 1922; W. W.
Payne, A re-evaluation of the Genus Ambrosia (Compositae),
Journ. Am. Arb. 45: 401-438. 1964.
Annual or perennial, monoecious herbs or subshrubs, glandular-aromatic, with
usually rough pubescence; leaves opposite or alternate, petiolate or sessile, mostly
lobate or dissected; staminate heads pedicellate or sessile, disposed in a racemose or
spicate inflorescence, the pistillate heads borne in leaf axils at the base of the
inflorescence; involucres funnel-shaped, urn-shaped, or saucer-shaped, commonly 5-
12-lobate or -dentate (tips of fused phyllaries); receptacle paleaceous, the pales
usually filiform, often with dilated tips; corollas hyaline, campanulate, 5-4 -cleft;
stamens 5, the anthers distinct or scarcely united, the apices appendaged; pistillate
heads erect, the involucres surrounding the nutlike fruit obovoid or urn-shaped, with
one or more series of tubercles or spines and usually with a truncate, 3-4-dentate
beak; pistil solitary, without a corolla, the style branches elongated, linear, the inner
stigmatic surface papillose; staminate heads more or less nutant, the involucres
saucer-shaped, the style short, truncate, its circular terminus penicillate with
unicellular hairs.
Perhaps 30, often closely related species, most of them in
temperate and tropical America, with one or two in warmer regions
of the Old World. Only the following two are known in Central
America. In the United States, species of this genus are abundant
weeds in many regions and are often the cause of hay fever; plants
of other families produce the same affliction but never on so vast a
scale.
Although the plants are characteristic of some alpine meadows,
the type was collected in an area that certainly is not alpine. The
plants are very small, and growing among mosses, grasses, and other
plants, are not easily seen or distinguished.
Plants usually erect (rarely decumbent); leaves pinnate-pinna tisect, deeply pinna tif id
but not divided to the costa, the segments relatively large; found mostly in the
interior, not growing on seashores A. cumanensis.
Plants prostrate or procumbent; leaves bipinnatifid to tripinnatifid, divided to the
costa into very small and numerous segments, although the ultimate segments
not usually divided to the costa; growing in sand on or near sea beaches.
A. hispida.
NASH: FLORA OF GUATEMALA 191
Ambrosia cumanensis HBK. Nov. Gen. & Sp. 4: 276. 1820.
Cakiax, tus pirn (Quecchi, Alta Verapaz). Figure 51.
Wet or dry, sometimes stony fields, dry grassy slopes, or a weed
in cultivated ground, 800-2,000 m.; Alta Verapaz; Baja Verapaz;
Huehuetenango; Jutiapa; El Quiche. Mexico; British Honduras;
Honduras; El Salvador; Nicaragua; Panama; West Indies; South
America.
Erect or rarely decumbent perennials, usually less than 1 m. tall, simple or
branched, the stems white-hirsute with mostly long, spreading hairs; leaves on short
or long petioles, the blades deeply pinnatifid but not divided to the costa, triangular-
ovate in outline, mostly 3-10 cm. long, densely strigose, paler beneath, hirsute on the
veins, the segments large and broad, obtuse or acute, variously lobate or dentate;
staminate heads numerous, sessile or on pedicels to 2 mm. long, disposed in long,
dense, spikelike racemes; involucres saucer-shaped, 3-4 mm. broad, shallowly crenate
or lobate, hispidulous, commonly 1 5- 30-f lowered; pales filiform; corollas greenish
yellow, more or less puberulent; pistillate heads fasciculate in the upper leaf axils;
fruit obovoid, angulate, puberulent and glandular-atomiferous, the short beak about
0.5 mm. long, the 4-7 spines short, stout, conic.
The plants have a strong, rank odor like that of the common
ragweed (Ambrosia artemisiifolia L.) of the United States. They
flower only during the rainy season.
Ambrosia hispida Pursh, Fl. Amer. Sept. 743. 1814.
In sand on or near sea beaches; British Honduras. Florida;
Yucatan; Panama; West Indies; northern coast of South America.
Herbaceous or somewhat suffrutescent perennials, the stems prostrate or
procumbent, mostly 20-50 cm. long, branching, densely hirsute or pilose with white
hairs; leaves petiolate, the blades bipinnatifid or more deeply divided, densely
grayish-pubescent on both surfaces, triangular-ovate in outline, mostly 3-5 cm. long,
the ultimate segments very small, the rachis not or very inconspicuously winged;
staminate heads on pedicels 1-2 mm. long, disposed in very dense, terminal, spikelike
racemes; involucres about 3 mm. broad, saucer-shaped or hemispheric, softly
pubescent, commonly 9-15-flowered; pales filiform, slightly thickened and puberulent
at the apex; corollas more or less puberulent; pistillate heads in fascicles of 3-5 in the
upper leaf axils; body of the fruit about 2 mm. long, ovoid, densely pubescent when
young, the beak about 0.5 mm. long, the spines very short and small.
Called "margarita del mar" and "malvarosa de la playa" in
Yucatan, where the plants are employed in domestic medicine.
APHANACTIS Weddell
Very small herbs, annual or perennial, usually cespitose or prostrate and repent,
glabrous or pubescent; leaves opposite, the margins entire or obscurely denticulate, 3-
5-nerved; heads small, heterogamous, radiate, few or numerous, sessile or pedunculate
at the ends of branches or among the leaves of the basal rosette; involucres
campanulate; phyllaries biseriate, subequal or graduated, thin, herbaceous, oblong or
192 FIELDIANA: BOTANY, VOLUME 24
ovate, obtuse or subacute; receptacle convex or becoming conic; pales thin, rigid; ray
flowers pistillate, uniseriate, the ligules yellow, suberect or spreading, 2-3-dentate at
the apex; disc flowers hermaphrodite, fertile, the corollas regular, tubular, 5-dentate;
anthers entire at the base; style branches of the disc flowers truncate or with short
appendages; achenes oblong or cuneate, obtusely tetragonous or somewhat
compressed, striolate; pappus none.
Three species, one in Guatemala, the others in the Andes of
Ecuador and Peru.
Aphanactis standleyi Steyerm. Field Mus. Bot. 22: 390, f. 3.
1940. Figure 52.
Damp or wet, open meadows, most common in alpine
situations, 2,700-3,700 m.; Chimaltenango (type from Cerro de
Tecpam, region of Santa Elena, Standley 58674); Huehuetenango
(Sierra de los Cuchumatanes); San Marcos; Solola; Totonicapan
(Sierra Madre Mountains).
Small, acaulescent annuals, more or less cespitose; leaves opposite, sessile,
decussate, flat on the ground, forming small rosettes, broadly ovate or almost
rounded, mostly 6-20 mm. long and 5-18 mm. wide, obtuse or almost rounded at the
apex, 3-5-nerved, the margins entire or obscurely denticulate and more or less ciliate,
green above and villous, almost glabrous and whitish beneath; heads 3-5 in the center
of each rosette of leaves, sessile, crowded, 3-4 mm. high, 2-3 mm. broad; principal
phyllaries about 6, biseriate, subequal, ovate or broadly oblanceolate, acute, glabrous
or nearly so, the outer ones ciliolate, scarious-marginate, one of the outer phyllaries
larger, broadly ovate; receptacle conic; pales variable in size and shape, 1.5-2.5 mm.
long, acuminate or aristate, the inner ones sparsely serrate or densely ciliate; ray
flowers commonly 5, the ligules yellow, about 1 mm. long; disc flowers 8-10, the
corollas yellow, about 1.5 mm. long, the tube villous at the base; achenes 1-1.3 mm.
long, turgid, obscurely 4-5-angulate, glabrous, blackish, truncate at the apex.
BALTIMORA Linneaus
Reference: Tod F. Stuessy, Revision of the genus Baltimora
(Compositae, Heliantheae), Fieldiana: Botany 36: 31-49. 1973.
Erect, much branched annuals with rough pubescence; leaves opposite, petiolate,
the blades thin, broad, the margins crenate-serrate or biserrate, heads heterogamous,
radiate, disposed in few-flowered racemes or broad, open panicles; involucres ovoid or
campanulate; phyllaries few, 3-seriate, more or less scarious toward the base, usually
herbaceous at the apex; receptacle convex; pales conduplicate, lanceolate; ray
flowers uniseriate, pistillate, fertile, the ligules yellow, spreading, the style with
stigmatic lobes about 1 mm. long; disc flowers hermaphrodite, sterile, the corollas
yellow, the 5 lobes partially reflexed; anthers black, auriculate at the base; style
undivided; ray achenes thick, triquetrous, truncate at the apex, sometimes winged;
pappus a crown of minute scales.
The genus consists of two species, with only one in Central
America.
MELCHERT: FLORA OF GUATEMALA 193
Baltimora recta L. Mant. PL 288. 1771. B. alba Pers. Syn. 2:
489. 1807. Scolospermum baltimoroides Less. Linnaea 5: t. 2, f. 19-
31. 1830. Fougerouxia alba DC. Prodr. 5: 510. 1836. F. recta DC. I.e.
Wedelia populifolia Hook. & Arn. Bot. Beechey Voy. 435. 1841.
Figure 53.
Damp thickets, open fields, grassy slopes, rocky hills, sometimes
a weed in waste or cultivated ground, sea level to 1,200 m.; Alta
Verapaz; Chimaltenango; Chiquimula; Escuintla; Guatemala; Pe-
ten; Retalhuleu; San Marcos; Santa Rosa; Zacapa. Mexico;
Honduras and El Salvador to Panama.
Much branched herbs, commonly about 1 m. high, rarely as much as 3 m., the
branches slender, angulate, hispidulous or pilose; leaves on petioles mostly 1-7 cm.
long, the blades broadly ovate to lance-ovate, mostly 5-10 cm. long, long- acuminate,
truncate at the base or broadly cuneate, the margins crenate-serrate, both surfaces
more or less strigose and scabrous; heads on slender, strigose pedicels, disposed in
racemes or corymbiform panicles; involucres campanulate, commonly about 6 mm.
broad; phyllaries ciliate near the apex, the outer ones ovate, subfoliaceous, densely or
weakly strigose, the inner ones lanceolate, rigid; ray flowers 3-8 (commonly 5), the
ligules 4-5 mm. long; disc flowers 16 or more; pales 3.5-4 mm. long; achenes about 3
mm. long, truncate at the apex, puberulent; pappus reduced to a small crown or cup
of minute scales.
One of the common, weedy plants of Central American
lowlands, often abundant in cornfields.
BIDENS Linneaus
By THOMAS E. MELCHERT
Erect annuals, erect to sprawling or procumbent, herbaceous perennials, or
scandent vines and shrubs, the stems terete, angulate or tetragonal, commonly striate
or multisulcate; leaves opposite, entire, 3-7-partite, or variously bipinnatisect (several
of these forms frequently segregating within single populations), ours mostly petiolate
and serrate, the margins narrowly hyaline, setose, the surface glabrous to densely
pilose, the pubescence exceedingly variable within given species; heads mostly radiate,
showy, less frequently discoid or disciform, ours conspicuously pedunculate, solitary
to clustered, corymbose- or cymose-paniculate; involucre dimorphic, biseriate, outer
phyllaries green, herbaceous, minute to subfoliose, linear to oblanceolate or even
ovate, the surface commonly dark-lined, margins entire, glabrous or ciliate, the inner
phyllaries paler, membranous, striate, becoming chaffy, the margins hyaline, clear or
yellow; ray florets commonly neutral, occasionally styliferous, ligules 3-16, normally
yellow to white, rarely purple-lavender, when yellow the proximal portion usually
deep, golden-yellow, the distal portion paler, often pressing whitish yellow, the
transition between these zones rather abrupt, some pale yellow throughout, white
ligules usually with faint yellow chalcone pigment along the major longitudinal veins,
anthocyanin spots may be present at the ligule base, apex rounded to subtruncate,
mostly 3-denticulate; disc florets few to numerous, the corollas 5-toothed, yellow (one
rosaceous) above, the tube becoming whitish below, the veins sometimes reddish;
194 FIELDIANA: BOTANY, VOLUME 24
anthers brown to shiny black, their terminal appendages ovate-lanceolate; style
branches tipped with short-acute to elongate-attenuate appendages; palae usually
similar to, but narrower than, the inner involucre, becoming linear-lanceolate toward
the center of the head, chaffy with yellowish, reddish or dark green striations, some
reddish or black-tipped; achenes black or olivaceous at maturity, the surface stippled,
glabrous or antrorsely setose, linear to linear-attenuate or linear- cuneate, in cross-
section basically tetragonal, each face longitudinally 3-ribbed, however, frequently
obcompressed, achenes then trigonal with a convex dorsal surface, some becoming
flattish, each face then seemingly 5-ribbed with a long, more or less raised, central
midrib, more rarely clavate or completely flattened, wing margined and rostrate, all
achenes in one head essentially equal-sized (monomorphous), erect or supernally
recurved, or, more commonly, graded monomorphous, i.e., tending toward dim-
orphous, the central achenes longest, erect, narrowly linear, becoming gradually
shorter, more incurved and obcompressed toward the periphery, the outermost about
one-half to three-fifths the size of the inner, sometimes truly dimorphous, 5-9 of the
peripheral achenes more or less clavate, 2-4 mm. long, rubrocastaneous or yellowish,
frequently exaristate, or several of the peripheral achenes densely erect-tomentose,
mostly three fifths to three-fourths the size of the central ones, all succeeding whorls
black, these graded monomorphous, margins occasionally tuberculate or tuberculate-
alate, apically of full width or attenuate, rarely rostrate, commonly aristate; awns 2-4
(-7), retrorsely or antrorsely barbed, rarely smooth, all erect or some divergent to
reflexed, sometimes exaristate.
A complex, worldwide genus with about 75 taxa in Mexico and
Central America, 20 of which (18 species and two varieties) occur in
Guatemala. The majority of these are weedy species.
Perennials.
Rays yellow; stems terete or angled.
Rays pistillate, 5 or 6 (8), yellow, large, oval; achenes obcompressed, linear-
cuneate, pallid tan to olivaceous at maturity (never black), glabrous,
longitudinally multinerved; sprawling subshrubs with coarsely dentate,
usually trifoliate leaves B. ostruthioides.
Rays neutral, 2-12; achenes linear-tetragonal to narrowly cuneate, becoming
blackish at maturity, commonly setose, each face 3-nerved, or superficially
5-nerved if strongly obcompressed.
Scandent vines or shrubs; achenes linear, often flattish, their margins white-
ciliate throughout, these commonly in groups of 2-4 arising from single
tuberculae.
Heads radiate or discoid, usually small, short-peduncled, congested; radiate
heads 1.5-3 (rarely -4) cm. across; ligules (0-) 2-5, 7-15 (-20) mm. long,
(2.5-) 3-5 mm. wide; discs 7-10 (-12) mm. wide, at anthesis 8-11 mm.
high, in fruit 10-12 mm. high; achenes 6-9 (-10) mm. long, about 0.8
mm. wide; awns 2-4 (-5) mm. long, smooth or barbed B. squarrosa.
Heads radiate, large, (4-) 5-6 (-7) cm. across, long-peduncled, in open, few-
flowered clusters; ligules mostly 2-2.5 (-3) cm. long, 5-8.5 mm. wide;
discs 12-16 mm. wide, at anthesis 12-16 mm. high, in fruit 20-22 mm.
high; achenes 12-18 mm. long, 1 mm. wide; awns 2 (3 or 4), 4-6.5 (-10)
mm. long; retrorsely barbed B. holwayi.
MELCHERT: FLORA OF GUATEMALA 195
Erect or sprawling to procumbent herbs, their bases perhaps slightly ligneous;
achenes (or ovaries) glabrous or erect- setose, never marginally tuberculate
or white-cilia te.
Plants procumbent or sprawling with only one or a few long-peduncled
heads on each main branch (reportedly rarely ascending to erect);
achenes narrowly linear- tetragonal, gradually narrowed above; mesic,
high-mountain species.
Leaves trifoliate, 4-9 (-13) cm. long; heads relatively small, only 1-2 cm.
across including the rays, these inconspicuous, pale yellow with
brownish veins, narrowly linear-elliptic, only 8-15 mm. long, to 5 mm.
wide; outer involucral bracts narrowly linear, (6-) 9-14 mm. long,
spreading or recurved at maturity, often equalling the rays; achenes
monomorphic, some or all supernally recurved at maturity.
B. chiapensis.
Leaves (in ours) commonly bipinnatisect, rarely trifoliate, 1.2-5 (-7) cm.
long; heads large, to 5 (-6) cm. across, rays showy, two-toned-yellow,
golden-yellow proximally, distally paler and narrowed, (12-) 18-25
mm. long, to 13 mm. wide; outer bracts appressed, 3.8-7 mm. long;
inner bracts usually bearing conspicuous, elongate, multicellular hairs;
achenes somewhat dimorphous, several at the periphery yellow or
rubrocastaneous, 4-5 mm. long, subclavate, the inner erect, linear-
tetragonal B. triplinervia.
Plants erect herbs, typically of wet to subaquatic habitats, lower stems or
rhizome with dense clusters of thickened, elongate, pale, adventitious
roots; achenes broadest at or near the summit, cuneate to linear-
cuneate, at least the outer ones flattened.
Stems tetragonal; leaves winged-petiolate, variably shaped, now lanceo-
late, now 3-7-partite, now bipinnatisect; heads clustered; rays 5 (8);
outer phyllaries (5-) 9-13 (-16), 3-4 mm. long, narrowly linear, 0.5-0.9
mm. wide; achenes linear-cuneate B. aurea.
Stems terete; leaves broadly lanceolate, sessile, their bases connate; heads
hemispheric; outer phyllaries 5 (8), usually foliose; achenes cuneate.
B. laevis.
Rays white; stems tetragonal to multi-angled.
Heads small; rays 5, short, to 13 mm. long; achenes exaristate, uniformly linear-
clavate, 3-4.5 mm. long; leaves 1-2-pinnatifid with narrowly linear segments,
rare B. steyermarkii.
Heads rather large; rays 5-8, 12-25 mm. long; achenes aristate, linear-attenuate-
tetragonal or obcompressed and linear-cuneate; leaves highly variable, many
unlobed or 2-5-partite, their blades lanceolate to ovate.
Plants erect, wet habitat herbs; rhizomes with clustered adventitious roots;
heads numerous, clustered; achenes monomorphous, obcompressed, linear-
cuneate; outer phyllaries narrowly linear, mostly 9-13; inner phyllaries
ovate-triangular with broad, yellow-scarious margins B. aurea.
Plants procumbent to ascending (rarely erect); 1-3 long-peduncled heads
terminating each main branch; achenes dimorphous, the inner linear-
tetragonal, narrowed above; involucre usually with scattered to dense
multicellular hairs; outer phyllaries linear to linear-spathulate; inner
phyllaries frequently becoming blackish when pressed.
B. chrysanthemifolia.
196 FIELDIANA: BOTANY, VOLUME 24
Annuals.
Plants inconspicuous; rays tiny, purple-lavender; leaves linear-filiform, unlobed,
entire; achenes rostrate; awns antrorsely barbellate B. rostrata.
Plants conspicuous; rays white or yellow, may be rose-tinged or spotted, or heads
discoid; leaves not as above; achenes beakless or if rostrate the fertile portion
lacking cartilaginous-winged margins.
Achenes with 1 erect and about 3 divergent to reflexed awns; heads tiny; rays
pale yellow with brownish lines, linear-oblanceolate, 4-6 mm. long (often
becoming white when dried); outer phyllaries spreading, often equalling the
rays B. riparia.
Achene awns all erect or ascending; heads if tiny, with white, obovate or oval
rays; outer phyllaries appressed.
Rays yellow; stems terete or obscurely angled.
Rays normally with a reddish anthocyanin spot at the base of each large,
two-toned-yellow ligule; awns retrorsely barbed; achenes dimorphous,
the blackish inner ones linear-tetragonal, very gradually narrowed
above; margins wingless B. bicolor.
Rays never red-spotted; awns antrorsely barbellate; achenes monomorphous,
all flat, rostrate, margins with stramineous, tuberculate wings.
Leaf segments linear- filiform, glabrate; rays 5 or 6; outer phyllaries 7 or 8.
B. alata.
Leaf segments linear, minutely hispid; rays 10-12; outer phyllaries 8-12.
B. blakei.
Rays white, some rose-tinged; stems tetragonal, often sharply so.
Achenes dimorphous, the outer club-shaped, rubrocastaneous to yellowish
red; short-rayed (occasionally rayless) B. bigelovii var. angustiloba.
Achenes graded monomorphous, i.e., the outer gradually somewhat shorter
and sometimes slightly more obcompressed and incurved than the inner.
Achenes 2-3 or 3-5 awned in single heads; heads discoid, disciform or in
Central America, short-radiate.
Achenes 3-5-awned, ray florets usually lacking or minute and disciform
(2-3 mm. long); California, Mexico, and Central America; n = 36.
B. pilosa var. pilosa.
Achenes 2- or 3-awned; heads invariable radiate, rays 5-8 mm. long;
Central America B. pilosa var. minor.
Achenes 0-2-awned; heads invariably radiate.
Achenes 2-awned, outer phyllaries 8-13, broadly spathulate, often
apically extrorse, 2-4 mm. long, 0.6-1.3 (averaging 0.8) mm. wide;
leaflets simple, oblong-lanceolate; rays 5-8; tropical mountains, n =
24 B. alba var. radiata.
Achenes 0-2 awned; outer phyllaries 6-12, commonly 8, linear or linear -
subspathulate, 1.2-5 mm. long, 0.3-0.7 mm. (averaging 0.4 mm.)
wide; leaflets simple or highly dissected; montane; n = 12.
Ligules 8-18 (averaging 13) mm. long, 6-12 mm. wide, outer phyllaries
6-10, 3-5 (averaging 3.7) mm. long, 0.5-1 mm. wide.
B. odorata var. odorata.
Ligules 4-8 (averaging 6.3) mm. long, 3-6 mm. wide; outer phyllaries 7-
11, 1.2-3 (averaging 2) mm. long, 0.2-0.5 mm. wide.
B. odorata var. calcicola.
MELCHERT: FLORA OF GUATEMALA 197
Bidens alata Melchert, Phytologia 32: 292. 1975. Cosmos
steyermarkii Sherff, Field Mus. Bot. 22: 438. 1941, not B.
steyermarkii Sherff, 1944.
Known only from the type, Chiquimula: 1,200-1,500 m.,
Montana Castilla, vicinity of Montana Cebollas, along Rio Lucia
Saso, 3 miles S.E. of Quezatepeque, dry rocky slope of glade, Nov. 6,
1939, Steyermark 31341.
Slender, erect, sparingly branched herbs, 3-5 dm. tall, seemingly annual, the
stems terete, obviously multinerved and sulcate, mostly glabrous but with scattered
patches of spreading hairs, the branches narrowing above into elongate peduncles,
these delicate above, to about 18 cm. long, microcephalous, linear-bracteate below;
leaves few, 1-2-pinnatifid with narrowly linear segments, these 0.5-1 (-1.3) mm. wide,
sharply apiculate, the terminal ones to 2.5 cm. long, lateral segments in 1 or 2 pairs;
heads radiate, yellow, 2-3.5 cm. across; involucre strongly biseriate; outer phyllaries
linear-spathulate, 3-5.5 mm. long, ciliate, usually with a broad amber to blackish
central line; inner phyllaries striate, chestnut-brown with hyaline margins, oblong to
oblanceolate-oblong, glabrous or apically pubescent, 6-10 mm. long, 1.8-2.5 mm. wide;
ray florets 5 or 6, golden-yellow, distally somewhat lighter, probably two-toned, the
limb linear-oblong, to 1.5 cm. long, apically 2- or 3-denticulate or weakly incised; disc-
florets yellow, about 5 mm. long; achenes all completely flattened, 15-26 mm. long, to
3 cm. wide, the fertile portion brownish black, tapering above into a broad, flat,
yellowish, biaristate rostrum, margins stramineous, thickened and indurated, more or
less tuberculate-alate, antrorsely setose, the tuberculae more or less separate below,
becoming continuous above, ventral midnerve also more or less tubercu late-setose,
peripheral achenes dorsally pubescent; awns 2, antrorsely barbed, to 3 mm. long.
Bidens alba var. radiata (Sch.-Bip.) Ballard in Webb & Berth.
Phyt. Canar. 2: 242. 1844. Coreopsis leucanthema L. Cent. I. 29
(excl. syn. Tournef.) 1755. C. coronata L. Sp. PI. ed. 2: 1281. 1763
(ex syn. Vaill. & Plum., but not to descr. or type locality). C.
leucantha L. op. cit.: 1282. Kerneria tetragona Moench, Meth. 595.
1794 (ex descr.). Bidens leucantha Willd. Sp. PI. 3: 1719. 1804.
Kerneria leucantha Cass. Diet. Sci. Nat. 24: 398. 1822. Bidens
abortiva Schum. & Thonn. Beskr. Guin. PL 381. 1827. (ex descr.). B.
adhaerescens Veil. Fl. Flum. 348, pro parte. 1827. B. striata Sweet,
Brit. Fl. Gard. pi. 237. 1828. B. oxyodonta DC. Prodr. 5: 600. 1836.
B. leucantha Meyen & Walp.; Walp. Nova Acta Acad. Leop.-Carol.
19 (Suppl. 1): 271. 1843. B. pilosa f. radiata Sch. Bip. Flora 27: 673.
1844. (ex syn. B. leucantha Willd., excl. Krauss' spec.) B. pilosa var.
radiata Sch. Bip. in Webb & Berth. Phyt. Canar. 2: 242. 1844. B.
pilosa var. leucantha Harv.; Harv. & Sound. Fl. Cap. 3: 133. 1864.
Acocotli quauhuahuacensis Hernandez; Altam. Mat. Med. Mex. 2:
154. 1898. B. pilosa var. humilis Walp.; Reiche, Anal. Univ. Chile
112: 153. 1903.
198 FIELDIANA: BOTANY, VOLUME 24
Like B. odorata, B. alba var. radiata is a weed of roadsides,
cultivated fields, around towns, etc., but normally occurring in
moist subtropical to tropical areas between 240-600 m.; a few
collections along the highway between Antigua and Guatemala City
at about 2,000 m.; Alta Verapaz; Chiquimula; Guatemala; Izabal;
Peten; El Quiche; Retalhuleu; Suchitepequez; Zacapa. In moist,
tropical lowlands and lush adjacent mountains along the eastern
coast of Mexico from southernmost Tamaulipas, southward into
Guatemala and Honduras, in tropical Florida and sporadically in
the Caribbean and Central America.
Annuals; stems square, erect, 0.5-1.5 m. tall, sparsely to moderately pilose; leaves
petiolate 1-pinnately compound, 6-17 cm. long, 6-13 cm. wide, leaflets oblong-
lanceolate, sparsely pilose on both surfaces, not coriaceous; heads 6-18, cymosely
disposed, 2.2-4.2 cm. wide at an thesis, involucre sparsely to moderately hispid, outer
phyllaries 8-13, spathulate, 2-4 mm. long, 0.6-1.3 mm. wide, inner phyllaries 8-10, 4-6
mm. long, 1.5-2 mm. wide, brown, margins hyaline; ray florets 5-8, white, limb
elliptic-obovate, apically subtruncate, 6-16 mm. long, 3-9 mm. wide, 7-10 striate; disc
florets 26-69, yellow, 3-5 mm. long; achenes linear, obcompressed-quadrangulate,
flattish, glabrous below, tuberculate-strigose above, achene body 4-12 mm. long, dark
brown to black, interior achenes supernally elongate, much longer than the marginal
ones; awns 2, retrorsely barbed, 1-2 mm. long, brown or yellow; chromosome number,
n = 24.
B. alba var. radiata typically bears once-pinnately compound
leaves with large, thin, simple leaflets and broadly spathulate,
apically extrorse, outer phyllaries. These features, together with the
fact that their heads are often eight-rayed, and the ligule limbs two
times as long as broad, combine to distinguish this taxon from the
grossly similar B. odorata var. odorata.
Bidens aurea (Ait.) Sherff, Bot. Gaz. 59: 313. 1915. Coreopsis
aurea Ait. Hort. Kew. 3: 252. 1789 (not C. aurea auct. al.). C. lucida
Cav. Descr. 224, 615. 1802. C. nitida Hort.; Destr. Elench. PI. Hort.
Bot. 10, as syn. 1806. B. luxurians Willd. Enum. 847. 1809 (not B.
luxurians auct. al.). B. arguta HBK. Nov. Gen. & Sp. 4: 231. 1820.
B. decolorata HBK. torn. cit. 233. C. tetragona Cerv. in La Llave &
Lex. Nov. Veg. Descr. 1: 31. 1824. B. arguta var. luxurians DC.
Prodr. 5: 596. 1836. B. tetragona DC. I.e. B. longifolia DC. torn. cit.
597. Diodonta aurea Nutt. Trans. Am. Philos. Soc. II. 7: 360. 1841
(ex syn. Ait.); Walp. Rep. 2: 614. 1843 (ex syn. Ait.). B.
warszewicziana Regel. Flora 32: 183. 1849 (based upon plants
grown at Zurich from seeds said to have been collected in
Guatemala by Warszewicz). B. heterophylla var. wrightii A. Gray,
Proc. Amer. Acad. 19: 16. 1883. B. aurea var. wrightii Sherff, Bot.
Gaz. 85: 16. 1928.
MELCHERT: FLORA OF GUATEMALA 199
Normally a plant of wet habitats on intermountain plateaus,
primarily between 1,800-2,700 m., now particularly common in wet
roadside ditches in cultivated fields, marshy areas along streams,
rarely a weed in dry open habitats; Chimaltenango; Huehue-
tenango; Quezaltenango; El Quiche; Solola. Arizona southward
throughout most of Mexico into Guatemala, common in Chiapas.
Erect, tetragonal-stemmed perennials from elongate rhizomes which are
normally covered with dense clusters of fibrous, annual- like, adventitious rootlets, the
stems simple or branched, erect (rarely subprostrate when growing along highway
shoulders), green to purplish, commonly 0.5-1 m. tall, glabrous or with inconspicuous
scattered pubescence; leaves extremely heterophyllous, in ours most frequently
bipinnatisect with narrowly linear segments, these entire, about 1 mm. wide,
sometimes 3-5-parted with lanceolate, to linear segments, these sharply serrate to
entire, sometimes undivided, submembranous, linear with entire margins or, more
commonly, lanceolate to ovate-lanceolate with numerous, prominent, subcrenate
serrations, to 15 cm. long, 2.5 cm. wide, their petioles more or less alate (the
lanceolate leaf form is the dominant one throughout Mexico); heads numerous,
clustered, slenderly pedunculate, radiate, 3-5 cm. across; involucre diagnostic, outer
phyllaries (5-) 9-13 (-17), narrowly linear, only 3-6 mm. long, 0.5-0.9 mm. wide, a
slight medial expansion often noted in each phyllary, the inner one comparatively
broad, lance-ovate, 4-5 (-6) mm. long, 2-2.8 mm. wide, the greenish brown, lanceolate,
central portion of each very narrow compared with the broad, yellow, rounded,
translucent margins, these 0.5-0.8 (-1) mm. wide; ray florets in ours white or perhaps
pale, creamy yellow (commonly drying yellow), limb linear-obovate, the apex
subtruncate, usually 3-denticulate, 17-25 mm. long, 8-15 mm. wide (in Chiapas and
much of Mexico populations with ligules that are proximally golden-yellow and
distally either paler yellow or white are more common than white-rayed
populations); disc florets yellow, numerous, about 50, their corollas narrowly
cylindrical, (3.5-) 4-6 mm. long; anthers brownish, 2-2.7 mm. long; style branch
appendages short-triangular, 0.5-0.8 mm. long; palae to 13 mm. long, about equalling
teh disc florets, the emergent portion narrowly linear, about 0.3 mm. wide, the tip
rounded, with reddish brown markings; achenes graded monomorphic, linear-cuneate,
blackish, the outer mostly 3.5-4 mm. long, about 1 mm. wide, notably compressed
dorsal-ventrally, but remaining subquadrangulate due to the raised longitudinal
midnerves, the setose apex rather abruptly flattened between the aristae, interior
achenes to 6.5 (-7.2) mm. long, narrower, about 0.7, mm. wide, but remaining flattish
apically; awns 2 (3), yellow, often slightly incurved, 1.5-2 (-2.5) mm. long, retrorsely
barbed supernally; chromosome number n = 23.
Bidens bicolor Greenman, Proc. Amer. Acad. 39: 114. 1903.
Montane weed of pine-oak and oak-mixed hardwood zones,
locally abundant along roadsides and in cornfields on high
intermountain plateaus, also on hillsides, exposed rocky slopes and
open grassy areas within woodlands; Chimaltenango; Guatemala;
Jalapa; Quezaltenango; Sacatepequez; San Marcos; Solola;
Totonicapan. South-central Oaxaca and Chiapas, Mexico.
200 FIELDIANA: BOTANY, VOLUME 24
Erect, terete-stemmed annuals, 1-10 dm. tall at anthesis, the stems branched,
frequently becoming reddish at maturity, scattered hirsute-pubescent to glabrate, the
lowermost portions sometimes developing adventitious rootlets; leaves typically with
3 (5) ovate to ovate-lanceolate segments, rarely bipinnatisect, conspicuously petiolate,
6-20 cm. long including the 3.5-6.5 cm. petiole, leaflets coarsely serrate, pubescent
above and below, sometimes densely so, rarely glabrous, the terminal segment 2-6.5
(-7.5) cm. long, 1-3.5 cm. wide; heads radiate, (3.5-) 4-6 cm. across, with numerous
disc florets; involucre with a tuft of hairs at the base; outer phyllaries mostly 8,
linear-oblong, commonly spreading at maturity, (3.5-) 5-10 mm. long, 0.5-1.5 (-2) mm.
wide, usually glabrous, the midvein and margins with interrupted, dark green lines,
the inner phyllaries lanceolate, usually shorter, frequently becoming blackish when
pressed, commonly, but not necessarily, pubescent, the hairs multicellular; ray florets
5 (7), two-toned yellow, the distal third of the limb lemon-yellow (drying whitish),
the proximal portion orange-yellow with a reddish anthocyanin spot at its base (the
size and definition of this spot varies considerably within single populations with
given individuals lacking this spot entirely, also rays of poorly preserved specimens
may appear uniformly yellow), the limb elliptic-oblanceolate, to 30 mm. long, 18 mm.
wide, apex rounded with a single, inconspicuous, terminal indentation; disc florets
numerous, the corolla yellow, often red-spotted at the base of the tube; anthers dark
brown, 2-3 mm. long, terminal appendages sharply triangular, glandular; style
appendages pointed, about 5 mm. long; palae linear or linear-lanceolate, black-tipped,
yellow with amber and blackish striations; achenes dimorphous, the outermost whorl
clavate, incurved, 3-5 mm. long, yellowish to rubro-castaneous, sometimes slightly
warty, exaristate or biaristate (this outer whorl inconspicuous or undifferentiated in
immature fruiting heads), central achenes blackish, linear, gradually narrowed and
weakly erect -setose above, the outer incurved, the inner slender and erect, to 13 mm.
long; awns 2 (rarely 3), 1-3 mm. long, retrorsely barbed, these sometimes orange
yellow; chromosome number, n = 12.
Bidens bigelovii var. angustiloba (DC.) Ballard in Melchert,
Phytologia 32: 297. 1975. B. anthriscoides var. angustiloba DC.
Prodr. 5: 601. 1836. B. duranginensis Sherff, Bot. Gaz. 70: 90. 1920.
B. bigelovii var. publensis Sherff, op. cit. 88: 287. B. amphicarpa
Sherff, torn. cit. 290. B. oligocarpa Sherff, op. cit. 92: 206. 1931.
Although collected but twice in Guatemala [Guatemala: 5,000
pp. Feb. 1890, Smith 2350 (US); Ciudad Vieja, Oct. 1913, Tyada 95
(US)], B. bigelovii can be expected more frequently because this
self-fertile species is now known from several collections from
Chiapas, Mexico, Costa Rica and Panama. Its main distribution,
however, is in central and northern Mexico and the southwestern
United States.
Plants annual, the stems erect, square, slender, branched, 0.4-1 m. tall, sparsely
pilose; leaves thin, 3-5-partite, 8-13 cm. long, leaflets simple or 2-3 lowest leaflets
tripartite, ovate or ovate-lanceolate, margins serrate or dentate, sparsely to
moderately pilose on both surfaces; heads terminal, 12-24, short-radiate or rarely
discoid, cymosely disposed, 1.5-2.8 cm. wide at anthesis; involucre basally hispid;
outer phyllaries 5-11, linear, 2-4 mm. long, 0.3-0.8 mm. wide, green, cilia te, glabrous to
MELCHERT: FLORA OF GUATEMALA 201
sparsely pubescent, the inner ones 7-9, lanceolate, 2.5-5 mm. long, 1-1.7 mm. wide,
dark brown, glabrous to sparsely pubescent, margin hyaline; ray florets 0-8, neutral
or styliferous, ligule oblong-obovate, 3-10 mm. long, 1.4-4.5 mm. wide, white or
yellow, 5-9-striate, often with a red dot at the base of the limb; disc florets 12-42, 2-
4.5 mm. long, yellow; achenes dimorphic; outer achenes 1-4, linear-cuneate, truncate,
papillose-hispidulous and very scabrous, yellowish or castaneous, achene body 5-7
mm. long, awnless to 2-3-awned, awns retrorsely barbed, 0.3-1.5 mm. long; inner
achenes 12-30, linear, black, glabrate below but commonly erect-hispid above, achene
body 8-12 mm. long; awns 2-3, retrorsely barbed, 1.5-3 mm. long; chromosome
number, n = 24.
B. bigelovii is usually misidentified as one of the varieties of the
B. pilosa complex (sensu Sherff, 1955). Though superficially very
similar, discoid and radiate forms of B. bigelovii can be dis-
tinguished from the discoid B. pilosa var. pilosa and the radiate B.
odorata, B. alba, and B. pilosa var. minor by the fact that its
fruiting heads have a peripheral row of clavate, castaneous or
rubrostramineous achenes. Their small heads (these 1.5-2.8 cm.
wide) and short, sometimes styliferous, rays are also key features.
Bidens blakei (Sherff) Melchert, Phytologia 32: 292. 1975.
Cosmos blakei Sherff, Bot. Gaz. 82: 334. 1926.
Known only from the type, collected at Retalhuleu, Jan. 1871,
Bernoulli and Cairo 1476 (type, Kew; cotype, Petrop.; fragment
including mature achenes at US). Illustration Bot. Gaz. 82, plate 22.
Following description from Sherff, 1955:
Erect herb, probably annual, glabrous below, pubescent above, + 5 dm. tall (the
root not seen), the stem obscurely tetragonal, with long internodes, only moderately
branched; leaves slenderly petiolate with petioles 1.5 cm. long, with petiole included
± 1 dm. long, bipinnatisect, the segments linear, sharply apiculate, minutely hispid
and spinulose-ciliate, 0.5-1.5 mm. wide; heads terminating the branches, coarsely
pedunculate with peduncles to 1.2 dm. long, at anthesis ± 2.5 cm. wide, ± 1 cm. tall;
involucre pubescent, more or less campanulate, the outer bracts spatulate and near
the top often widely rotund-dilated, ciliate, at the apex weakly mucronate, 4-6 mm.
long, the inner ones much larger, conspicuously yellowish when dry, widely
lanceolate, at the apex narrowed and often minutely glandular-ciliate, about 1 cm.
long; ligulate florets about 10-12, yellow, about 1.5 cm. long, the ovary linear,
membranous, sterile, glistening-whitish when dry, erect-hispid on the margins and
along the upper part of the median rib, the body about 1 cm. long, the apex erect-
hispid and weakly aristate with 1 or 2 aristae, these slender, antrorsely hispid, ± 0.5
mm. long; corolla of the disk-florets yellow, only about 5 mm. long; achenes strongly
obcompressed, in the lower part their outline (with the broad wings included)
conspicuously oblanceolate, very glabrous on the blackish-brown faces, or above and
along the median rib erect-setose, on the margins stramineous and erect-ciliate with
minute, spinulose setae, narrowed above into a rostrum, this somewhat stramineous,
erect-ciliate, commonly 3-7 mm. long, the body (including the rostrum) 1.4-2.4 cm.
long, (including the wings) 2.7-3.4 mm. wide, the apex erect-hispid and biaristate, the
aristae erect or scarcely diverging, slender, antrorsely hispid, 1.5-2.5 mm. long.
202 FIELDIANA: BOTANY, VOLUME 24
Bidens chiapensis Brandg. Univ. Calif. Publ. Bot. 6: 76. 1914.
Primarily in montane forests, 1,900-3,400 m., most frequently
sprawling over banks and thickets at the forest's edge, on shaded
slopes of steep ravines, volcanic slopes, and in oak, pine, and /or
mixed hardwood forests, particularly cutover ones, one collection
from a swampy meadow; Chimaltenango; Huehuetenango; Quezal-
tenango; El Quiche; San Marcos; Solola; Totonicapan. Chiapas
(and doubtfully Michoacan) Mexico.
Weak, scrambling half-shrubs up to 1 m. tall or rarely erect and to 1.5 m., the
stems terete, often purplish, glabrous throughout, internodes of sprawling forms
elongate; leaves uniformly trifoliate, long-petiolate, upper surface dark green,
conspicuously ciliate, otherwise essentially glabrous (trailing edge of the lateral
leaflets rarely lobed or divided), leaflets ovate or lance-ovate, tips long-acuminate,
margins serrate to incised-serrate throughout, indurate-mucronate, teeth pointing
forward; heads pale yellow, short-radiate, only 2-3 cm. across, long-pedunculate, 1-3
terminating each main branch; outer involucre of (8-) 12-16, narrowly linear
phyllaries (6-) 9-14 mm. long, 0.8-1.8 (-2) mm. wide, at maturity typically spreading or
recurved, 1-3 lined, usually equalling to much exceeding the inner involucre, some
equalling the rays, the inner phyllaries mostly 6-8 mm. long, chaffy-yellow,
multistriate, frequently red pigmented, especially toward the obtuse apex; ray florets
pale yellow, relatively inconspicuous, 8-10, the limb narrowly elliptic or linear -elliptic,
8-14 (-15) mm. long, to 5 mm. wide; disc florets numerous, yellow, corollas 7-9 mm.
long, their teeth sharply lanceolate; anthers black, 3.5-4 mm. long; style branch
appendages attenuate, 1-1.2 mm. long; mature achenes, particularly the outer,
supernally recurved, all linear-tetragonal, slightly narrowed above, black, glabrous, 7-
13 mm. long, 2-4 aristate within single heads; major awns 3-5 (-6) mm. long, one or
two shorter, retrorsely barbed throughout; chromosome number, n = 12.
Bidens chrysanthemifolia (HBK.) Sherff, Bot. Gaz. 61: 501.
1916. Cosmos chrysanthemifolius HBK. Nov. Gen. & Sp. 4: 239, t.
382. 1820. Cosmea chrysanthemifolia Spreng. Syst. 3: 615. 1826.
Cosmos chrysanthemoides HBK.; DC. Prodr. 5: 607. 1836 (sphalm.).
Bidens kunthii Sch. Bip., Seem. Bot. Voy. Herald 308. 1856. B.
parvulifolia Sherff, Bot. Gaz. 56: 490. 1913. B. chrysanthemifolia
var. parvulifolia Sherff, Field Mus. Bot. 22: 436. 1941. B.
chrysanthemifolia var. typica Sherff, Brittonia 6: 340. 1948.
Wet, montane forests, primarily 2,000-3,500 m., one collection
at 900 m., particularly common at the edge of, or in open spots
within, pine and/or oak woods, roadsides, mountain pastures,
meadows, moist thickets; Baja Verapaz; Chimaltenango; Guate-
mala; Huehuetenango; Jalapa; Quezaltenango; El Quiche; Sacate-
pequez; San Marcos; Solola. Chiapas, Mexico; El Salvador.
Sprawling, procumbent to somewhat ascending, perennial herbs with several
main stems radiating from a narrow central rootstock, these commonly with
MELCHERT: FLORA OF GUATEMALA 203
adventitious rootlets near their bases, the stems tetragonal to subtetragonal, angulate
when pressed, up to 1 m. long, often branched, densely short-pilose to nearly
glabrous, often reddish striped; leaves highly variable, mostly lobed, lance-ovate, apex
acute, bases rhombic-truncate, crenate-serrate throughout, 3-9 cm. long including the
petiole (to 14 cm. long in certain Chiapas populations), 1.5-4 cm. wide, most densely
short-pubescent, some nearly glabrous, sometimes trifoliate with lanceolate segments,
or more rarely, 5-partite or evenly bipinnatisect with narrow, linear-lanceolate
segments; heads showy, 2.5-4 (-4.5) cm. across, rays cream-white, disc yellow, either
solitary, or, perhaps more commonly, with about 3 long-peduncled heads terminating
each branch, the main peduncles 5-23 cm. long, sometimes once-branched but the
heads never tightly clustered; outer phyllaries 8-12, linear to linear-subspathulate,
two-thirds as long to subequal with the inner phyllaries, 4-6 mm. long, 0.8-1 mm.
wide, or when dilated supernally to 2 mm. wide, the surface marked with an amber
midnerve which sometimes becomes blackish above, similar marginal nerves are
usually, but not invariably, present, ciliate; the inner phyllaries lanceolate, apex
obtuse, the chaffy surface frequently drying glossy grey to blackish, usually with
scattered to rather dense, multicellular hairs, very rarely glabrous; ray florets 5-8
between or within populations, creamy white, but frequently drying yellowish (such
individuals repeatedly misidentified as B. triplineruia), ligules linear- or elliptic-
obovate, (12-) 15-20 mm. long, mostly 9-10 mm. wide, the apex somewhat truncated,
obscurely 3-denticulate; disc florets numerous, yellow, sometimes orangish with age,
4-6 (-7) mm. long; anthers 2-3 mm. long, black; style branches about 2 mm. long,
their terminal appendages narrow, long-tapered, never decidedly triangular; chaff
linear-lanceolate, to 8 (11) mm. long, yellow with orange and/or black stripes, when
confluent the entire central portion appearing blackish, the apex obtuse, short-ciliate;
achenes somewhat dimorphous, few or several at the periphery of the head
stramineous to orangish or rarely even purple, short, mostly 3-4 mm. long, strongly
obcompressed to subclavate, short-awned or exaristate (these outer achenes falling
quickly, only present on complete heads), all other achenes becoming black, 3.5-6 (-7)
mm. long, linear-tetragonal, the outer black ones obcompressed, slightly incurved, the
elongate central ones erect, narrowly fusiform; awns 2 (3), rarely exaristate, 0.9-2.5
mm. long, orangish yellow, their barbs retrorse, supernal only; chromosome number,
n = 24.
Bidens holwayi Sherff & Blake in Sherff, Bot. Gaz. 64: 39.
1917.
Plants of damp montane forests in southwestern Guatemala,
commonly climbing high into trees, hanging over moist shaded
slopes or streambanks, scrambling in ravines or over thickets, some
at the forest edge shrublike, 1,500-3,500 m.; Chimaltenango (?);
Huehuetenango; Quezaltenango (type, Holway 816)', San Marcos;
Solola.
Large, scandent vines with stems to 20 or 30 m. long, or scrambling and
shrublike, to 3.5 m. tall, the stems terete below, usually angular or even subtetragonal
toward the inflorescence; leaves 3-5-partite with large, lanceolate to lance-ovate
segments, occasionally simple, 9-15 (-18) cm. long, distinctly petiolate, these (2.5-) 3-5
cm. long, leaflets darker above (usually pressing brownish green), glabrate or with
scattered hairs, paler and more densely pilose-hispid below, particularly so along the
204 FIELDIANA: BOTANY, VOLUME 24
veins, margins sharply serrate to the abruptly rounded base, the tip narrowly long-
acuminate, entire, terminal segments of tripartite leaves mostly 8-10 (-14) cm. long, 2-
4 (-4.5) cm. wide, simple leaves to 5 cm. wide; heads large, radiate, yellow throughout,
mostly (4-) 5-6 cm. across, to 2.2 cm. tall in fruit (awns included), discs 1.2-1.6 cm.
wide at anthesis; inflorescence of heads in cymosely-branched, paniculate clusters,
peduncles of the terminal clusters mostly 3-7 (-13) cm. long, becoming coarse in fruit;
outer phyllaries (5-) 8-11, broadly linear to linear-oblanceolate, 7-9 (-14) mm. long,
1.8-2.5 mm. wide, cilia te, with 3-5 dark nerves, glabrous or pubescent on the upper
mid-portion; ray florets (5-) 6-8, the limb linear-elliptic or narrowly oblanceolate,
pale yellow with 7-9 dark lines, mostly 2-2.5 (-2.8) cm. long, 0.5-0.8 cm. wide, the apex
often irregularly toothed; disc florets numerous, the corollas (5.5-) 6-8 (-9) mm. long,
pale yellow; anther tube black, 4-4.5 mm. long; achenes linear, black, flattish, but the
dorsal and ventral nerves somewhat raised, margins with conspicuous, spreading or
ascending, white cilia, these usually arising in groups from marginal tuberculae,
achene body 12-18 mm. long, about 1 mm. wide throughout, the apex antrorsely
setose, sometimes greenish; awns 2 (rarely 3 or 4), erect or at maturity the outer
frequently spreading or recurved, 4-6.5 (-10) mm. long, retrorsely barbed.
See discussion following B. squarrosa.
Bidens laevis (L.) B.S.P. Prel. Cat. N. Y. 29. 1888. Helianthus
laevis L. Sp. PL 906 (ex cit.). 1753. Coreopsis radiata Mill. Gard.
Diet. ed. 8, Coreopsis no. 5 (ex descr.). 1768. Bidens chrysanthe-
moides Michx. Fl. Bor.-Am. 2: 136. 1803. B. helianthoides HBK.
Nov. Gen. & Sp. 4: 230. 1820. B. quadriaristata DC. Prodr. 5: 595.
1836. B. speciosa Parish, Zoe 5: 75. 1900 (not B. speciosa Gardn.
1845). B. elegans Greene, Pittonia 4: 254. 1901. B. lugens Greene,
loc. cit. B. formosa Greene, torn. cit. 264 (not B. formosa Sch.-Bip.
1856). B. parryi Greene, torn. cit. 265. B. expansa Greene, torn. cit.
266. B. persicaefolia Greene, loc. cit.
Not reported from Guatemala, but may be expected there since
it is established on reservoir and pond banks in San Cristobal de las
Casas, Chiapas, Mexico. Eastern United States coastal plain, west
to California, widely scattered in Mexico and South America, also
Hawaii.
Erect or suberect, large-yellow-headed, glabrous, subaquatic plants with clusters
of elongate, adventitious roots from their lower nodes, the base often procumbent,
seemingly perennial, to about 1 m. tall, the stems terete, often purplish, usually with
regularly disposed, opposite branches in the inflorescences; leaves sessile, unlobed,
broadly lance-linear, gradually narrowed at each end, the base broad, often
somewhat connate, the apex acuminate, regularly serrate throughout, the teeth often
slender, mostly 5-15 cm. long; heads radiate, yellow, mostly 5-7 cm. across, erect at
anthesis but usually nodding in fruit, the large disc hemispheric, 2-2.5 (-3) cm. across,
1.3 cm. tall; outer phyllaries 5-8, subfoliose, broadly linear, 8-12 (-25) mm. long, 2-5
mm. wide, tip usually obtuse, conspicuously veined, often spreading, inner phyllaries
broadly ovate, acute, mostly 10-11 mm. long, 5-7 mm. wide, central portion chaffy,
multistriate, margins yellow; ray florets mostly 8, yellow, deeper proximally, the limb
MELCHERT: FLORA OF GUATEMALA 205
oval, large, to 3 cm. long, 1-1.5 cm. wide, the rounded apex barely, if at all, toothed;
disc florets very numerous, corollas yellow, 4.5-5.5 mm. long, the tube slender, about
2.5 mm. long; palae much narrower than the inner phyllaries, becoming linear,
usually but not necessarily orange- tipped, lighter below; achenes narrowly cuneate,
flattened dorsal-ventrally, the edges retrorsely barbed, 5-6 mm. long, the outer widest,
the flattened apex about 2 mm. wide, tapering to about 0.8 mm. at the base, the
dorsal surface longitudinally multistriate with thickened, yellow margins, midrib of
the ventral surface sometimes raised, the achenes then 3 (or 4) angulate; awns 2-3
(-4), erect, (2-) 3-5 mm. long, retrorsely barbed throughout; chromosome number, n
= 12.
Bidens odorata Cav. Ic. 1: 9. 1791. Coreopsis odoratissima
Cav. in Pers. Syn. PI. 2: 477. 1811. C. odorata Poir. in Lam. Encyc.
Suppl. 2: 350. 1811. Cosmos tennellus HBK. Nov. Gen. & Sp. 4: 420.
1820. B. daucifolia DC. Prodr. 5: 601. 1836. B. caucalidea DC. torn,
cit. 604. B. bonplandii Sch.-Bip. in Seem. Bot. Voy. Herald 308.
1856. B. inermis S. Wats. Proc. Amer. Acad. 23: 278. 1888. B. deamii
Sherff, Bot. Gaz. 56: 490. 1913. B. ramosissima Sherff, torn. cit. 491.
B. pilosa var. bimucronata f. odorata (Cav.) Sherff, op. cit. 81: 41.
1926. Coreopsis ferulaefolia var. odoratissima Pers.; Sherff, Field
Mus. Publ. Bot. 16: 444. 1937.
Aggressive, widespread plants of roadsides, cultivated fields, and
numerous other highly disturbed habitats in montane and high
plateau communities, between 1,500-3,500 m.; Chimaltenango;
Guatemala; Huehuetenango; Quezaltenango; Sacatepequez; Solola;
Totonicapan. Found primarily in northern and central Mexico
(southwestern Chihuahua and Nuevo Leon in the north, southward
into Mexico and Puebla) with disjunct populations in New Mexico,
Chiapas, and Guatemala. The latter are effectively isolated
ecologically by the tropical lowlands of the Isthmus of Tehuantepec
and a range of arid, lower mountains in Oaxaca.
Plants annual, the stems erect, branched, square, 0.3-1.5 m. tall, sparsely pilose;
leaves petiolate, opposite, 3-10 cm. long, 5-8 cm. wide, leaflets simple to pinnatifid,
sparsely pilose on both surfaces; heads terminal, cymosely disposed, 21-42, 2.5-4.6 cm.
wide at anthesis; involucre basally hispid; outer phyllaries 6-10, linear to linear-
spathulate, 3-5 mm. long (averaging 3.7 mm.), 0.5-1 mm. wide, green, ciliate, surface
glabrous to moderately pubescent, inner phyllaries 6-8, lanceolate, 3-6 mm. long, 1.3-2
mm. wide, brown, margin hyaline, glabrous to moderately pubescent; ray florets 5,
ligules elliptic-obovate, apically subtruncate, 8-18 mm. long (averaging 12.7 mm.), 6-
12 mm. wide, white, rosaceous, or rarely light yellow, 8-14-striate; disc florets 30-53,
3-6 mm. long, yellow; achenes 30-53, linear, obcompressed-quadrangulate or flattish,
glabrous below, tuberculate-strigose above, achene body 5-14 mm. long, brown to
black, interior achenes supernally elongate and much longer than marginal ones;
awns 0-2, retrorsely barbed, 1-3 mm. long, yellow; chromosome number, n = 12.
Specimens of B. odorata var. odorata can be distinguished from
206 FIELDIANA: BOTANY, VOLUME 24
var. calcicola and the grossly very similar B. alba var. radiata by
the following features: (1) heads 2.5-4.6 cm. wide at an thesis; (2)
heads 5-rayed; (3) 12.7 cm. average length of the ray floret limbs,
and (4) 3.7 mm. average length of the outer phyllaries.
Bidens odorata var. calcicola (Greenm.) Ballard in Melchert,
Phytologia 32: 1975. Cosmos pilosus HBK. Nov. Gen. & Sp. 4: 241.
1820. B. exaristata DC. Prodr. 5: 600. 1836. B. brachycarpa DC. loc.
cit. B. rosea Sch.-Bip. in Seem. Bot. Voy. Herald 308. 1856. B. rosea
var. calcicola Greenman, Proc. Amer. Acad. 41 : 264. 1905. B. pilosa
var. calcicola Sherff, Bot. Gaz. 80: 377. 1925. B. orendainae M. E.
Jones, Contr. W. Bot. 18: 82. 1933. B. barrancae M. E. Jones, loc.
cit.
Weed of roadsides and other disturbed habitats between 1,100-
1,800 m.; Chimaltenango; Guatemala; Jalapa; Jutiapa; Santa Rosa.
B. odorata var. calcicola is centered in the Sierra Volcanica
Transversal of southwestern and central Mexico (particularly in
Michoacan and Jalisco) with small populations scattered into
northern Mexico, in the vicinity of the Oaxaca-Chiapas border, and
in Guatemala.
Similar to var. odorata, but heads 1.8-2.6 cm. wide at anthesis; outer phyllaries
7-11, 1.2-3 mm. long (2.3 mm. average), 0.2-0.5 mm. wide, inner phyllaries 7-9, 2-4 mm.
long, 1-1.5 mm. wide; ray florets 5-8, limbs 4-8 mm. long (5.8 mm. average), 3-6 mm.
wide, white to subrosaceous, 6-9-striate; disc florets 20-44; achene body 2.5-14 mm.
long; chromosome number, n = 12.
These populations were included by Sherff as B. pilosa var.
calciola.
Bidens ostruthioides (DC.) Sch.-Bip. in Seem. Bot. Voy.
Herald 308. 1856. Delucia ostruthioides DC. Prodr. 5: 633. 1836. B.
costaricensis Benth. in Oerst. Vidensk. Meddel. 1852: 94. 1852. B.
guatemalensis Klatt, Bot. Jahrb. 8: 44. 1887 (type from San Marcos,
Lehmann 1560). B. ostruthioides var. costaricensis Sherff, Bot. Gaz.
88: 298. 1929. B. ostruthioides var. typica Sherff, Brittonia 6: 341.
1948. B. ostruthioides var. matritensis Sherff, Am. Journ. Bot. 38:
66. 1951.
Plants of openings in high altitude, montane forests, partic-
ularly "cloud forests"; especially common on moist roadside or in
stream bank thickets, open mountain summits, etc. in Pinus-
Quercus, or Pinus- Abies and Abies-Cupressus communities; Chi-
maltenango; Huehuetenango; Jalapa; Quezaltenango; El Quiche;
MELCHERT: FLORA OF GUATEMALA 207
San Marcos; Solola; Totonicapan. Jalisco and Veracruz, Mexico,
southeastward at high elevations into northwesternmost Panama.
Sprawling, multistemmed, suffruticose herbs from a thickened, woody rootstock,
usually decumbent (ours rarely erect), the stems elongate, 1-1.5 or even 2 m. long, the
basal portion terete, woody, usually glabrous, current growth subterete, multinerved,
the nerves often purplish, glabrous, or more commonly scattered-pubescent; leaves
mostly 3- (5-) partite with broad, ovate to rhomboid-ovate segments which have only
1-5 coarse, dentate-serrate, indurate-mucronate teeth per segment margin, sometimes
simple above (in Mexico rarely simple throughout), membranous, dark green above,
pale below, ours 3-6.5 cm. long including the ciliate petiole (to 10 cm. in Chiapas),
lateral segments 2-3 cm. long, 1-2 cm. wide, the terminal ones 2-3.5 cm. wide, to 4 cm.
wide when leaves simple; heads large, radiate, 3-5 (-6) cm. across, mostly solitary on
elongate peduncles, these commonly to 2 cm. long; outer involucre of 5 (7) foliaceous
phyllaries, these elliptic-ovate to broadly linear or lanceolate, ours mostly (5-) 7-12
mm. long, (1.8- ) 2.5-4 mm. wide (to 11-17 mm. long and 4.5-10 mm. wide and much
exceeding the inner phyllaries in certain Chiapas populations), 3-5 pale-amber nerves
seen when pressed, commonly ciliate; inner involucre normally orange-yellow, chaffy-
membranous, the individual phyllaries lanceolate to ovate-lanceolate, the broad
central portion multistriate, orange-yellow, the narrow, scarious margins yellow; ray
florets 5 or 6 (rarely 8), fertile, each with a T-shaped style protruding about 4 mm.
above its tube, the ligules yellow, sometimes paler distally, elliptic- to linear-obovate,
(14-) 20-26 mm. long, (8-) 10-15 mm. wide, the rounded apex 3-denticulate; disc
florets numerous, the corollas yellow above, notably paler than the orange-yellow
chaff which project between them, whitish below, the tube elongate, about two-fifths
the corolla length; anthers black, 2.6-3.2 mm. long; stigmatic areas short attenuate-
triangular, about 0.5 (0.6) mm. long; palae orange-yellow, similar to the inner
involucre, the inner palae linear-lanceolate, to 12 mm. long; achenes pallid tan to
light olivaceous, never becoming black, linear- cuneate, obcompressed, the dorsal
surface slightly rounded, longitudinally multinerved, the nerves equal sized, the
ventral surface with a raised midnerve, the two adjacent faces each with several
smaller nerves, outer achenes 4-8 mm. long, to 2 mm. wide, the inner to 12.5 mm.
long, about 1 mm. wide, all glabrous, 2-3-aristate; major awns 3-5.5 mm. long, each
having 6-8 sets of antrorse barbs spaced evenly over their entire length (rarely
quadriaristate, these much shorter); chromosome number, n = 23.
Bidens ostruthioides collections with pinnatisect rather than
tripartite leaves have traditionally been segregated as either var.
costaricensis or var. matritensis (Sherff, 1955). Though occurring
mainly from Costa Rica to Panama, i.e., in the southern part of B.
ostruthioides range, pinnatisect forms with narrow leaflets and
outer bracts have been found as far northwest as Jalisco. Since the
available collections demonstrate various degrees of leaf lobing
between tripartite and narrowly bipinnatisect, since no other
distinguishing morphological traits were found to correlate with a
particular leaf pattern, and since tripartite and bipinnatisect leaf-
form segregates are commonplace within single populations of most
of our Bidens species, all B. ostruthioides collections are treated
here as part of a single, variable taxon.
208 FIELDIANA: BOTANY, VOLUME 24
Bidens pilosa L. Sp. PL 832. 1753. B. reflexa Link, Enum.
Hort. Berol. 2: 306. 1822. B. adhaerancens Veil. Fl. Flum. 348, pro
parte. 1825. B. hirsuta Nutt. Trans. Am. Philos. Soc. II. 7: 369. 1841.
B. pilosa var. discoidea Sch.-Bip. in Webb. & Barth. Phyt. Canar. 2:
242. 1844. B. leucantha f. discoidea Sch.-Bip. Flora 27: 673. 1844. B.
leucanthus var. pilosus Griseb. Cat. PI. Cub. 155. 1886. Kerneria
pilosa Lowe, Man. Fl. Madeira 1: 474. 1868. K. pilosa var. discoides
Lowe, loc. cit. B. montaubani Philippi, Anal. Mus. Nac. Chile Bot.
1891: 49. 1891.
An eminently successful pantropic weed; ours in a variety of
disturbed habitats between sea level and 2,500 m.; one collection
from Santa Rosa erroneously reported at 4,000 m.; very common;
Alta Verapaz; Chimaltenango; Chiquimula; Escuintla; Guatemala;
Huehuetenango; Izabal; Jalapa; Peten; Quezaltenango; Retalhuleu;
Sacatepequez; Santa Rosa; Solola; Suchitepequez; Zacapa. In
North America this primarily discoid, self-fertile hexaploid has a
widespread, more or less continuous distribution from California
southward through Mexico and Central America.
Plants annual, the stems erect, branched, square, 0.3-1.8 m. tall, sparsely pilose;
leaves petiolate, opposite, 2.5-13.5 cm. long, 2-11 cm. wide, leaflets simple, ovate or
lanceolate, apically acuminate, margins serrate, sparsely pilose on both surfaces;
heads terminal, cymosely disposed, 21-42, 0.7-1 mm. wide at anthesis; involucre
basally hispid; outer phyllaries 7-10, linear to linear-spathulate, 2-7.5 mm. long, 0.5-
1.2 mm. wide, green, ciliate, surface glabrous; inner phyllaries 8-10, lanceolate, 3-5
mm. long, 1-2 mm. wide, brown, glabrous, margin hyaline; ray florets usually absent,
when present minute, 2-3 mm. long, tubular, fertile, white; disc florets 35-75, 3-4 mm.
long; achenes 25-65, linear, obcompressed-quadrangulate, glabrous below, tubercu-
late-strigose above, achene body 8-16 mm. long, dark brown to black, interior achenes
supernally elongate and much longer than marginal ones, awns 3 (occasionally 2, 4,
or 5), retrorsely barbed, yellow; chromosome number, n = 36.
Morphologically, B. pilosa is easily recognized by a
combination of the following features: (1) simple, ovate or
lanceolate leaflets, (2) heads discoid or pseudoradiate, (3) when
present the ray florets (actually slightly flattened disc florets) are
white, 2-3 mm. long, tubular in shape, and fertile, and (4)
monomorphic achenes, these usually 3-awned.
Bidens pilosa var. minor (Blume) Sherff, Bot. Gaz. 80: 387.
1925. Coreopsis leucorrhiza Lour. Fl. Cochinch. 508. 1790 (ex descr.
et patria.). Bidens hispida HBK. Nov. Gen. & Sp. 4: 237. 1820.
Kerneria dubia Cass. Diet. Sci. Nat. 24: 398, in part. 1822. B.
sundaicus Blume, Bijdr. 913. 1826. B. sundaicus var. minor Blume,
op. cit. 914. B. lleucorhiza DC. Prodr. 5: 605. 1836. B. leucantha
MELCHERT: FLORA OF GUATEMALA 209
var. sundaica Hassk. Cat. Hort. Bog. 100. 1844. B. pilosa var.
brevifoliata Hieron. Bot. Jahrb. 29. 48. 1900.
Not known from Guatemala but collected several times in
weedy lowland areas of British Honduras, and from 800 to about
1,500 m. in El Salvador. Common in Costa Rica and Panama.
Like B. pilosa var. pilosa, but leaflets more often sharply tipped and sharply
serrate; heads subradiate; ray florets 4-7, white or yellow, irregularly 3-5-lobulate
above, ligules 5-8 mm. long; style lacking or present; awns 2-3, 1-2 mm. long;
chromosome number, n = about 36.
Excepting its ligules, var. minor is virtually identical to, and
perhaps only a segregating form of, var. pilosa.
Bidens riparia HBK. Nov. Gen. & Sp. 4: 236. 1820. B. refracta
Brandg. Zoe 1: 310. 1890. B. riparia var. refracta (Brandg.) O. E.
Schulz, Symb. Ant. 7: 132. 1911. B. ambrosioides Willd. in O. E.
Schulz, loc. cit. B. riparia var. typica Sherff, Brittonia 6: 340. 1948.
A tropical weed, ours occurring primarily in damp thickets
between 200-900 m., also in ravines, along arroyas, on brushy, rocky
slopes; Chimaltenango; Chiquimula; Guatemala; Jutiapa; Santa
Rosa; Zacapa. Common throughout Central America; in Mexico
along the gulf and Pacific coastal plains to Veracruz and Baja
California, respectively; South America.
Erect, square-stemmed, small-headed, thin-leaved, annual herbs, to 1.5 m. tall,
branched; leaves mostly trifoliate with ovate to lance-ovate segments, occasionally 7-
partite or bipinnatisect, 2-8 (-16) cm. long, the petiole narrow, mostly two-fifths to
one-half the entire leaf length, leaflets of the trifoliate forms acuminate, sharply
serrate to their cuneate bases, upper surface dark green, scabrous ciliate, lower
surface paler, scattered pubescent, dark reticulate, terminal segments usually largest,
to 3 (4) cm. wide; heads few, inconspicuously radiate, often appearing discoid, small,
only 8-10 (-13) mm. wide at anthesis, but becoming greatly enlarged in fruit; outer
phyllaries 8-13 (-16), linear, some gradually expanded supernally, glabrous, mostly 4-6
mm. long (to 10 mm. in fruit), 0.5-1 mm. wide, normally spreading and slightly to
much exceeding the inner phyllaries (rarely subfoliose, to 20 mm. long, 3 mm. wide),
the inner phyllaries lanceolate, only 3-4 mm. long, red-brown striate, white margined;
palae similar but narrower; ray florets mostly 5 (reportedly rarely discoid), ligules
pale yellow, becoming dark striate and often whitish when pressed, the limb narrowly
linear-oblanceolate, 4-6 mm. long, to 2 mm. wide, barely exceeding the disc when
dried, apex obtuse, barely lobed; disc florets 20-30, the corolla yellow with dark veins,
only 3-3.5 mm. long; anthers brown; style appendages minute; achenes all elongate,
linear- tetragonal, with one erect and 3 (rarely 4) divergent to reflexed, retrorsely
barbed awns, the body brown to olivaceous, stippled with black, never becoming black
throughout, dimorphous in size, color, and texture, 6-9 of the peripheral achenes in
each head densely an trorse- hispid throughout, 7-9 (-13) mm. long, mostly one-half to
three-fourths the length of, and darker than, the central achenes (one specimen seen
with purplish-castaneous outer achenes), central achene 10.5-16 (-19) mm. long,
210 FIELDIANA: BOTANY, VOLUME 24
narrowed slightly supernally, with scattered, antrorse, hispid hairs above; awns 4 (or
5), retrorsely barbed, the erect one to 4 mm. long, the 3 or 4 divergent ones 2-3 mm.
long, the central one usually shortest; chromosome number, n — 12.
Bidens rostrata Melchert, Phytologia 32: 291. 1975 Cosmos
exiguus A. Gray; S. Wats. Proc. Amer. Acad. 22: 429. 1887, not B.
exigua Sherff, 1920.
Plants of open, grassy areas, on rocky hillsides, or limestone
ledges, or outcrops, in pine-oak woods. Collected only once in
Guatemala; Huehuetenango, Rio Pucal, about 14 km. south of
Huehuetenango, about 1,780 m., pine-oak forest, scarce. Several
collections from Central Chiapas; in Mexico scattered collections
across the Volcan Transversal into Jalisco then northward along
the Sierra Madre Occidental into Chihuahua.
Inconspicuous, slender annuals with tiny, short-radiate, rosaceous heads, the
stems sharply tetragonal, glabrous, only 1.5-4.5 (-6) dm. tall, becoming almost filiform
above, normally unbranched below the inflorescence; leaves few, narrowly linear to
linear- filiform, entire, 1.5-9.5 cm. long, margins sometimes subrevolute, lower surface
glabrous, margins and upper surface often setose; heads several, well separated on
slender, naked peduncles, minute in bud, to only 8 mm. across at full anthesis (rays
included), disc only 2-3 mm. across, becoming greatly expanded in fruit; involucre
dimorphic in size only, outer as well as inner phyllaries and palae greenish with
numerous, amber to greenish-black striations; outer phyllaries subulate to narrowly
triangular, 1-2 (-2.5) mm. long, a tuft of multicellular hairs at their base, these
sometimes purplish, the inner phyllaries much larger, oblong to lance-oblong, 4-6 mm.
long, their rounded tips and hyaline margins sometimes rosaceous; palae still longer,
to 7 mm.; ray florets mostly 5 (2-5 within single populations), purplish- la vender with
darker veins, drying dark-rosaceous, tiny, 4-6 mm. long including the tube of about 2
mm., only slightly exceeding the disc, the limb truncate-obovate, 3-denticulate; disc
florets few, about 13, whitish or the upper portion partially to totally rosaceous, teeth
occasionally yellow, 3-4.5 mm. long; achenes strongly flattened, linear, attenuate-
rostrate, lower fertile portion multinerved, becoming black, somewhat concave
dorsally, rostrum flat, olivaceous, biaristate, marginally antrorse-hispid; awns 2,
erect, slender, 1.5-4 mm. long, antrorsely barbellate; chromosome number, n = 12.
Bidens squarrosa HBK. Nov. Gen. & Sp. 4: 238. 1820. B.
tereticaulis DC. Prodr. 5: 598. 1836. Coreopsis trifoliata Bertol.
Novi Comm. Bonon. 4: 436. 1840 (type from Volcan de Agua,
Velasquez s.n.). B. antiquensis Coult. Bot. Gaz. 16: 100. 1891 (type
from Antigua, Sacatepequez, J. D. Smith 2354). C. scandens Sesse &
Moc. Fl. Mex. ed. 2. 1894. B. tereticaulis var. sordida Greenm. Proc.
Amer. Acad. 39: 115. 1903. P. tereticaulis var. indivisa Robins. Proc.
Bost. Soc. Nat. Hist. 31: 270. 1904. B. coreopsidis var. procumbens
Donn.-Sm. Bot. Gaz. 42: 299. 1906 (type from Secanquim, Alta
Verapaz, Maxon & Hay 3162). B. tereticaulis var. antiguensis O. E.
Schulz in Urban, Symb. Ant. 7: 142. 1911. Figure 54.
MELCHERT: FLORA OF GUATEMALA 211
Common, usually abundant; scrambling in thickets or climbing
in trees in a variety of open or disturbed, woodland habitats from
100-2,000 m.; particularly common in shrubby, roadside slopes or
thickets, hedgerows, etc., wooded or shrubby slopes, cliffs, in
secondary growth mountain forests; Alta Verapaz; Baja Verapaz;
Chimaltenango; Chiquimula; Escuintla; Guatemala; Huehue-
tenango; Izabal; Jalapa; Jutiapa; Peten; Quezaltenango; El Quiche;
Retalhuleu; Sacatepequez; Santa Rosa; Solola; Suchitepequez.
British Honduras; El Salvador; Veracruz and Jalisco, Mexico
southeastward throughout Central America; West Indies; South
America.
Scrambling or climbing, herbaceous vines or shrublike, to 2 m. tall, the stems
terete, multistriate and mostly glabrous below, new growth often pilose-pubescent
and somewhat angled (especially when pressed); leaves mostly trifoliate with
lanceolate or ovate-lanceolate segments, sometimes unlobed, sometimes ovate-
lanceolate, 5-partite with narrowly lanceolate segments, 5-10 (-15) cm. long, including
the narrow, 2.5-4 (-5.8) cm. petiole, pubescence variable, glabrous above with
scattered hairs below, to densely short-pilose throughout, the lower surface more
pubescent than the upper, occasionally glabrous throughout; leaflets usually abruptly
rounded at the base, sharply serrate throughout excepting the long-acuminate, entire
apex, the teeth with forward-pointing, mucronate tips, terminal leaflet usually
largest, stalked, 4.5-9 cm. long, 1.5-4 cm. wide; heads numerous, usually densely
clustered, the inflorescence of cymosely- branched racemes or paniclas, radiate or
occasionally discoid; radiate heads mostly 2.5-2.7 (-3) cm. wide, the disc 7-10 (-12)
mm. wide, 8-10 mm. tall at anthesis, in fruit to 10 mm. wide, 10-12 mm. tall (awns
included); ray florets none or 3-5 (-6), the number per head often varying within
individual plants, pale yellow with 7-11 darker veins, narrowly oblong to linear-
oblanceolate, 7-14 (-19) mm. long, (2.5-) 3-5 mm. wide, the apex regularly, but
obscurely, 3-denticulate, outer phyllaries 5-8, mostly linear to linear-spathulate, 0.5-
0.9 (-1.2) mm. long, 3-lined, commonly ciliate, glabrous to densely short-pilose, the
inner phyllaries linear-lanceolate, 5-7 mm. long, chaffy and reddish brown in fruit;
disc florets yellow, small, narrowly cylindrical, numerous, their corollas only (3.5-) 4-
5 mm. long; anthers brown to blackish, 2.3-3 mm. long; palae similar to, but narrower
than, the inner involucre; achenes linear, flattish, black, the margins densely invested
with spreading or ascending, white cilia, these pilose hairs usually arising in clusters
from small marginal tuberculae, achene bodies 6-9 (-10) mm. long, about 0.8 mm.
wide throughout, only slightly narrowed above, the apex antrorsely setose, becoming
greenish black; awns 2, initially erect, but some in each head becoming recurved or
horizontally spreading, smooth or retrosely (very rarely antrorsely) barbed.
The vast majority of the Guatemalan B. squarrosa collections
have numerous, small heads (discs 8-10 mm. wide) in congested,
short-pedunculate panicles. In the heavily collected central high-
lands most of these plants are short-radiate (the limbs 8-14 mm.
long) and have smooth awns. Though mostly 5, the number of ray
florets frequently varies from 3-5. Similar, but completely discoid
plants, are found along the southern tier of states in Guatemala
212 FIELDIANA: BOTANY, VOLUME 24
from Retalhuleu to El Salvador. Plants of this small-headed, short-
rayed type from the highland area near San Cristobal, Chiapas are
known to be diploid (n = 12).
Scattered collections from lowland regions of Alta Verapaz,
Izabal, Peten, and British Honduras tend to have thick, unlobed,
sometimes coriaceous leaves, and fewer, larger heads in rather open
inflorescences (rays mostly 14-16 mm. long). B. squarrosa plants of
this type are also known from the Yucatan peninsula in Mexico. It
must be emphasized, however, that many intermediates are known.
A few large-headed (to 4 cm. across), long-peduncled plants
have also been collected in the central highlands (Chimaltenango,
Huehuetenango, El Quiche, Solola, and Zacapa). Unlike their
"lowland" counterparts these plants all have 3-5-partite leaves with
elongate, lanceolate segments and rather stout, retrorsely barbed
awns, features which combine to give them the appearance of a
"small B. holwayi."
Large-headed B. squarrosa-type plants are common in Panama
and Costa Rica. Like our B. holwayi, the ray number of these
plants varies from 5-8, sometimes on a single plant. Comparatively
large-headed, 5-8-rayed B. squarrosa plants also occur in Mexico
(Oaxaca westward along the Sierra Madre del Sur into Jalisco). One
such Oaxaca collection is known to be tetraploid (n = 24, Melchert,
unpublished).
In brief, a modern analysis of section Greenmanii, designed to
ascertain the relationships between B. reptans, B. holwayi, and the
polyploid complex known as B. squarrosa, is sorely needed.
Bidens steyermarkii Sherff, Am. Journ. Bot. 31: 278. 1944.
Known only from the type; Huehuetenango, around bluffs at
an altitude of 1,900 m., rocky slopes above La Libertad, on Cerro
Pueblo Viejo, Aug. 20, 1942, Steyermark 50998 (type, F, isotypes, G,
US).
A perennial herb, slender, about 3.5 dm. tall, the stems prostrate or procumbent,
very delicate, glabrous, about 2 dm. long, the branches erect or decumbent, very
slender, subquadrangulate, minutely grooved, becoming purple, under 2 mm. thick,
glabrate or moderately puberulent, ramulose, the ultimate branchlets suberect,
subcapilliform, ending in peduncles, these glabrous or toward the apex sparsely
hispidulous, to 1 or 1.5 dm. long, monocephalous; leaves petiolate (the petioles
narrow but margined, weakly hispidulous, most often 7-13 mm. long), 3-5.5 cm. long
including the petiole, the blade very membranous, principal leaves pinnate or more
often bipinnatifid, the lateral leaflets one or two pairs, the ultimate segments more or
less linear-oblong, very sharply apiculate, a little paler below, margins minutely and
MELCHERT: FLORA OF GUATEMALA 213
antrorsely ciliolate, glabrate to sparsely hispidulous; heads radiate, at anthesis about
2.5 cm. wide and 5-7 mm. tall or sometimes only about 1.5 cm. wide and about 5 mm.
tall; phyllaries more or less whitish hispidulous, the outer 10-12, narrowly linear,
finally 4-4.5 mm. long, sharply indurate-apiculate, scarcely longer than the inner
lanceolate ones; ligulate florets short, about 5, white, to 13 mm. long, to 8 mm. wide,
the limb widely oblong or obovate, apex irregularly 2- or 3-denticulate; disc florets
about 30, surpassing the supernally orange-colored and twisted palae, the corolla
orange with erect lobes; submature achenes small, reddish black, linear-clavate,
exalate, subquadrangulate, 3-4.5 mm. long, 2-grooved on each face, glabrous or
supernally very sparsely setulose, exaristate.
Bidens triplinervia HBK. Nov. Gen. & Sp. 4: 231. 1820. B.
crithmifolia HBK. torn. cit. 234. B. delphinifolia HBK. loc. cit. B.
humilis HBK. loc. cit. B. glaberrima DC. Prodr. 5: 601. 1836. B.
canescens Bertol. Novi Comm. Bonon. 4: 431. 1840 (type from
Sacatepequez, Velasquez s.n.) B. artemisiaefolia Poepp. & Endl.
Nov. Gen. & Sp. 3: 49. 71843. B. consolidaefolia Turcz. Bull. Soc.
Nat. Mosc. 24: 185. 1851. B. humilis var. tenuifolius Sch.-Bip.
Griseb. Abh. Ges. Wiss. Gott. 24: 198. 1879. B. attenuata Sherff, Bot.
Gaz. 61: 495. 1916. B. triplinervia var. macrantha (Wedd.) Sherff,
op. cit. 80: 383. 1925. B. triplinervia var. mollis (Poepp. & Endl.)
Sherff, tom. cit. 384. B. triplinervia var. macrantha f. octoradiata
Sherff, op. cit. 92: 203. 1931. B. triplinervia var. typica Sherff,
Brittonia 6: 341. 1948. B. triplinervia var. eurymera Sherff, Am.
Journ. Bot. 38: 66. 1951.
High, wet-montane plants, mostly at 3,000-4,000 m. but
occuring as low as 2,300 m., common near or at the summit of
volcanoes, in meadows above timberline or on open wooded slopes;
in sandy soil among grass hummocks in open pine-juniper forests;
Chimaltenango; Huehuetenango; Quezaltenango; Sacatepequez;
Totonicapan. Widespread and common from Chihuahua and Nuevo
Leon, Mexico, southeastward through Mexico and Central America
well into South America.
Lax, procumbent or sometimes ascending, perennial herbs with several stems
radiating from a thickened, woody, central rootstock, the branches basically terete,
but multiribbed and often somewhat angulate, especially when pressed, each
normally terminated by showy, solitary heads on elongate, upturned peduncles; stem
(and leaf) pubescence exceedingly variable, densely villous-pubescent or canescent to
virtually glabrous; leaves (in ours) normally much dissected, bi- or even
tripinnatisect, rarely trifoliate (these as well as undivided leaf- forms known
throughout Mexico), rather small, with petiole included only 1.2-5 (-7) cm. long, the
ultimate segments short, narrow, linear to linear-sublanceolate, pubescence variable
between and within populations, canescent to variously scattered pubescent or
essentially glabrous, sometimes pubescent above, glabrous below; heads radiate, quite
showy, to 5 (6) cm. across, usually solitary on elongate, ascending or upturned
214 FIELDIANA: BOTANY, VOLUME 24
peduncles; involucre with a conspicuous covering of elongate, multicellular hairs,
these usually forming a dense tomentum near the peduncle (in bud this tomentum
will sometimes virtually hide the inner phyllaries); outer phyllaries 8-12 (-13), linear
to linear -subspathulate, somewhat shorter to subequal with the inner, 3.8-7 mm. long,
0.8-1.5 mm. wide, ciliate, green with black middle (and usually marginal), longitudinal
lines, the inner phyllaries lanceolate, commonly drying glassy black, their chaffy
margins nearly obsolete, 5-7 mm. long, 2-3 mm. wide, the tips rounded and short-
pubescent; ray florets mostly 5 or 6, sometimes 8, rarely even 10, their ligules
proximally deep golden-yellow, paler yellow distally (sometimes becoming whitish in
age or when pressed), oblong-elliptic, (12-) 18-25 mm. long, to 13 mm. wide, gradually
narrowed over the apical half to an obtuse or rounded, obscurely 3-denticulate apex;
disc florets numerous, their corollas yellow or orangish, 4-6 mm. long; anthers black,
2-3.7 mm. long, their terminal appendages triangular, about 0.5 mm. long; style
branches about 2 mm. long; palae narrowly linear above, the central ones to 9 mm.
long, their apices marked with a black triangular spot, the body with yellowish or
amber and blackish, longitudinal stripes; achenes somewhat dimorphic, a few of the
short, 4-5 mm. long, obcompressed-tetragonal peripheral ones commonly yellow or
rubrocastaneous and awnless, the remainder black, linear-tetragonal, gradually
narrowed above, but never rostrate, sparsely antrorse-hispid, the central ones very
narrow, to 9 (11) mm. long; awns 2 (3), erect, short, to 2 (4) mm. long, retrorsely
barbed supernally; chromosome number (of ours), n = 12, 24, 36.
Throughout its wide range, B. triplinervia (sensu lato) is
extremely heterophyllous. All but a very few of the many
Guatemalan collections have small, bipinnatisect leaves with
narrowly lanceolate or oblanceolate segments. Such plants have
typically been referred to var. macrantha (Sherff, 1955). Similar
plants with eight rays rather than the usual five, six (or seven) were
distinguished as var. macrantha from octoradiata. B. canescens
Bertol. is a name applied to plants of the var. macrantha type from
Volcan de Agua which have 8-10 rays and dense, whitish-pubescent
leaves, upper stems, and phyllaries. Numerous collections seen from
the type area show that canescent and essentially glabrous or
scattered pubescent plants occur together (J. R. Johnston 815, US),
as do plants with five and eight rays. Accordingly, B. canescens is
treated here simply as a highly pubescent form of B. triplinervia.
BORRICHIA Adanson
Erect, branching shrubs of seacoasts, glabrous or more or less sericeous with
whitish hairs; leaves opposite, sessile or short-petiolate, the blades carnose, usually
narrow, the margins entire or denticulate; heads radiate, terminal, long-pedunculate;
involucres hemispheric; phyllaries more or less unequal, imbricate in 2-3 series, the
inner ones coriaceous; receptacle convex, paleaceous, the pales rigid, enclosing the
disc flowers; ray flowers fertile, the ligules yellow; disc flowers perfect, the corollas
tubular, the limb 5-dentate; style branches elongated, hispidulous; anthers dark-
colored, entire or minutely sagittate at the base; achenes of the ray flowers trigonous,
those of the disc flowers tetragonous; pappus a short, dentate crown.
NASH: FLORA OF GUATEMALA 215
Two species, both in tropical America, with one in Central
America.
Borrichia arborescens (L.) DC. Prodr. 5: 489. 1836. Buph-
thalmum frutescens L. Syst. ed. 10. 1227. 1759. Borrichia argentea
DC. 5: 489. 1836. Figure 55.
British Honduras, beaches or dune areas bordering mangrove
swamps. Southern Florida; Mexico (Yucatan); West Indies.
Branching shrubs to about 1 m. high, green and glabrous or densely white-
sericeous; leaves thick and fleshy, narrowly oblanceolate or spathulate, mostly 4-10
cm. long, acute or obtuse and cuspidate, attenuate to the base, sessile or with a
petioliform base, the margins entire; heads solitary on long, stout peduncles;
involucres 1-1.5 cm. high; phyllaries closely appressed, stria te, obtuse or acute, the
outer ones usually mucronulate, the inner ones rounded and more or less ciliolate;
ray flowers commonly 12-20, the ligules yellow, 6-9 mm. long; disc flowers numerous;
achenes about 3 mm. long; pappus crown usually less than 0.5 mm. high.
One of the characteristic strand plants in some parts of the
Caribbean region, but apparently rare on the mainland of North
America.
CALEA Linneaus
References: B. L. Robinson and J. M. Greenman, Revision of
the Mexican and Central American species of the genus Calea,
Proc. Amer. Acad. 32: 20-30. 1896; S. F. Blake in Standl. Contr. U.
S. Nat. Herb. 23: 1591-1596. 1926.
Shrubs, small trees, or perennial herbs, erect or rarely subscandent; leaves
opposite, or the uppermost ones sometimes alternate, sessile or petiolate, not lobate,
usually ovate, triangular-ovate, or lanceolate, sometimes elliptic, the margins serrate
or dentate, rarely entire, usually more or less pubescent; inflorescences cymose, often
corymbose, sometimes umbelliform; heads radiate or discoid; involucres ovoid,
cylindrical, campanulate, or hemispheric; phyllaries 2-several-seriate, usually gradu-
ate, scarious or membranaceous or the outer ones herbaceous or with herbaceous tips;
receptacles small, convex or almost flat; pales scarious, concave, rigid or thin and
hyaline; ray flowers (when present) fertile, the ligules yellow, white, or rarely pink,
entire or denticulate at the apex, sometimes inconspicuous and/or only one or two
present; disc flowers fertile, usually yellow, the corolla limb deeply 5-cleft; anthers
shallowly sagittate at the base, appendaged at the apex; style branches obtuse to
truncate; achenes 4-5-angulate, usually pubescent or ciliate along the angles; pappus
scales 4-20, subequal, usually fimbriate or ciliolate.
About 90 species, all American and chiefly in tropical regions.
Of the 16 species treated here, 14 are in Guatemala; 2, which occur
in British Honduras and in Chiapas, Mexico, may be expected in
Guatemala.
216 FIELDIANA: BOTANY, VOLUME 24
Plants herbaceous; inflorescences terminating elongated peduncles, these mostly 4-30
cm. long.
Heads discoid; leaves sessile (plants of higher elevation, 1,600-2,250 m.).
Achenes without pappus C. scabra.
Achenes crowned with fimbriate pappus scales C. scabra var. peduncularis.
Heads radiate; leaves on winged petioles (plants of low elevation, near sea level to
900 m.) C. megacephala.
Plants shrubs or trees (except C. integrifolia which may be either herbaceous or
suffrutescent); inflorescences not terminating elongated peduncles, often leafy,
often shorter than the leaves.
Leaves penninerved C. guatemalensis.
Leaves 3-nerved or triplinerved.
Ray flowers with bright yellow ligules 3-5 mm. long.
Leaves densely pilosulous beneath; petioles 0.5-4 cm. long; inflorescences often
large, convex or flat, corymbiform panicles; pappus considerably shorter
than the achenes C. skutchii.
Leaves hispidulous or scabrous beneath; petioles usually less than 0.5 cm. long;
inflorescences small, umbelliform panicles; pappus longer than the
achenes C. urticifolia.
Ray flowers with white ligules, or the heads discoid.
Leaves linear, linear-lanceolate, oblong-lanceolate, elliptic, or ovate-elliptic.
Outermost phyllaries green, more or less herbaceous.
Leaf blades narrowly linear, acuminate, 1-2 mm. wide; involucres about 5
mm. high C. fluviatilis.
Leaf blades elliptic, broadly elliptic, or oblong-lanceolate, mostly 1.5-4 cm.
wide; involucres about 10mm. high C. longipedicellata.
Outermost phyllaries membranaceous C. integrifolia.
Leaves broadly ovate, triangular-ovate, lance-ovate, or suborbicular.
Lower leaf surfaces commonly densely tomentose with more or less matted
or crisped hairs.
Outermost phyllaries herbaceous throughout or at the apex.
Heads sessile or if short-pedicellate, the pedicels less than 0.5 cm. long.
C. hypoieuca.
Heads on pedicels 0.5-2 cm. long C. trichotoma.
Outermost phyllaries membranaceous, very short, suborbicular.
C. pringlei.
Lower leaf surfaces hispidulous, sparsely or moderately pilose, or glabrous
and glandular-punctate, never tomentose with matted or crisped hairs.
Leaves usually more or less hispidulous or pilose beneath (sometimes the
indument confined to costae and/or veins); heads sessile or on
pedicels mostly less than 5 mm. long.
Outermost phyllaries herbaceous throughout; pappus longer than the
achenes C. crassifolia.
Outermost phyllaries membranaceous, short, orbicular; pappus shorter
than the achenes C. zacatechichi.
Leaves always glabrous beneath; heads on pedicels 5-20 mm. long.
C. peckii.
Calea crassifolia Standl. & Steyerm. Field Mus. Bot. 23: 256.
1947.
NASH: FLORA OF GUATEMALA 217
Known only from the type, Alta Verapaz, along knife-edge of
limestone ridge, Cerro Chinaja, between Finca Yalpemech and
Chinaja, 150-700 m., Steyermark 45627.
A shrub, 1.5-2.5 m. high, the branches stout, terete, very densely hispidulous with
short, spreading, brownish hairs; leaves sessile or nearly so, the blades coriaceous or
subcoriaceous and rigid, broadly triangular-ovate or rounded-ovate, 5-7 cm. long, 3.5-
6 cm. wide, obtuse and obscurely mucronulate at the apex, shallowly cordate or
rounded at the base, very scabrous above, densely hispidulous beneath with short,
spreading, rough hairs, the margins coarsely crenate-serrate with mucronulate teeth,
3-nerved, the veins prominent and conspicuous beneath and laxly reticulate;
inflorescences small, 1.5-2.5 cm. broad, sessile at the ends of the branches; heads few,
apparently discoid, closely aggregate and sessile or on petioles about 1 mm. long;
involucres 6-7 mm. high; phyllaries 3-seriate, the outer ones herbaceous, green, oblong
or lance-oblong, acute or subacute, usually spreading, mostly about equalling or
slightly longer than the inner ones, densely hispidulous, ciliate, the inner phyllaries
stramineous, oblong, obtuse or subacute, ciliate, glabrous or glabrate; corollas yellow,
4-4.5 mm. long; achenes narrow, 2-3-angulate, about 2.6 mm. long, sparsely
pubescent; pappus dissected into many setaceous, long-attenuate scales about 4 mm.
long.
Calea fluviatilis Blake, Journ. Wash. Acad. Sci. 22: 385. 1932.
Known only from the type, British Honduras, growing on
stones in Rio Privation, Mountain Pine Ridge, El Cayo District,
Bartlett 11790.
A small shrub, about 25 cm. high, the several, erect, branching stems arising from
a thick, woody caudex, minutely hispidulous to glabrate; leaves opposite, sessile or on
petioles 1 mm. long, the blades narrowly linear to linear-lanceolate, 1.3-2.8 cm. long,
1-2 mm. wide, acuminate, attenuate to the base, the margins subentire or very
remotely denticulate and more or less revolute, triplinerved, glandular-punctate on
both surfaces, glabrous above, glabrous or sparsely strigillose beneath; heads discoid,
on slender peduncles 8-12 mm. long, disposed in terminal clusters of 3-7; phyllaries 3-
4-seriate, the outermost ones green, coriaceous-herbaceous throughout or at least
above the middle, triangular, 3-5 mm. long, the inner ones yellowish and
longitudinally striate, sometimes tipped with purple; pales oblong, yellowish, striate;
corollas yellow, glabrous, 4-5 mm. long; achenes more or less hirsutulous, about 2
mm. long, pappus scales about 20, longer than the achenes.
Calea guatemalensis Donn.-Sm. Bot. Gaz. 23: 9. 1897. Lluvia
de oro (Huehuetenango).
Damp or wet thickets, mixed forest, or pine-oak forest, 2,100-
3,700 m.; Huehuetenango (type collected between San Martin and
Todos Santos, Nelson 3624); El Quiche.
Shrubs or trees, 2-6 m. tall, the branches dark brown, striate, glabrous or very
sparsely short-hirsute; leaves on stout petioles 1-2 cm. long, the blades rather thick,
ovate to rather narrowly lanceolate, mostly 7-16 cm. long and 2.5-5.5 cm. wide, acute
to long-acuminate, acute at the base and sometimes narrowly decurrent on the
petiole but not to its base, the margins entire, subentire, or rarely some of them
218 FIELDIANA: BOTANY, VOLUME 24
serrate, glabrous above, much paler beneath and glabrous or sometimes hirsute along
the veins, penninerved; inflorescences mostly 10-26 cm. broad, corymbose, the
numerous bracts similar to the leaves but much smaller, sessile; heads radiate, on
rather stout, usually pubescent pedicels; involucres hemispheric, about 4.5 mm. high
and broad; phyllaries 2-3-seriate, oblong, membranaceous, obtuse, yellowish, ciliate;
ray flowers commonly 7-8, the ligules yellow, 6-8 mm. long, spreading, conspicuous;
disc flowers numerous, yellow, about 6 mm. long; achenes about 3 mm. long, oblong,
ciliate on the angles; pappus scales 4, triangular-linear, lacerate, those of the ray
flowers about 1 mm. long, those of the disc flowers about 2 mm. long.
Easily recognized by the penninerved leaves, those of other
local species being three-nerved or triplinerved. This may not be
distinct from C. orizabaensis Klatt of Mexico (Veracruz), but
authentic material of that species is not now available for study. If
the two should prove to be synonymous, then the name C.
orizabaensis (Leopoldina 23: 145. 1887) would, of course, take
precedence.
Calea hypoleuca Robins. & Greenm. Proc. Amer. Acad. 32: 24.
1896. C. sororia Blake, Contr. U. S. Nat. Herb. 22: 645. 1924 (type
collected near Nenton, Huehuetenango, Nelson 3544).
In Guatemala known only from Huehuetenango, 900-1,200 m.
Mexico (Oaxaca and Chiapas).
Branching shrubs to at least 1 m. tall, the branches pubescent; leaves short -
petiolate or subsessile, the blades broadly ovate or suborbicular, mostly 3-5 cm. long,
acute or obtuse, obtuse to rounded or sometimes subcordate at the base, the margins
coarsely crenate-serrate, scabrous above, densely tomentose beneath with matted or
crisped hairs; inflorescences small, cymose, dense, the heads obscurely radiate, short-
pedicellate or subsessile; involucres narrowly campanulate, 5-6 mm. high; phyllaries
closely appressed, broad, obtuse, the outermost ones with herbaceous, pubescent tips,
the others essentially glabrous except usually more or less pubescent at the apex; ray
flowers inconspicuous, but at least one or two always present, sometimes more, the
ligules white, scarcely 2 mm. long; disc flowers about 10; achenes 2-3 mm. long, black
and shining when mature, usually foveolate, pubescent or at least ciliate on the
obscure angles; pappus scales about 10, obtuse or acute, 1-1.5 mm. long.
Calea integrifolia (DC.) Hemsl. Biol. Centr. Am. Bot. 2: 205.
1881. Allocarpus integrifolius DC. Prodr. 5: 676. 1836. C. integrifolia
var. dentata Coulter, Bot. Gaz. 20: 51. 1895 (type from Nebaj, El
Quiche, Heyde & Lux 4506). Cola de zorro (Huehuetenango); sac-
kilocuj, subub (Quecchi, Alta Verapaz).
Damp or wet thickets or open forest, common in pine-oak
forest, 450-2,700 m.; Alta Verapaz; Chimaltenango; Guatemala;
Huehuetenango; Jalapa; Peten; El Progreso; Quezaltenango; El
Quiche; Sacatepequez; San Marcos; Santa Rosa; Solola; Suchitepe-
quez. Southern Mexico; Honduras; El Salvador; Nicaragua.
NASH: FLORA OF GUATEMALA 219
Herbaceous or suffrutescent plants, the stems 1-3 m. long, erect or weak and
reclining or scrambling on other plants, terete, glabrous to densely hirsute, usually
sparsely branched; leaves on petioles 1-10 mm. long, the blades mostly oblong-
lanceolate, sometimes lance-ovate or linear-lanceolate, commonly 6-12 cm. long,
acuminate to long-acuminate, rounded to very obtuse at the base, often lustrous and
usually rugose, the margins coarsely serrate to almost entire, usually hirsute and
scabrous on the upper surface, rarely smooth and glabrous, the lower surface rather
densely pilose to almost glabrous; inflorescences leafy at the base and often shorter
than the leaves; heads usually very numerous, short-pedicellate, disposed in small,
usually dense, cymose panicles at the ends of the branches, 15-20-flowered; involucres
broadly campanulate, 3-4 mm. high; phyllaries pale, membranaceous, striate, oblong
or oval, obtuse or rounded at the apex, ciliate, glabrous or nearly so; ray flowers
commonly 5-8, the ligules white, mostly 3-5 mm. long, tridentate; disc flowers yellow;
ray achenes without pappus; disc achenes scarcely more than 1 mm. long, black,
pubescent; pappus scales white, about 3 mm. long.
Although this species exhibits considerable variation in pubes-
cence and leaf form the variation is so gradual that the forms
scarcely seem to merit special designation.
Calea longipedicellata Robins. & Greenm. Proc. Amer. Acad.
32: 28. 1896.
In savannas or pine forest, near sea level to 1,000 m.; Alta
Verapaz; Huehuetenango. British Honduras; Honduras; Mexico
(Chiapas).
Shrubs 1-4.5 m. high, the branches glabrous or sparsely pilose; leaves on very
short petioles or subsessile, the blades elliptic to ovate-elliptic or oblong-lanceolate,
mostly 3-8 cm. long and 1.5-4 cm. wide, acute, at the base obtuse or acute, the
margins remotely and obscurely denticulate or sometimes with a few large teeth,
glabrous or nearly so, triplinerved, often lustrous; heads discoid, usually 2-4 at the
ends of the branches (rarely solitary), the pedicels naked, 3-6 cm. long, sparsely
puberulent; involucres about 1 cm. high, 1-1.5 cm. broad, hemispheric; phyllaries few,
the outer ones herbaceous, appressed, oblong or elliptic, very obtuse or rounded at
the apex, usually somewhat exceeding the inner ones, these coriaceous, pale,
glabrous; corollas orange-yellow; achenes about 2 mm. long, glabrous; pappus scales
about 20, linear-attenuate, 5-6 mm. long.
Calea megacephala Robins. & Greenm. Proc. Amer. Acad. 32:
21. 1896. Tonalanthus aurantiacus Brandegee, Univ. Calif. Publ.
Bot. 6: 75. 1914.
Not reported from Guatemala, but included here because the
type collection of Tonalanthus aurantiacus (Purpus 7002) was
made in nearby Chiapas, Mexico.
Erect perennials, the simple, thick stems to almost 1 m. high, pilose with
multiseptate hairs; leaves opposite, on winged petioles, the blades thin, broadly ovate
or triangular-ovate, mostly 7-10 cm. long, 6-8 cm. wide, the upper ones acute to
acuminate, the lowermost ones rounded at the apex, all truncate to subcordate at the
220 FIELDIANA: BOTANY, VOLUME 24
base and then abruptly decurrent on the petiole, the margins coarsely crenate-
dentate, rather sparsely pubescent on both surfaces but more or less pilose on the
veins beneath; peduncles 1-3, axillary, to about 30 cm. long; heads solitary, about 15
mm. high and 15-20 mm. broad (excluding the ray flowers); phyllaries 2-3-seriate,
lance-oblong, obtuse, about 1 cm. long, striate, pubescent; pales scarious, acuminate;
ray flowers 15-20, the ligules orange, about 8 mm. long; disc flowers numerous, yellow
to orange; achenes 4-5 mm. long, pubescent; pappus scales numerous, fimbriate,
almost as long as the body of the achene.
Calea peckii Robinson, Proc. Amer. Acad. 44: 624. 1909.
Usually in damp or wet, open, pine forest, at or a little above
sea level; Peten. British Honduras (exact type locality not
indicated, Peck 64). Mexico (Yucatan); Honduras; Nicaragua; Costa
Rica.
Erect, arching, or subscandent shrubs, 1-2 m. high, the branches slender, striate,
scabrous or puberulent; leaves on petioles 2-9 mm. long, the blades broadly ovate to
lance-ovate, 2-7 cm. long, acute or acuminate, rounded or obtuse at the base, the
margins subentire to coarsely crenate-serrate, scaberulous above, glabrous beneath
but glandular-punctate, triplinerved from near the base; heads discoid, on puberulent
pedicels mostly 0.5-2 cm. long, disposed in small, sometimes dense cymes, the
inflorescences often appearing umbellate, in the upper leaf axils; involucres cylindric-
campanulate, 5-6 mm. high; outer phyllaries broadly ovate and puberulent, ciliolate,
more or less herbaceous, somewhat squarrose, the middle ones ovate-oblong, reddish-
striate, the innermost narrowly lanceolate, yellowish, acute; disc flowers about 14,
the corollas yellow; achenes narrowly obconic, pubescent, about 2 mm. long; pappus
squamellae 20 or more, narrowly linear, 4-5 mm. long.
Calea pringlei Robinson, Proc. Amer. Acad. 36: 488. 1901. C.
pringlei var. rubida Greenman, Field Mus. Bot. 2: 272. 1907. C.
tejadae Blake, Contr. U. S. Nat. Herb. 22: 646. 1924 (type from
Agua Blanca, Escuintla, Tejada 80). Oregano, San Julian (Es-
cuintla); mosca blanca, tasiscob bianco, vara blanca (Guatemala).
Damp or dry, often rocky, thickets or forest, sea level to 1,500
m.; Escuintla; Guatemala; Izabal. Mexico; Honduras.
Erect shrubs, the stems stout, 1-2.5 m. high, the young branches pilose-
tomentose; leaves short-petiolate, the blades rather thick and rigid, ovate to broadly
ovate, mostly 2-6 cm. long, 1.5-4 cm. wide, acute or acuminate, rounded to cuneate at
the base, triplinerved from near the base, the margins usually coarsely crenate-
serrate, rarely subentire, rugose above and usually densely hispid-pilose, densely
tomentose beneath with matted or crisped hairs; inflorescences terminal,
corymbiform; heads obscurely radiate, disposed in small, aggregate, more or less
umbellate, densely flowered corymbs, the pedicels 2-6 mm. long; involucres 4-5 mm.
high, 2.5-3 mm. broad; phyllaries 5-6-seriate, appressed, the outermost ones very
short, suborbicular, the others oblong or oval, obtuse or rounded at the apex, without
herbaceous tips, pale, ciliate, glabrous; ray flowers commonly 2, rarely more,
inconspicuous, the ligules white, about 2 mm. long; disc flowers 10-14, the corollas
glabrous, about 3.5 mm. long; achenes blackish, first angulate but becoming
NASH: FLORA OF GUATEMALA 221
subterete, 2.5-3 mm. long, shining, usually with 2-4 lines of pubescence; pappus scales
8-10, subequal, 1-1.5 mm. long.
Calea scabra (Lag.) Robinson, Proc. Amer. Acad. 44: 625.
1909. Calydermos scaber Lag. Gen. & Sp. Nov. 25. 1816. Calydermos
longifolius Lag. I.e. Calydermos peduncularis var. epapposa HBK.
ex DC. Prodr. 5: 669. 1836. Calea scabra var. longifolia Robinson,
Proc. Amer. Acad. 44: 625. 1909. Cola de zorro (Huehuetenango);
lengua de gato (Solola).
Open pine or oak forest, 1,600-2,250 m.; Baja Verapaz;
Chimaltenango; Huehuetenango; Jalapa; Solola. Mexico.
Erect, perennial herbs, the stems mostly simple, to about 1 m. tall, hirsute and
scabrous; leaves rather remote, sessile, ovate to oblong- lanceolate or rarely almost
linear, mostly 4-10 cm. long, acute or acuminate, rounded or subcordate and
somewhat clasping at the base, the margins serrate-dentate, usually remotely so,
hirsute, scabrous, 3-5-nerved; inflorescences terminal, on peduncles mostly 5-20 cm.
long, more or less corymbiform, dense, usually 3-5 cm. broad; heads discoid, sessile or
short-pedicellate; involucres 4-6 mm. high; phyllaries glabrous or nearly so,
stramineous, more or less striate, membranaceous, rounded at the apex, ciliate;
corollas yellow, the tubes pubescent; achenes cuneate, almost glabrous or
conspicuously pubescent, scarcely 2 mm. long; without pappus.
Calea scabra var. peduncularis Robinson, Proc. Amer. Acad.
44: 625. 1909. C. peduncularis HBK. Nov. Gen. & Sp. 4: 295, t. 408.
1820. C. peduncularis var. livida Robins. & Greenm. Proc. Amer.
Acad. 32: 24. 1896. C. purpusii Brandg. Univ. Calif. Publ. Bot. 10:
419. 1924. Sojoyan (Huehuetenango).
Open, damp forest or on dry, rocky slopes, 1,400-2,000 m.;
Huehuetenango; El Quiche; Totonicapan. Mexico.
Differs from the typical variety in its achenes being crowned
with fimbriate pappus scales, about one-half as long as or equalling
in length the body of the achene.
Calea skutchii Blake, Journ. Wash. Acad. Sci. 24: 438. 1934.
Bilil (San Marcos). Figure 56.
Damp or wet forest or thickets, often in pine, pine-oak, or oak
forest, sometimes in Alnus or Cupressus forest, often in sandy soil,
1,500-3,800 m.; Chimaltenango; Huehuetenango; Quezaltenango; El
Quiche; San Marcos; Solola; Totonicapan. Mexico (Chiapas).
Shrubs or small trees, 1.5-8 m. tall, the branches pilose with brownish, somewhat
matted hairs; leaves commonly opposite but sometimes the upper ones alternate, on
petioles 0.5-4 cm. long, the blades ovate to ovate-lanceolate or oblong-lanceolate,
mostly 8-17 cm. long and 3.5-7 cm. wide, acuminate, rounded and then rather
abruptly cuneate at the base, or sometimes attenuate to the base, the margins
222 FIELDIANA: BOTANY, VOLUME 24
subentire or serrate, triplinerved far above the base, scaberulous or glabrate above,
usually densely pilosulous beneath; heads radiate, very numerous, disposed in large,
convex or almost flat, corymbiform panicles, the pedicels slender, mostly 4-10 mm.
long; involucres campanulate, 5-8 mm. high, 4.5-7 mm. broad; outermost phyllaries
herbaceous, pubescent, oblong or oblong-ovate, obtuse or subacute, the inner ones
narrowly oblong to cuneate-obovate, obtuse or apiculate, yellowish; ray flowers
commonly 8-9, the ligules bright yellow, mostly 3-5 mm. long; disc flowers about 30,
the corollas to about 5 mm. long; achenes narrowly cuneate, 3-3.5 mm. long, dark
brown, hirsutulous or ciliate on the angles; pappus squamellae 6-10, lanceolate,
acuminate, 0.5-1.5 mm. long.
Showy shrubs, common in the mountains of the Occidente,
mostly at rather high elevations.
Calea trichotoma Donn.-Sm. Bot. Gaz. 13: 289. 1888.
Usually in damp or wet, pine forest, near sea level to 1,450 m.;
Alta Verapaz (type from Coban, Tuerckheim 1353}; Huehue-
tenango; Izabal; Peten. Southern Mexico; British Honduras;
Honduras.
Erect shrubs, 1-2 m. high or sometimes the stems arching or scandent, the
branches densely sordid-pilose with short or long hairs; leaves on very short petioles,
the blades thick, ovate, triangular-ovate, or lance-ovate, mostly 2-6 cm. long, acute or
obtuse, subcordate to rounded or obtuse at the base, the margins coarsely crenate
crenate-serrate, or subentire, scabrous-pilose or densely and softly short-pilose on the
upper surface, densely tomentose beneath with matted, multiseptate, grayish or
brownish hairs; heads discoid, on densely pubescent pedicels mostly 0.5-2 cm. long,
disposed in small, umbellate cymes or panicles, these terminal or in the upper leaf
axils; involucres 5-6 mm. high; outer phyllaries short, herbaceous throughout or at
the apex, somewhat squarrose, densely pubescent, the inner ones oblong, oval, or
ovate-oblong, obtuse or rounded at the apex, reddish-striate, the innermost
lanceolate, yellowish, glabrous, ciliate; disc flowers 11-18, yellow; achenes 2-2.5 mm.
long, pubescent; pappus scales 20 or more, linear, 4-5 mm. long.
Calea urticifolia (Mill.) DC. Prodr. 5: 674. 1836. Solidago
urticifolia Mill. Gard. Diet. ed. 8, no. 30. 1768. C. axillaris DC.
Prodr. 5: 673. 1836. C. axillaris var. urticaefolia Robins. & Greenm.
Proc. Amer. Acad. 32: 27. 1896. C. urticifolia var. axillaris Blake,
Contr. Gray Herb. 52: 57. 1917. Mosca amarilla (Guatemala).
Damp or dry thickets, sometimes in oak-pine forest, 100-1,900
m.; Alta Verapaz; Chiquimula; Escuintla; Guatemala; Huehue-
tenango; Peten; Santa Rosa; Zacapa. Mexico; British Honduras and
Honduras; Nicaragua; Costa Rica; Panama.
Erect shrubs, 1-2 m. high, the branches sparsely or densely pilose; leaves short -
petiolate, the blades ovate to lance-oblong or lance-elliptic, mostly 4-12 cm. long and
1-6 cm. wide, acute or acuminate, acute or obtuse at the base, triplinerved, the
margins serrate, scabrous above, often rugose, the veins conspicuous, thinly or
NASH: FLORA OF GUATEMALA 223
densely hispidulous or scabrous beneath, often lustrous on both surfaces; heads
numerous, radiate, disposed in small, umbelliform panicles, these often shorter than
the leaves, the pedicels slender, mostly 0.5-2.5 cm. long; involucres 6-7 mm. long;
outermost phyllaries oblong or lance-oblong, herbaceous at the apex or throughout,
acute or obtuse, puberulent, often equalling the inner ones, the inner phyllaries
oblong or obovate-oblong, membranaceous or sometimes with herbaceous tips,
glabrous or nearly so, pale yellowish; ray flowers 3-8, the ligules bright yellow,
spreading, 4-5 mm. long; disc flowers commonly 20-26 (rarely 8-14); achenes about 2.5
mm. long, short-pilose; pappus scales 3-4 mm. long.
Calea zacatechichi Schlecht. Linnaea 9: 589. 1834. C. nelsonii
Robins. & Greenm. Proc. Amer. Acad. 32: 25. 1896. C. zacatechichi
var. macrophylla Robins. & Greenm. torn. cit. 26 (type from Coban,
Alta Verapaz, Tuerckheim 1345). Amargoso (Guatemala); canilla
de zanate (Jutiapa); ore/a de conejo and vara negra (Jalapa); vara
blanca (Chimaltenango).
Damp or dry, brushy slopes, often in pine-oak forest, 65-2,500
m. (most common at elevations above 1,500 m.); Alta Verapaz; Baja
Verapaz; Chimaltenango; Chiquimula; Escuintla; Guatemala;
Huehuetenango; Izabal; Jalapa; Jutiapa; Peten; Quezaltenango;
Sacatepequez; Santa Rosa; Solola; Zacapa. Southern Mexico;
British Honduras; Honduras; El Salvador; Nicaragua; Costa Rica.
Erect or sometimes arching shrubs, 0.5-3 m. high, the branches glabrous,
puberulent, or pilose; leaves short-petiolate, the blades thick, often very rugose and
lustrous, broadly ovate to lance-ovate or triangular-ovate, mostly 3-7 cm. long, acute
or acuminate, obtuse or rounded to cuneate at the base, triplinerved from near the
base, the margins usually coarsely dentate or crenate-serrate, sometimes subentire,
more or less scabrous on the upper surface, conspicuously or obscurely glandular-
punctate beneath and usually sparsely to moderately pilose beneath (rarely
essentially glabrous); heads obscurely radiate, very numerous, sessile or short-
pedicellate, disposed in small, dense cymes; involucres narrowly campanulate, 4-5
mm. high; phyllaries closely appressed, broad, very obtuse or rounded at the apex,
subcoriaceous, without herbaceous tips, glabrous, the outer ones short and
suborbicular, the inner ones oval to oblong, usually ciliate; ray flowers usually 2,
rarely more, inconspicuous, the ligules white, scarcely 2 mm. long; disc flowers
commonly (8-) 10-14; achenes 2-3.5 mm. long, black, shining, subterete, more or less
pubescent; pappus scales about 12, about 1 mm. long.
The achenes of this species and of some related ones often
appear to be angulate and ciliate on the angles until quite mature.
In age they become subterete, the "cilia" or pubescence then is
disposed in vertical lines.
The specific name is derived from a Nahuatl vernacular name
used in Mexico, based on the Nahuatl zaca-chichic, "bitter grass"
or "bitter weed."
224 FIELDIANA: BOTANY, VOLUME 24
Reported from Guatemala as C. salmeaefolia Hemsl., a species
of Mexico, which differs in having pedicels considerably longer than
the heads.
CALYPTOCARPUS Lessing
Reference: R. McVaugh and N. Smith, Calyptocarpus vialis
and C. wendlandii (Compositae), Brittonia 19: 268-272. 1967.
Low annuals or perennials, usually procumbent or prostrate, the pubescence of
appressed or spreading hairs; leaves opposite, petiolate, the blades thin, the margins
crenate-serrate; heads heterogamous, radiate, solitary or clustered, some of them
long-pedicellate, others sessile; phyllaries about 5, the outer ones herbaceous;
receptacle paleaceous, the pales flat or somewhat concave, acute, not carinate; ray
flowers 5-8, pistillate, fertile, uniseriate, the ligules yellow, oblong; disc flowers
hermaphrodite, tubular, the limb 4-5-dentate; stamens 4 or 5, the anthers sagittate,
conspicuously appendaged at the apex; achenes of ray and disc flowers similar,
compressed, usually more or less tuberculate when mature, sometimes with short,
irregular, marginal projections but not winged, those of the ray flowers flatter;
pappus of 2 stout, hard, spinelike awns.
The genus consists of two species, with one in Guatemala.
Calyptocarpus wendlandii Sch. Bip. Bot. Zeit. 24: 165. 1866.
Cachillo (Quezaltenango); cachito bravo (Guatemala and Quezal-
tenango); sitit q'en (Alta Verapaz, Quecchi). Figure 57.
Damp thickets or forest or on open banks and in fields, often a
weed in waste or cultivated ground, 300-2,000 m.; Alta Verapaz;
Chimaltenango; Chiquimula; Escuintla; Guatemala; Huehue-
tenango; Jalapa; Jutiapa; Quezaltenango; Retalhuleu; Sacatepe-
quez; San Marcos; Suchitepequez. Southern Mexico; El Salvador;
Honduras; Nicaragua; Costa Rica.
Plants annual or perennial from a small, hard, woody base, the stems branched,
slender, mostly prostrate of creeping, often rooting at the nodes, to about 35 cm. long,
hirsute with white, spreading or ascending hairs; leaves on long, slender petioles, the
blades ovate or rhombic-ovate, mostly 2-6 cm. long, acute or obtuse, cuneate or
obtuse at the base and decurrent on the petiole, more or less strigose, more densely so
beneath, the margins inconspicuously crenate-serrate; some heads sessile and some on
slender, naked pedicels 1-4 cm. long; phyllaries 6-12 mm. long, appressed, oblong or
obovate-oblong, acute or acuminate, green, densely strigose; pales essentially
glabrous; ray flowers 5-8, the ligules yellow, 3-4 mm. long; disc flowers few; achenes
4.5-6.5 mm. long; pappus of stout, glabrous awns 3-5 mm. long, usually spreading or
reflexed when mature.
In general appearance much like Synedrella, which may be
distinguished by the fact that its flowering heads are never long-
pedicellate, and by it ray achenes which are smooth and have
winglike, lobate margins.
NASH: FLORA OF GUATEMALA 225
CHRYSANTHELLUM L. Richard
Glabrous, annual herbs, erect to prostrate, often diffusely branched; basal leaves
mostly incised-dentate, the middle, cauline ones alternate, variously dissected, the
uppermost ones smaller and usually little dissected; heads heterogamous, radiate,
long-pedunculate, terminal or in the upper leaf axils; involucres campanulate or
hemispheric; phyllaries 1-2-seriate, subequal, membranaceous; receptacle flat; pales
very narrow, flat, scarious; ray flowers pistillate, uniseriate, the ligules spreading,
entire or bidentate; disc flowers hermaphrodite, fertile, the corollas regular, the tube
short, the limb 5-cleft; anthers obtuse and entire at the base; style branches of the
hermaphrodite flowers slender, attenuate into a long subulate appendage; achenes
linear-oblong, glabrous, smooth or nearly so, the outer ones more or less cylindrical,
8-10-sulcate, crowned by an epigynous disk, epappose, the innermost achenes often
much compressed, with 2 membranous margins, epappose or with 2 very minute
apical teeth.
About eight species, in the tropics of both hemispheres, with
two in Guatemala.
Peduncles mostly 3-6 cm. long; heads 4-6 mm. high, 6-10 mm. broad; achenes about 3
mm. long C. americanum.
Peduncles mostly 1-3 cm. long; heads 3-4 mm. high, 3-6 mm. broad; achenes about 2
mm. long C. mexicanum.
Chrysanthellum americanum (L.) Vatke, Bremen Abh. 9:
122. 1885. Anthemis americana L. Sp. PL 895. 1753. C. procumbens
L. Rich, ex Pers. Syn. PL 2: 471. 1807. Figure 58.
Open, grassy, often stony fields and hillsides, 900-1,400 m.;
Escuintla; Huehuetenango; Jalapa; Jutiapa; El Quiche; Santa
Rosa. Mexico; El Salvador; Honduras; Costa Rica; Panama;
Greater Antilles; northern South America.
Ascending, decumbent, or prostrate herbs, often much branched from the base or
above, the stems mostly 5-30 cm. long, striate; leaves on broad petioles, the blades
somewhat succulent, the larger ones mostly 2-5 cm. long, pinnatisect with entire or
dentate lobes or segments, these obtuse or acute, the uppermost leaves sometimes
merely dentate; peduncles slender, mostly 3-6 cm. long, usually solitary, sometimes 2
or 3 in a very lax cyme; heads 4-6 mm. high, 6-10 mm. broad; phyllaries broadly
oblong, obtuse, striate, scarious-margined, about 4 mm. long; pales linear, about 3
mm. long; ray flowers few, often inconspicuous, the ligules about 3 mm. long; disc
flowers yellow, 2.5-3 mm. long; achenes about 3 mm. long, the outer ones more or less
cylindrical to almost trigonous, the inner ones much compressed and very narrowly
winged.
Chrysanthellum mexicanum Greenm. Proc. Amer. Acad. 39:
114. 1903.
Open fields and slopes, 1,000-2,000 m.; El Quiche. Mexico.
Erect or ascending herbs, the stems simple or much branched, 4-20 cm. high,
glabrous or somewhat hirtellous, sometimes purplish; leaves petiolate, the blades
226 FIELDIANA: BOTANY, VOLUME 24
mostly 1-2.5 cm. long, mostly tripinnatisect, the narrow segments acute, apiculate,
glabrous or nearly so; inflorescences laxly cymose, sometimes becoming sub-
corymbose; heads 3-4 mm. high, 3-6 mm. broad, the peduncles mostly 1-3 cm. long;
involucres broadly campanulate; phyllaries ovate-oblong, acute, striate, scarious-
margined, glabrous; pales linear; ray flowers few, inconspicuous, the ligules about 1
mm. long, bidentate; disc flowers yellow, about 1 mm. long; achenes about 2 mm.
long, the outer ones subterete, the inner ones obcompressed, with narrow,
membranous, ciliolate margins.
CLIBADIUM Linneaus
Reference: O. E. Schulz, Beitrage zur Kenntnis der Gattung
Clibadium, Bot. Jahrb. 46: 613-628. 1912.
Shrubs or small trees, rarely herbs, usually scabrous-pubescent; leaves opposite,
petiolate, the blades mostly ovate to lanceolate, often triplinerved, the margins
serrate or crenate; heads small, disciform, few-many-flowered, usually numerous and
disposed in more or less corymbiform panicles; involucres ovoid or subglobose;
phyllaries few, pale, ovate or suborbicular, longitudinally striate; receptacles small,
paleaceous near the margins, sometimes naked near the center; outer pales enclosing
the 3-37 fertile, pistillate flowers, the inner pales narrow or none; pistillate corollas
tubular, the limb 2-4-dentate, the style branches subacute, minutely papillose on the
outer surface; hermaphrodite flowers 3-11, sterile, the corollas short, 5-dentate, the
styles minutely bifid; anthers linear, subsagittate at the base, ovate-appendaged at
the apex; achenes obovoid, somewhat compressed, convex on the outer surface, often
carinate on the inner, at maturity black or dark purple; pappus reduced or none.
Of the more than 25 reported species, two are in Guatemala,
and a third, found in Chiapas, Mexico, is also treated here.
The fruits in this genus often appear baccate, although this is
not at all obvious in dried specimens. Some of the South American
species are said to be employed as fish poisons.
Pistillate flowers 20-30 C. pituieri.
Pistillate flowers 3-6.
Principal leaves ovate to broadly ovate, mostly 12-20 cm. long, 9-16 cm. broad.
C. arboreum.
Principal leaves lanceolate to lance-ovate, mostly 6-10 cm. long, 1.5-5 cm. broad.
C. leiocarpum.
Clibadium arboreum Donn.-Sm. Bot. Gaz. 14: 26. 1889. C.
donnell-smithii Coulter, op. cit. 16: 98. 189 (type from Guatemala,
J. D. Smith 2347). C. oligandrum Blake, Brittonia 2: 342. 1937 (type
from Quezaltenango, Skutch 927). Figure 59.
Damp or wet forest or thickets, near sea level to 2,500 m.; Alta
Verapaz (type from Pansamala, Tuerckheim 929); Chimaltenango;
Escuintla; Guatemala; Izabal; Peten; Quezaltenango; San Marcos;
NASH: FLORA OF GUATEMALA 227
Solola; Suchitepequez. Southern Mexico; British Honduras;
Honduras.
Shrubs or small trees, 2-8 m. high, densely branched, the branches densely pilose
with spreading hairs; leaves petiolate, the blades thin, ovate or broadly ovate, the
principal ones mostly 12-20 (-25) cm. long, 9-16 cm. broad, acuminate or long-
acuminate, usually more or less rounded at the base and rather abruptly contracted
and then cuneate, triplinerved far above the base, the margins serrate, scabrous on
the upper surface, softly and densely short-pilose beneath; panicles small or large,
usually densely flowered and compact, leafy at the base; heads numerous, sessile or
short-pedicellate; phyllaries 3-4, broadly ovate, acute or obtuse, 3-5.5 mm. long, more
or less strigose or glabrate, striate; pistillate flowers 3-6; hermaphrodite flowers 4-11;
corollas 2-3.5 mm. long; anthers blue; achenes blackish, broadly obovoid, about 2.3
mm. long, hirsutulous at the apex.
This species has been reported from Guatemala as C. asperum
DC., certainly a similar species. I have not seen authentic material
of this species, but if the two should prove to be synonymous, then
the name C. asperum (1836) would take precedence.
Clibadium leiocarpum Steetz in Seem. Bot. Voy. Herald; 152.
1852. C. schulzii Blake, Contr. U. S. Nat. Herb. 22: 602. 1924. C.
leiocarpum var. strigosum Blake, Journ. Wash. Acad. Sci. 27: 382.
1937.
Not reported from Guatemala but to be expected there as it
occurs in Chiapas, Mexico, as well as in Costa Rica and Panama.
Shrubs to about 3 m. tall, usually densely branched, the branches striate, densely
tomentose; leaves on petioles 0.3-4 cm. long, the blades thin, lanceolate to lance-
ovate, mostly 6-10 cm. long, 1.5-5 cm. broad, acuminate, almost rounded and then
rather abruptly cuneate at the base, triplinerved above the base, the margins serrate,
more or less scabrous above or becoming almost smooth in age, softly pilose-
tomentose beneath; panicles densely flowered, often compact, mostly 3-6 (10) cm.
broad; heads numerous, sessile or short-pedicellate; phyllaries 3-4, broadly ovate,
acute or obtuse, 3-4.5 mm. long, striate, more or less strigillose or glabrate, ciliate;
pistillate flowers 3-6; hermaphrodite flowers 10-14; corollas glabrous, white, 2-3 mm.
long; achenes obovoid, 2-2.5 mm. long, pubescent at the apex.
Clibadium pittieri Greenman, Proc. Amer. Acad. 39: 98. 1903.
C. pittieri f. phrixium Greenman, op. cit. 40: 38. 1904. C. polygynum
Blake, Contr. Gray Herb. 52: 32. 1917. C. terebinthinaceum var.
pittieri O. E. Schulz, Bot. Jahrb. 46: 626. 1912.
Wet thickets or open forest near sea level to 350 m.; Alta
Verapaz. British Honduras; Honduras; Nicaragua; Costa Rica;
Panama; West Indies.
Shrubs 1.5-3 m. high, the rather slender branches densely hispid; leaves petiolate,
the blades ovate to lance-ovate, mostly 7-12 cm. long, acuminate or long-acuminate,
228 FIELDIANA: BOTANY, VOLUME 24
cuneate at the base or abruptly cuneate and decurrent on the petiole, triplinerved far
above the base, the margins serrate, scabrous and hispidulous on both surfaces;
panicles dense or rather open, with rather few heads on pedicels 3-7 mm. long; heads
globose, 5-6 mm. in diameter; phyllaries 2 or more, broadly ovate or oval, subacute,
striate, about 3 mm. long; receptacles paleaceous almost throughout; pistillate
flowers 20-30, the hermaphrodite ones about 8; corollas of the hermaphrodite flowers
1.8 mm. long, the 5 teeth pilosulous; achenes purplish to black, obovoid, marginate,
pilosulous at the apex, about 2 mm. long.
COREOPSIS Linneaus
References'. E. E. Sherff, Revision of the genus Coreopsis, Field
Mus. Bot. 11: 279-475. 1936; D. J. Crawford, Systematic Studies on
Mexican Coreopsis (Compositae). Coreopsis mutica: Flavenoid
Chemistry, Chromosome Numbers, Morphology, and Hybridization,
Brittonia 22: 93-111. 1970.
Herbs or shrubs, glabrous or pubescent; leaves chiefly opposite, petiolate, the
blades simple or lobate or 2-3-parted, or 2-3 times ternately or pinnately divided, the
margins entire or dentate; heads radiate, pedunculate or pedicellate, solitary or
disposed in lax, corymbose panicles; phyllaries mostly biseriate, more or less connate
at the base, the outer ones herbaceous, appressed or often spreading, the inner ones
mostly larger, brown or yellow, membranaceous; receptacle flat or somewhat convex;
ray flowers uniseriate, sterile or rarely styliferous and fertile, the ligules spreading,
entire or dentate, commonly yellow; pales flat or concave, membranaceous,
subtending the disc flowers; disc flowers perfect and fertile or the innermost sterile,
the corollas tubular, the limb usually 5-dentate; anthers entire or subsagittate at the
base; style branches truncate or conic at the apex or short-appendaged; achenes
obcompressed, orbicular to oblong or oblong-linear, often with 2 wings, these
membranaceous or indurate, entire or pectinate-dentate, flat or incurved; pappus
none or of 2 bristles, teeth, or scales, never of retrorse-scabrous setae.
Species about 100, in both hemispheres, mostly in tropical
regions, but in North America also in temperate areas. Only one
variable species is known in Central America, in the wild state, and
only one of the seven varieties recognized by Crawford is found in
Guatemala.
Coreopsis mutica DC. Prodr. 5: 571. 1836.
Erect or arching shrubs, 1-3 m. tall, often forming dense clumps, or sometimes
more or less scandent, the branches terete; leaves petiolate, the blades simple, ovate
to lanceolate, or 2-3-lobate, the margins serrate, mostly 3-12 (-18) cm. long,
acuminate, acute at the base, essentially glabrous above, glabrous beneath or more or
less villosulous or pubescent, penninerved; heads few or numerous, mostly 2-5.5 cm.
wide (including the ray flowers), on pedicels 1-4 cm. long, disposed in corymbose
panicles; involucres usually glabrous; outer phyllaries about 5, oblong to spathulate-
oblanceolate, often mucronulate, 4-13 mm. long, the inner ones almost twice as long,
ovate-oblong to rounded-ovate; pales glabrous, oblong-linear; ray flowers 5-12, the
ligules bright yellow, 1-2 cm. long; disc flowers yellow; achenes obcompressed, black,
MELCHERT: FLORA OF GUATEMALA 229
pale-marginate, glabrous, 6-13 mm. long, 1-3 mm. broad; pappus none or rarely the
inner achenes with 2 bristles.
Coreopsis mutica var. microcephala Crawford, Brittonia 22:
109. 1970. Santo Domingo (Guatemala). Figure 60.
Damp thickets or open forest, often in pine-oak forest, 900-2,400
m.; Alta Verapaz; Baja Verapaz; Chimaltenango; Chiquimula;
Escuintla; Guatemala; Huehuetenango; Jalapa; El Progreso; Que-
zaltenango; El Quiche; Sacatepequez; Solola; Zacapa. Mexico
(Chiapas); Honduras; El Salvador.
Differs from the typical variety in its large, commonly simple
leaves, mostly 6-18 cm. long (those of var. mutica mostly 4-8 cm.
long and usually many of them 2-3-lobate or 2-3-parted), and in its
smaller heads, 2-3.5 cm. wide (those or var. mutica commonly about
4 cm. wide).
COSMOS Cavanilles
By THOMAS E. MELCHERT
Tap-rooted annuals, rhizomatous-suffruticose perennials, or herbaceous per-
ennials with tuberform roots, the stems terete, multiribbed (sometimes angled when
pressed); leaves opposite, ours commonly 1-2-pinnatifid, rarely simple, unlobed; heads
radiate, meduim size to rather large, long-pedunculate, usually solitary on rather long
peduncles, sometimes corymbose-paniculate; involucre subhemispheric, biseriate,
dimorphous; outer phyllaries 5-8, green, herbaceous, inner phyllaries membranous,
striate, clear or slightly pigmented with yellow (chalcone) or reddish (anthocyanin)
pigments; receptacle essentially flat; ray florets 5-8 (double forms in cultivated forms
of C. sulphureus and C. bipinnatus), neutral, ligules commonly rosaceous to light
lavender or white, some orange-yellow, sulphureous or sanguineous; disc florets
yellow above, whitish below, some with masking rosaceous pigments; stamen
filaments with dense to scattered pilose hairs; style branches thickened above,
hirtellous, tipped with short, acute appendages; chaff flat to slightly concave, or, in
the annuals, expanded below, abruptly narrowing to an elongate, yellow, subfiliform
tip which protrudes between the disc florets or achenes; achenes fusiform -tetragonal,
each of the four faces with a median longitudinal sulcus, in mature, fully expanded
achenes a slender nerve seen extending the length of this sulcus, the mature central
achenes plump, erect, round-edged, the peripheral achenes usually slightly incurved,
somewhat obcompressed, like immature achenes their corners often angular, apex
gradually narrowed or, in the annual species, abruptly and conspicuously beaked,
antrorsely setose or smooth, aristate or exaristate; pappus awns in ours (1-) 2-5,
retrorsely barbed, erect to divergent.
A genus with about 35 taxa centering in central Mexico, only
five of which extend southeastward into Chiapas, Guatemala and
beyond. None are dominant members of the Guatemalan flora.
230 FIELDIANA: BOTANY, VOLUME 24
Plants perennials; leaves simple, unlobed to once-pinnatifid; achenes beakless.
Rhizomatous, suffruticose perennials; leaf segments narrowly linear; rays mostly 5.
C. crithmifolius.
Herbaceous perennials with tuberous roots; leaf segments broad, ovate to lance-
spathulate or leaves simple, coarsely toothed; rays mostly 8 C. diversifolius.
Plants annuals; leaves 2-3-pinnatisect; achenes beaked.
Rays orange-yellow C. sulphureus.
Rays purple-lavender to pink or partially to totally white.
Heads relatively inconspicuous, 2-2.5 cm. across; rays small, only 1-1.6 cm. long;
ultimate leaf segments lanceolate to lance-ovate, 2-6 mm. wide; achenes long
beaked C. caudatus.
Heads showy, 3.5-8 cm. across; rays mostly 2-3.5 cm. long; ultimate leaf segments
elongate, linear-filiform, 0.4-1 (-1.5) mm. wide; achenes short beaked.
C. bipinnatus.
Cosmos bipinnatus Cav. Ic. 1: 10. 1791. Coreopsis formosa
Bonato, Pis. Autom. 22. 1793. Cosmea bipinnata Willd. Sp. PL 3:
2250. 1804. Georgia bipinnata Spreng. Syst. 3: 611. 1826. Cosmos
bipinnatus var. exaristatus DC. Prodr. 5: 606. 1836. Cosmus
tenuifolius Lindl. Bot. Reg. 23: pi. 2007. 1837. Cosmea tenuifolia
Lindl., Heynh. Nom. 1: 223. 71840. Bidens formosa Sch.-Bip., Seem.
Bot. Voy. Herald 307. 1856. B. lindleii Sch.-Bip., foe. cit. Cosmos
bipinnatus var. typicus Sherff, Brittonia 6: 341. 1948.
Native to the wet, intermountain "llanos" of central Mexico,
this showy annual is widely cultivated in a variety of single- and
double-rayed forms; ours seemingly escaped from cultivation; Alta
Verapaz; Huehuetenango; Jalapa. Adventive or escaped in Central
and South America and the United States.
Annual herbs, branched, to 2 m. tall, the stems usually with minute, antrorsely
dissected, scabrous hairs, dense to scattered, some essentially glabrous; leaves 2-3-
pinnatifid, mostly 5-15 cm. long including the petiole, this inconspicuous, subsessile to
10 (-15) mm. long, winged, leaf segments linear-filiform to narrowly linear, 0.5-1 (-1.7)
mm. wide, their tips acute, indurated, but not particularly sharp; heads very showy,
mostly 5-7 (-8) cm. across; ligules commonly 8, obovate, (16-) 20-35 (-40) cm. long,
(8-) 12-20 (-25) cm. wide (pressed), apically subtruncate with about 3 broad, undulant
dentations, markedly narrowed below, rosaceous to lilac or white, intensely
pigmented anthocyanin spots may occur at the ligule base; outer phyllaries usually 8,
ovate to lance-caudate, 7-15 mm. long, 3-5 (-6) mm. broad; inner phyllaries ovate-
lanceolate, 8-12 mm. long, hyaline with numerous black stria tions and a broad, to 1
mm. wide, clear border, sometimes with yellowish or rosaceous pigment, apex ciliate;
disc florets numerous, corollas yellow, becoming white below, 5-6 mm. long; anthers
brownish black, about 3 mm. long, their terminal appendages short-triangular, 0.5-0.8
mm. long, hyaline; style branch tips short, rather blunt, about 0.5 mm. long; chaff
with golden-yellow, filiform-caudate tips protruding between the disc florets or
achenes, their expanded bases hyaline, yellow-lined; achenes becoming blackish,
smooth or very short-setose, linear-fusiform, the outer ones 5-8 mm. long, abruptly
rounded to a short, 0.5-1.7 mm. long, but distinct rostrum, the inner achenes to 18
MELCHERT: FLORA OF GUATEMALA 231
mm. long, their yellowish beaks to 4 or 5 (rarely 10) mm. long; pappus or 2 or 3 awns
or exaristate, awns to 3 mm. long.
Cosmos caudatus HBK. Nov. Gen. & Sp. 4: 240. 1820. Bidens
berteriana Spreng. Syst. 3: 454. 1826. Cosmea caudata Spreng. torn,
cit. 615. Bidens caudata Sch.-Bip. in Seem. Bot. Voy. Herald 308.
1856. Figure 61.
A tropical weed occurring in damp roadside thickets, wet
tropical grasslands, cultivated fields, etc., between sea level and 900
m., occasionally to 1,360 m.; Chiquimula; Escuintla; Guatemala;
Jutiapa; Retalhuleu; Santa Rosa. Very common throughout the
West Indies and Central America; in Mexico from the Yucatan
northward along the Gulf Coastal Plain through Veracruz; South
America; now widely adventive in the tropical Pacific, e.g., Hawaii;
Philippines; Fijis; Haiti; Borneo; Sumatra; Ceylon; India; etc.
Tall, branching annuals, 0.5-2 (-3) m. high; stems terete, striate, glabrate to
sparsely pubescent with elongate, plurilocular hairs; leaves 2-3-pinnatisect with
lanceolate segments, broadly triangular, 16-20 (-37) cm. long including the 2.5-13.5
cm. petiole, to 24 cm. wide, broadly triangular, ultimate segments lanceolate to
oblong-lanceolate, 3-8.5 cm. long, 2-5 (-6.5) mm. wide, sharp-pointed, margins entire,
spinulose-ciliate, surface glabrous or with inconspicuous, scattered, forward-pointing
hairs; heads relatively few and rather small relative to the plant size, 2.5-3 (-3.5) cm.
across, the rays of pressed specimens mostly equalling to slightly exceeding the disc,
heads single or in clusters of 2-4 on each main inflorescense branch; peduncles
slender, to 14 cm. long; outer phyllaries mostly 8, linear-subulate to lanceolate-
sharp-pointed, 6-9 (-25) mm. long, 1-1.8 (-2) mm. wide, midnerve and 2 or 4 laterals
prominent, ours ciliate; inner phyllaries oblong-lanceolate, acute 8-11 mm. long, 1.7-3
mm. wide, inconspicuously striate, usually rosaceous; rays rosaceous to nearly white,
oblong-oblanceolate, 10-15 (-18) mm. long, 6-8.5 mm. wide, apically 3-dentate, the
central lobe longest; disc florets numerous, yellow, 6.5-8 mm. long; anthers about 3
mm. long; style appendages about 1 mm. long; palae resembling the inner bracts,
usually yellowish, some overlaid with rose, 8.15-13 mm. long; achenes with elongate
narrow beaks extending 5-15 mm. above the palae, black, 13-33 mm. long including
the beak; beaks scabrous, one-half to one-third the achene length, usually aristate,
the peripheral achenes shorter, somewhat incurved, short-beaked; pappus awns 2,
slender, becoming divergent to reflexed, 2.5-4 mm. long, retrorsely barbed.
Cosmos crithmifolius HBK. Nov. Gen. & Sp. 4: 242. 1820.
Cosmea crithmifolia Spreng. Syst. 3: 615. 1826. Bidens sartorii Sch.-
Bip. in Seem. Bot. Voy. Herald 307. 1856. B. valladolidensis Sch.-
Bip. op. cit. 308. 1856. Coreopsis verticillata Sesse & Moc. PI. Nov.
Hisp. 147. 1890 (as to Mexican specimen in Sesse & Mocifio
collection at Madrid). Not Coreopsis verticillata L. 1753.
Grassy, rocky slopes in oak and/or pine montane woodlands,
now often on top of ledges formed by steep mountain roadcuts,
232 FIELDIANA: BOTANY, VOLUME 24
mostly 1,200-2,800 m.; Chimaltenango; Chiquimula; Huehue-
tenango; Jalapa. Widespread, Chihuahua and Nuevo Leon, Mexico
southeastward in montane forests throughout Central America,
rarely into Panama.
Rhizomatous perennials to 1 m. tall, the stems becoming lignescent basally,
terete with prominent longitudinal nerves, angulate and sulcate, particularly so when
pressed, the nerves often amber or rosaceous, glabrous or with scattered hairs; leaves
stiff, the principal ones characteristically 3-7-partite with narrowly linear, evenly and
widely spaced, sharp-pointed segments, 5-12 (-14) cm. long petioles included,
becoming simple above, rarely simple throughout or a few twice pinnatifid, the
segments evenly and widely spaced, opposite to subopposite, (0.5- ) 1-2 (-2.5) mm.
wide, thickish with a single, prominent midvein, surface often appressed-hispid,
becoming brown to reddish, margins scabrous-ciliate; heads radiate, showy,
commonly to 5.5 cm. across, long-pedunculate, usually only one on each main
branch; outer phyllaries 8, lance-subulate, 4-9 mm. long, 1.5-3 mm. wide, indurate-
tipped, with about 5 dark, elevated veins (or vein pairs) when dried; inner phyllaries
broadly oblong-ovate, chaffy-white to yellowish with black striations, 8-14 mm. long,
about 5 mm. wide; ray florets 5 (6), deep pink to rosy white, often drying purple-
rosaceous, limb broadly obovate, 1.7-3 cm. long, 1.5-2 cm. wide, sharply 3-denticulate,
prominently nerved, long white hairs near base of the dorsal surface; disc florets
numerous, yellow above, whitish below, 7.5-10 mm. long, teeth prominent, about 1.8
mm. long with numerous papillate hairs on their inner surface, anther tubes brownish
to black, (3.5-) 4-5 mm. long, style branches often reddish; palae yellowish, outer ones
oblong-lanceolate, inner linear-lanceolate, to 12 mm. long; achenes brown, tetragonal,
11-17 mm. long, narrowed above but never rostrate, antrorse-setose throughout,
strongly so above, aristate; pappus awns (2-) 3-5, erect, to 5 mm. long, unequal.
Cosmos diversifolius Otto, Knowles & Westc. Fl. Cab. 2: 3.
1838. Cosmos reptans Benth. PI. Hartw. 40. 1839. Dahlia repens
Hartw. in Benth. PI. Hartw. 40, as syn. 1839. Cosmea diversifolia
Hort., Heynh. Norn. 1: 222. 71840. Bidens diversifolia Sch.-Bip. in
Seem. Bot. Voy. Herald 308. 1856. B. reptans Sch. Bip., foe. cit. Not
B. reptans G. Don. B. dahUoides S. Wats. Proc. Am. Acad. 26: 142.
1891. Cosmos diversifolius var. pumilus Sherff, Bot. Gaz. 88: 305.
1929.
Plants of sandy, alpine meadows and open spots in thinly
forested, pine-oak woodlands, now frequent on tops of roadcuts,
mostly 2,000-3,400 m.; infrequent in Guatemala; Chimaltenango;
Huehuetenango; Quezaltenango; Solola; Sacatepequez. Chiapas and
Oaxaca to Mexico State, then northward along the Sierra Madre
Oriental into San Luis Potosi and Tamauliapas.
Scapose to subscapose herbs with fasicled, tuberform roots, the stems single or if
few, then branching at or near the base, ours mostly glabrous; leaves alatepetiolate or
sometimes spathulate, 5-12.5 cm. long, 1.5-2.5 (-3.5) cm. wide, membranous, margins
entire or with a few coarse teeth, gradually narrowed below, sometimes 3-7 (-10)
partite, segments more or less ovate to cuneate or oblong-lanceolate, 1-3.5 cm. long,
MELCHERT: FLORA OF GUATEMALA 233
(2.5-) 3-10 (-13) mm. wide, ours mostly glabrous; heads large, (3.5-) 4.5-7 cm. across,
normally solitary and long-pedunculate, at most one or two heads on each main
branch; peduncles 20-37 (-45) cm. long; involucre conspicuously striate; outer
phyllaries 8 (10), thick, subfoliose, oblong-lanceolate to ovate-oblanceolate, (7.5-) 10-
12.5 mm. long, (2-) 2.5-4 (-4.5) mm. wide, exterior surface with 6-10 conspicuous black
veins, these usually parallel, some branched, inner surface light green to tannish,
unveined; inner phyllaries white- to yellowish white- translucent, surface with 13-17
paired blackish striations, apex and margins sometimes rosaceous, 8-13 mm. long, 3-6
mm. wide; ray florets 8 (10), anthocyanin concentration highly variable, deep purple
to light lavender or white with rosaceous veins, limb oblanceolate to nearly obovate,
2-3 (-3.5) cm. long, (7.5-) 13-16 mm. wide, apex, at most, obscurely denticulate; disc
florets numerous, yellow, mostly 6 mm. long; anthers brownish black, 3-3.5 mm. long,
their terminal appendages triangular, 0.5-0.6 mm. long; palae clear to slightly
yellowish with 3-5 pairs of parallel black striations; achenes fusiform, never beaked,
weakly sulcate, brownish or castaneous, mostly 1-2 cm. long, aristate; pappus awns 2-
3 (-5), unequal, 2-3.5 mm. long, retrorsely barbed.
With one exception, the Guatemalan populations of C.
diversifolius have heads which are 4-6 cm. across, a feature which
allies them closely to diploid populations from Oaxaca and Chiapas.
One collection, however, from the plains near Tecpan, Chimal-
tenango, A. F. Skutch 568 (US), has notably larger heads with rays
about 3.5 cm. long, the outer phyllaries foliose, overlapping, 5 mm.
wide. Grossly, these specimens are very similar to tetraploid
populations of C. diversifolius from Mexico and Hidalgo, Mexico.
Cosmos sulphureus Cav. Ic. 1: 56. 1791. Coreopsis artemisiae-
folia Jacq. Ic. PL Rar. 3: 16. 1793. Cosmea sulphured Willd. Sp. PL
3: 2250. 1804. Bidens sulphured Sch.-Bip. in Seem. Bot. Voy. Herald
308. 1856. Cosmos aurantiacus Klatt, Leopoldina 25: 105. 1889.
Coreopsis artemisifolia Sesse & Moc. PL Nov. Hisp. 148. 1890.
Cosmos sulphureus var. exaristatus Sherff, Field Mus. Publ. Bot. 8:
411. 1932. Cosmos sulphureus var. hirsuticdulis Sherff, Am. Journ.
Bot. 24: 90. 1937.
Cosmos sulphureus is a pantropical weed which reaches its
maximum development in, and is doubtlessly native to hot,
subtropical regions of central and southern Mexico; ours seemingly
adventive, along roadsides, in damp meadows, on brushy slopes, also
cultivated and escaping as a garden weed, mostly 1,100-2,100 m.;
Chimaltenango; Huehuetenango; Escuintla; Guatemala. Through-
out Central America; West Indies, Florida, South America.
Commonly cultivated in temperate North America and Europe.
Annual herbs, to 2 dm. tall, the stems of larger specimens well-branched,
basically terete, becoming angled or even somewhat quadrate when pressed, but never
tetragonal, glabrate to densely spreading white-pilose, the hairs elongate,
234 FIELDIANA: BOTANY, VOLUME 24
conspicuously plurilocular, their walls often with rosaceous pigment; leaves 2 (-3)
pinnatisect with lanceolate segments, 5-20 (-35) cm. long including the 1.5-11.5 cm.
petiole, blade broadly triangular in outline, ultimate segments 2-4 (-8) mm. wide,
obscurely spinulose-ciliate, sharply apiculate, surface glabrous or with a few hairs
along the veins; heads showy, radiate, orange with yellow centers, 3-5 cm. across;
outer phyllaries commonly 8, linear-subulate, 3-9 mm. long, 1-2.1 mm. wide, (1) 3 or 5
lined; inner phyllaries oblong- lanceolate, mostly yellowish with rose to orange
spots, 7-12 mm. long, 1.5-2.5 mm. wide; ray florets 8 (10), orange (rarely yellow), the
limbs broadly obovate, usually overlapping, 1.6-2.5 cm. long, 8-18 mm. wide, apically
3-denticulate, some nearly truncate; disc florets numerous, corollas yellow with
conspicuously papillate teeth, 4.5-10 mm. long; anthers brownish to yellow, 3.2-4.5
mm. long; style branches yellow, their appendages pilose, about 1.4 mm. long; palae
similar to the inner involucre, chaffy-white with yellow tips or yellowish throughout
with reddish spots; achenes with prominent, narrow beaks which rise well above the
palae, 8-27 (-33) mm. long including the yellowish 2.5-12 mm. beak, body grey to
blackish, often sharply angled, antrorsely setose above, weakly 2-awned or more
commonly truly or developmentally exaristate; pappus awns 0-2, (2.5-) 3-5 (-7) mm.
long, retrorsely barbed, often fragile and falling at maturity.
CUCHUMATANEA Seidenschnur and Beaman
Reference: C. E. Seidenschnur and J. H. Beaman, Cuchuma-
tanea, a new genus of the Compositae (Heliantheae), Rhodora 68:
139-146. 1966.
Minute annuals; leaves opposite, spathulate, usually anisophyllous, the blades
narrowing into petioliform bases, these slightly connate, the margins entire; heads
homogamous, discoid, sessile or subsessile, solitary, terminal, closely subtended by a
pair of spathulate leaves; phyllaries 2, membranaceous but with herbaceous tips;
receptacle conic, paleaceous; pales membranaceous; not enclosing the flowers,
persistent; flowers 5-10, hermaphrodite, fertile, actinomorphic, the tubular corollas
narrowly campanulate, 3-4-lobate, stamens 3-4, the anthers sagittate at the base,
with broadly ovate to suborbicular apical appendages; style constricted below the
branches, the branches acute, papillose to the base; achenes ellipsoid-oblong, striate;
pappus none.
Cuchumatanea steyermarkii Seidensch. & Beaman, Rhodora
68: 139. 1966. Figure 62.
Open pine forest, in black soil near limestone outcrops, 3,680-
3,750 m. (type from Sierra de los Cuchumatanes, between Llano de
San Miguel and Todos Santos, Beaman 3962)', also collected in
vicinity of Chemal, summit of Sierra de los Cuchumatanes,
Steyermark 50260.
Minute annuals to about 1 cm. high, often forming open mats; stems sparsely
hirtellous or glabrate; leaves opposite, decussate, spathulate, obtuse or rounded at the
apex, 2-7 mm. long, 0.5-3 mm. wide, narrowing into petioliform bases, these slightly
connate, the margins entire, sparsely ciliate, glabrous above but usually with some
sparse pubescence on the costae beneath; heads sessile or subsessile, solitary,
NASH: FLORA OF GUATEMALA 235
terminal, 2-2.5 mm. high and about 1 mm. wide, subtended by a pair of leaves;
involucres campanulate; phyllaries 2, oblong-obovate, 2-2.5 mm. long, the herbaceous
tips reflexed, lacerate near the apex; receptacle to 0.5 mm. high; pales to about 2 mm.
long, glabrous, minutely lacerate above; disc flowers 5-10, the corollas about 1 mm.
long, yellow with purplish lobes; achenes 1.1 mm. long, brownish black, glabrous;
pappus none.
DAHLIA Cavanilles
Reference: Paul D. Sorensen, Revision of the genus Dahlia
(Compositae, Heliantheae-Coreopsidineae), Rhodora 71: 309-365,
367-416. 1969.
Herbaceous or suffrutescent perennials from tubers or tuberously thickened
rootstocks, sometimes epiphytic; leaves opposite or in whorls of 3, usually petiolate,
the blades simple to 3 times pinnately parted; heads heterogamous, radiate, long-
pedunculate; phyllaries biseriate, the outer ones smaller, more or less herbaceous,
spreading or reflexed at anthesis, somewhat carnose and appearing ceraceous, the
inner ones membranaceous, variously colored, longitudinally striate, accrescent in
fruit; receptacles flat; pales scarious, yellowish to purplish, subtending the disc
flowers; ray flowers neutral or pistillate and sterile, the ligules entire or minutely
tridentate, variously colored; disc flowers hermaphrodite, fertile, regular, tubular;
anthers entire and obtuse at the base; style branches with hairy appendages; achenes
linear-oblong to obovate or spathulate, obcompressed; pappus wanting or reduced to
2 minute awns, or 2 very slender, filamentous awns.
Of the 27 reported species in Mexico and Central America (two
of them probably introduced in South America), four are known in
Guatemala but five are treated here as D. purpusii Brandg. from
Chiapas, Mexico may be expected in Guatemala. A sixth species, D.
excelsa Benth. (the type from Mexico) reported by Sherff as
occurring in Guatemala (Am. Journ. Bot. 38: 54-73. 1951), is
omitted in this treatment, as I, like Sorensen (Rhodora 71: 329-330.
1969), have been unable to examine any material. Sorensen
attempted to obtain specimens for study by revisiting the
Guatemalan localities (Solola) reported by Sherff, but his search
was unsuccessful. He has stated, "On the characteristics displayed
in the photographs of the type, Dahlia excelsa may be dis-
tinguished from Dahlia imperialis by its 3-7 (not 9-15) primary
leaflets. . . . Despite these distinctions and despite the unfortunate
lack of material by which to judge, I strongly feel that Dahlia
excelsa and Dahlia imperialis are variants of the same species." He
did, however, retain D. excelsa in his treatment on a provisional
basis.
Rays commonly orange to scarlet, sometimes yellow D. coccinea.
Rays white, pink, lavender, or purple.
236 FIELDIANA: BOTANY, VOLUME 24
Plants commonly 2-6 m. tall; leaves large, 50-90 cm. long, primary leaflets 9-15;
achenes 13-17 mm. long D. imperialis.
Plants commonly about 1 m. tall, or less; leaves smaller, 3-40 cm. long, primary
leaflets 3-7 or the leaves simple; achenes 5-13 mm. long.
Plants cultivated; heads double- flowered (ray flowers in 2 or more series).
D. pinnata.
Plants native; heads with a single series of ray flowers.
Median leaves simple, not lobed nor divided D. purpusii.
Median leaves compound D, australis.
Dahlia australis (Sherff) Sorensen, Rhodora 71: 378. 1969. D.
scapigera var. australis et f. australis Sherff, Am. Journ. Bot. 34:
143. 1947. D. scapigera var. scapigera f. serratior Sherff, torn. cit.
142. D. scapigera var. liebmannii Sherff, torn. cit. 143. D. scapigera
var. australis f. purpurea Sherff, torn. cit. 145. D. australis var.
liebmannii Sorensen, Rhodora 71: 383. 1969. D. australis var.
chiapensis Sorensen, torn. cit. 384. D. australis var. serratior
Sorensen, torn. cit. 386.
Rocky banks, ledges, and cliffs, 2,500-3,700 m.; Huehuetenango;
El Quiche; Solola; Totonicapan. Mexico.
Perennial herbs to about 1 m. tall, simple or branched, the stems essentially
glabrous or more or less pubescent; leaves pinnatifid to bipinnate, 3-21 cm. long, the
primary leaflets 3-5 (-7), ovate or rhombic-ovate to lanceolate or oblong, acute or
acuminate, cuneate at the base, sessile or petiolulate, essentially glabrous above or
with scattered hairs near the margins or sometimes finely pubescent along the veins,
usually pubescent along the veins beneath, the margins serrate to crenate-dentate;
heads solitary or in groups of 2 or 3, erect or nearly so; outer phyllaries reflexed at
anthesis, 7-15 mm. long, the inner ones 10-18 mm. long; rays purple or pink, acute,
acuminate, or denticulate; disc flowers 39-67, yellow or purplish; achenes 7-11.4 mm.
long.
Dahlia coccinea Cav. Icon. PI. 3: 33, t. 266. 1794. D. popenovii
Safford, Journ. Wash. Acad. Sci. 9: 369, /. 3. 1919 (type from
Sacatepequez, Popenoe 682). D. geniryi Sherff, Am Journ. Bot. 29:
332. 1942. D. coccinea var. steyermarkii Sherff, op. cit. 31: 280. 1944
(type from Huehuetenango, Steyermark 50341). D. coccinea var.
palmeri Sherff, op. cit. 33: 508. 1946. D. coccinea var. gentryii
Sherff, op. cit. 34: 152. 1947. Chunay de zopa, chunis-boch, comida
de cache, dalia de monte (Huehuetenango); dalia (Guatemala).
Figure 63.
Damp or dry, often rocky, open or brushy slopes, often in oak
or pine-oak forest, 1,100-3,000 m.; Chimaltenango; Guatemala;
Huehuetenango; Jalapa; Sacatepequez; Santa Rosa; Solola;
Totonicapan; Zacapa. Mexico; British Honduras. Reported to be
naturalized in Peru.
NASH: FLORA OF GUATEMALA 237
Erect herbs, 0.5-1.5 m. tall, simple or sparsely branched, glabrous or pubescent;
leaves opposite, variable, simple and unlobed to tripinnate, mostly 12-35 cm. long, the
primary leaflets 3-11, sessile or petiolulate, thin, ovate to oblong- lanceolate or elliptic,
acute to acuminate, the base acute, rounded, truncate, or rarely subcordate, the
margins serrate, sparsely villous, mostly on the veins beneath, or essentially glabrous;
heads erect or cernuous, usually few, on peduncles 2-30 cm. long; outer phyllaries
spreading or reflexed at anthesis, herbaceous, linear to oblong or obovate to
spathulate, 6-15 mm. long, the inner ones narrowly oblong, 11-20 mm. long; pales 10-
14 mm. long; rays 1.6-4 cm. long, yellow, orange, or scarlet, acute or denticulate at
the apex; disc flowers 71-157, yellow or tipped with scarlet, 8-10 mm. long; achenes 8-
13 mm. long; pappus wanting or reduced to 2 minute awns, these rarely elongated
into delicate, filiform threads.
Handsome, showy, wide-ranging, variable plants.
Dahlia imperialis Roezl ex Ortgies in Regel, Gartenflora 12:
243. 1863. D. maximiliana Hort. ex Hooker f. Bot. Mag. t. 7655.
1899. D. lehmannii Hieron. Bot. Jahrb. 19: 55. 1894. D. dumicola
Klatt, Bot. Beibl. Leopoldina 6. 1895. D. maxonii Safford, Journ.
Wash. Acad. Sci. 9: 371. 1919 (type from Alta Verapaz, Maxon &
Hay 3295). D. lehmannii var. leucantha Sherff, Am. Journ. Bot. 38:
70. 1951. Cana de agua (Alta Verapaz); catarina, santa catarina
(Guatemala, Sacatepequez, San Marcos); c'olox (Quecchi, Alta
Verapaz); dalia (Jalapa, Totonicapan); dalia de palo (Chimal-
tenango); flor de la concebida (Alta Verapaz); runai (Guatemala);
tunay (Huehuetenango, Totonicapan); tzoloj (Alta Verapaz).
Damp thickets, steep slopes, roadside hedges, wet meadows,
cornfields, oak-pine and coniferous forest, 1,200-3,800 m.; Alta
Verapaz; Chimaltenango; Chiquimula; Guatemala; Huehue-
tenango; Jalapa; Jutiapa; Quezaltenango; El Quiche; Sacatepequez;
San Marcos; Santa Rosa; Solola; Totonicapan. Mexico; Honduras;
El Salvador; Nicaragua; Costa Rica; Colombia.
Tall, herbaceous or suffrutescent perennials, mostly 2-6 m. but sometimes as
much as 9 m. tall, stems glaucous when fresh, multistriate when dry, those of the
previous year sometimes 10 cm. in diameter; leaves 50-90 cm. long, bipinnate or
tripinnate, the primary leaflets 9-15, ovate to oblong-elliptical, acute to long-
acuminate, usually cuneate or rounded at the base but sometimes truncate or
subcordate, sessile or petiolulate, essentially glabrous or with multicellular hairs
mostly along the veins, the margins serrate; heads very numerous, suberect or
cernuous, usually long-pedunculate; outer phyllaries reflexed at anthesis, herbaceous,
6-14 mm. long, obovate or oblanceolate, obtuse to subacuminate, the inner ones 15-25
mm. long, ovate, obtuse; pales to 2 cm. long; ray flowers with pubescent tubes, the
ligules white, pale pink, or lavender to bright purple, 3.5-6 cm. long, acute or
subacute, denticulate; disc flowers 128-172, yellow or sometimes the tips red, 9-11
mm. long; achenes 13-17 mm. long.
238 FIELDIANA: BOTANY, VOLUME 24
Dahlia pinnata Cav. Icon. PL 1: 57, t. 80. 1791. D.
sambucifolia Salisb. Parad. Lond. t. 16. 1805. D. pinnata var. nana
Jackson in Andrews Bot. Repos. 7: t. 483. 1807. Georgina variabilis
Willd. et vars. Hort. Berol. 2: tt. 93-95. 1809. G. superflua DC., Ann.
Mus. His. Nat. Paris 15: 310. 1810. D. superflua Ait. f. Hort. Kew.
ed. 2, 5: 87. 1813. Coreopsis georgina Cass. Diet. Sci. Nat. 18: 442.
1820. D. variabilis Desf. Cat. PL Hort. Paris, ed. 3: 182. 1829. D.
pinnata var. variabilis Voss in Vilmorin, Blumengart. 1: 489. 1894.
Described by Cavanilles from plants grown in the gardens of
Madrid from seed collected in Mexico, the plants, even at that time
hybrids, producing multiseriate ray flowers. Frequently grown in
Guatemalan gardens and escaping into thickets and along road-
sides; Alta Verapaz; Sacatepequez; Santa Rosa.
Perennial herbs to about 1 m. tall, usually unbranched below the inflorescence;
stems often reddish, somewhat scabrous; leaves simple or pinnatisect to bipinnate, 13-
25 cm. long, the 3-5 leafltets ovate to oblong-ovate, acute, sparsely pubescent, often
scabrous, the margins serrate or dentate; heads few, long-pedunculate, in groups of 2
or 3; outer phyllaries reflexed at anthesis, 10-14 mm. long, obovate, acute, the inner
ones 15-21 mm. long; ray flowers multiseriate, the ligules commonly lavender or
purple, sometimes white or pink, 3.5-4.5 cm. long, acute, denticulate; disc flowers 96-
144, yellow or sometimes more or less suffused with purple; achenes 11-13 mm. long.
Dahlia purpusii Brandg. Univ. Calif. Publ. Bot. 6: 76. 1914.
At present known only from the type locality in Chiapas,
Mexico, but might be expected in Guatemala.
Herbaceous perennials less than 1 m. tall, the stems glabrous; leaves simple,
sessile or very short -petiolate, the blades oblong-ovate to elliptical, short-acuminate,
almost rounded at the base, 8-12 cm. long, glabrous, the margins serrate-crenate;
heads long-pedunculate, solitary or in groups of 2 or 3, erect or nearly so; outer
phyllaries reflexed at anthesis, 8-12 mm. long, obovate, acute or obtuse, glabrous, the
inner ones 14-15 mm. long; rays probably purple, about 3 cm. long; disc flowers
probably yellow; mature achenes not seen.
DELILEA Spreng.
Erect annuals, the stems slender, branching, thinly or densely hispidulous or
scabrous throughout; leaves opposite, short-petiolate, the margins dentate or
subentire; heads heterogamous, radiate, clustered in the leaf axils and at the ends of
the branches; involucres compressed and flat; phyllaries 2-4, thin, herbaceous, the
outer one large, orbicular, green, the inner 2 or 3 opposite it, smaller; receptacle
naked; ray flowers 1-3, pistillate, fertile, the tube slender, the ligule minute,
spreading, yellow; disc flowers 1-4, hermaphrodite, sterile, regular, the limb narrowly
campanulate, 5-dentate; style branches hirtellous; anthers almost entire at the base;
achenes obovate, compressed; pappus none.
Four species, three confined to the Galapagos Islands, and the
following widely distributed one.
NASH: FLORA OF GUATEMALA 239
Delilea berterii Spreng. Bull Sci. Soc. Philom. Paris: 54-55, t.
1. 1823; Syst. 3: 674. 1826. Milleria biflora L. Sp. PL 1301. 1753.
Elvira martyni Cass. Diet. Sci. Nat. 30: 68. 1824. E. biflora DC.
Prodr. 5: 503. 1836. D. biflora Kuntze, Rev. Gen. 1: 333. 1891. Figure
64.
Damp or rather dry thickets or fields, sometimes on open or
shaded, rocky slopes, sea level to 2,500 m. (more common below
1,500 m.); Chimaltenango; Chiquimula; Guatemala; Huehue-
tenango; Jalapa; Jutiapa; Santa Rosa; Zacapa. Mexico; British
Honduras and Honduras to El Salvador and Panama; tropical
South America.
Erect plants, commonly 20-50 cm. tall, the stems simple or much branched,
slender, subterete, often reddish or brown, strigose or hispidulous; leaves short-
petiolate or sometimes subsessile, the blades thin, oblong-ovate to lance-oblong,
mostly 1.5-5 cm. long, acute to long-acuminate, obtuse or cuneate at the base,
triplinerved, the margins crenate to subentire, the upper surfaces hirsute, the lower
surfaces paler, strigose and scabrous; heads often in dense clusters on short, slender
pedicels; outer phyllary orbicular, about 5 mm. in diameter, broadly rounded at the
apex, cordate at the base, reticulate-veined, green, glabrate, the margins subentire or
crenulate; ray flowers 1 or 2 (commonly 1), the ligule yellow, bifid; disc flowers 1-4
(commonly 1 or 2); ray achene obovate, compressed, glabrous or pubescent near the
apex; pappus none.
DESMANTHODIUM Bentham
Usually erect shrubs or coarse herbs, glabrous or pubescent; leaves opposite,
sessile or petiolate, commonly triplinerved, the margins serrate; heads heterogamous,
discoid, small, sessile in headlike glomerules, these few or numerous, disposed in
corymbiform-paniculate inflorescences; involucres ovoid; phyllaries 3-8, ovate, obtuse,
membranaceous, the innermost lageniform, dorsally compressed, closed, enclosing the
pistillate flowers; receptacles small, flat, naked or nearly so; pistillate flowers 1-3,
fertile, the corollas slender, shorter than the style, truncate or 3-4-dentate;
hermaphrodite flowers 4-6, sterile, regular, the limb 5-cleft; anthers obtuse at the
base; style of the hermaphrodite flowers simple, hirtous; achenes of the pistillate
flowers included in the subtending phyllary, obovate-oblong to fusiform, dorsally
compressed, naked, glabrous, bearing at the apex the persistent corolla; achenes of
the hermaphrodite flowers abortive; pappus wanting.
Perhaps seven species in the mountains of tropical America.
Only two are known in Guatemala, but three are treated here as
one occurs in adjacent Chiapas, Mexico, and may be expected in
Guatemala.
Principal leaves with petioles conspicuously and broadly alate throughout, these
connate-perfoliate at the base D. perfoliatum.
Principal leaves sessile or petiolate but the petioles neither alate throughout nor
connate-perfoliate at the base (sometimes the blades rounded and then abruptly
cuneate or abruptly attenuate to the base).
240 FIELDIANA: BOTANY, VOLUME 24
Stems essentially glabrous or sometimes bifariously pubescent, the indument, when
present, confined to narrow, often inconspicuous lines or bands; leaves
essentially glabrous; heads 5-7 mm. high D. guatemalense.
Stems densely villous-tomentose; leaves more or less pubescent on both surfaces,
with dense, longer tomentum on costae and veins beneath; heads about 4 mm.
high D. tomentosum.
Desmanthodium guatemalense Hemsl. Biol. Centr. Am. Bot.
2: 142, t. 45. 1881. Figure 65.
Damp or wet thickets or forest, 1,500-3,600 m.; Chimaltenango;
Chiquimula; San Marcos; Sacatepequez (type form Volcan de
Fuego, Salvin s.n.); Santa Rosa; Suchitepequez. Honduras; El
Salvador.
Shrubs to 2.5 m. high, the stems erect or sometimes sprawling and subscandent,
brittle, striate, glabrous or sometimes bifariously and inconspicuously pubescent;
leaves sessile or the lower ones sometimes short- petio late, the blades ovate, ovate-
lanceolate, oblong-ovate, or elliptic, 6-23 cm. long, 1.5-12 cm. broad, acuminate, acute
to rounded at the base or often abruptly narrowed to a cuneate base, glabrous or
nearly so, the margins serrate to denticulate; glomerules of heads numerous, disposed
in small or large corymbiform panicles, bracteate at the base; involucres 4-6 mm.
long; phyllaries 6-8, subscarious, ovate, striate, glabrous, the outermost ones
sometimes ciliate; flowers glabrous, the hermaphrodite ones long-stipitate; achenes
black, shining.
Desmanthodium perfoliatum Benth. in Hook. Ic. PI. 12: 14, t.
1116. 1876. Flaveria perfoliata Klatt, Leopoldina 23: 146. 1887. D.
caudatum Blake, Journ. Wash. Acad. Sci. 28: 488. 1938.
Not yet reported from Guatemala. Mexico (Chiapas, Guerrero,
Oaxaca), commonly in forest, about 1,400 m.
Erect shrubs to about 2 m. tall, the stems striate, bifariously pubescent or
glabrate; leaves ovate to ovate-lanceolate, the principal ones 10-25 cm. long,
acuminate to long-acuminate, the uppermost ones sessile, subsessile, or short-
petiolate, the others with the blade abruptly narrowed into a broadly winged petiole,
this conspicuously connate-perfoliate at the base, the margins serrate, essentially
glabrous or glabrate above and shining, sometimes with a few scattered hairs and
obscurely puberulent along the costae and veins, the lower surfaces usually with some
pubescence on costae and veins; inflorescences corymbiform-paniculate, sometimes
somewhat convex, the branches bifariously sordid-pilosulous, the glomerules
composed of few-many heads and subtended by 2 appressed, ovate, striate bracts 3-5
mm. long, these sometimes ciliolate near the base; involucres about 5 mm. high;
phyllaries usually 3-6, acute or subacute, glabrous or sparsely pilosulous on the upper
part of the costa; pistillate flowers 3-5; hermaphrodite flowers 6-7, sterile, long-
stipitate, the corollas more or less pubescent above; achenes black, shining.
Desmanthodium tomentosum Brandg. Univ. Calif. Publ. Bot.
6: 73. 1914.
NASH: FLORA OF GUATEMALA 241
Open, wet forest, 1,600-1,800 m.; Baja Verapaz. Mexico
(Chiapas).
Shrubs, the stems to 1 m. or more tall, densely sordid-tomentose; leaves
petiolate, the blades ovate to lauce-ovate, acuminate, mostly 5-15 cm. long, 3-8 cm.
wide, cuneate or attenuate to the base, the margins serrate, ciliate, puberulent or
pubescent above, pubescent beneath, the costae and veins densely tomentose;
inflorescences dense, cymose-paniculate, the glomerules composed of 3-5 sessile or
subsessile heads, the subtending bracts lanceolate; involucres 3-4 mm. high; phyllaries
scarious, lanceolate to oblong, striate, about equalling the pistillate flowers;
hermaphrodite flowers ling-stipitate; achenes oblanceolate to fusiform, black, shining.
ECLIPTA Linneaus
Annual herbs, usually much branched and often prostrate or procumbent, rough-
pubescent; leaves opposite, sessile or short-petiolate, the margins serrate, denticulate,
or subentire; heads terminal and axillary, small, heterogamous, radiate, short-
pedunculate; involucres hemispheric or broadly campanulate; phyllaries imbricate in
2 series, subequal or the outer ones longer, broad, more or less herbaceous; receptacle
flat or convex, paleaceous, the pales setaceous, subtending the achenes; ray flowers
pistillate, fertile; disc flowers hermaphrodite, mostly fertile, their corollas tubular, 4-
5-dentate; style branches obtuse; achenes thick, tuberculate (in ours), those of the
ray flowers trigonous, those of the disk somewhat compressed; pappus none or of a
few short teeth.
Species perhaps four, in both hemispheres, with only one in
North America.
Eclipta alba (L.) Hassk. PL Jav. Rar. 528. 1848. Verbesina
alba L. Sp. PL 902. 1753. V. prostrata L. I.e. E. erecta L. Mant. PL
286. 1771. E. prostrata L. I.e. E. brachypoda Michx. Fl. Bor. Am. 2:
130. 1803. Figure 66.
Damp, shaded places, often in thickets, wet fields, or waste or
cultivated ground, frequently on sandbars along streams, near sea
level to 1,500 m., most frequent at low elevations; Chiquimula;
Escuintla; Guatemala; Izabal; Jutiapa; Peten; El Progreso; El
Quiche; Retalhuleu; Zacapa. Widely distributed in the United
States; Mexico; British Honduras to El Salvador and Panama;
West Indies; South America; warmer regions of the Old World.
Annual herbs, rather densely strigose throughout with whitish hairs, usually
prostrate or procumbent, sometimes erect, often forming small mats, the stems
mostly 20-75 cm. long, rather stout, often much branched; leaves sessile or the lower
short-petiolate, the blades lanceolate, oblong-lanceolate, or linear-lanceolate, 2-10 cm.
long, 0.5-2 cm. wide, acute or acuminate, obtuse or acute at the base, the margins
rather remotely denticulate or subentire; heads usually numerous, 6-12 mm. broad,
slender-pedicellate or subsessile; ray flowers usually numerous but inconspicuous, the
ligules mostly about 1 mm. long, whitish; disc flowers numerous, about 1 mm. long;
242 FIELDIANA: BOTANY, VOLUME 24
anthers brown; achenes 2-2.5 mm. long, tuberculate, glabrous save for a few short
hairs at the apex; pappus reduced to a few minute, inconspicuous teeth or wanting.
ELEUTHERANTHERA Poiteau
Pubescent annuals, erect or prostrate, with slender branches; leaves opposite,
petiolate, the margins dentate, denticulate, or subentire; heads small, subsessile or
short-pedicellate in the forks of branches or in the axils of the upper leaves,
homogamous, discoid, the flowers all hermaphrodite and fertile or sometimes the
central ones sterile; involucres campanula te or ovoid; phyllaries 5-10, subequal, the
outer ones more or less herbaceous, hispid; receptacle small, convex; pales
membranaceous-scarious, embracing the flowers; corollas regular, tubular, yellow, the
limb 4-5-dentate; anthers subsagittate at the base, the auricles minute; style branches
rather long, acute, hirtous dorsally; achenes obovate-oblong, pilosulous near the apex
when young, often tuberculate when mature, especially near the apex or along the
angles, rather thick, obscurely angulate, rounded at the apex; pappus small,
cyathiform, ciliate-dentate, sometimes with 2-3 very short setae on the short neck or
cup of the achene.
The genus consists of a single species.
Eleutheranthera ruder alis (Swartz) Sch. Bip. Bot. Zeit. 24:
165. 1866. Melampodium ruderale Swartz, Fl. Ind. Occ. 3: 1372.
1806. Ogeira triplinervis Cass. Diet. 35: 445. 1826. Wedelia dis-
coidea Schlecht. Linnaea 6: 728. 1831. E. ovata Poit. ex Steud.
Nom. ed. 2, 1: 549. 1841. Kegelia ruderaUs Sch. Bip. Linnaea 21:
245. 1847. E. prostrata Sch. Bip. Bot. Zeit. 24: 239. 1866. Figure 67.
Grassy, open banks, sea level to about 500 m.; Izabal (Quirigua,
Standley 72201). Pacific coast of Honduras; Costa Rica; Panama;
West Indies; tropical South America.
Plants usually prostrate, sometimes erect, the stems slender, mostly 15-50 cm.
long, pilose; leaves on petioles to 1 cm. long, the blades ovate to oblong-lanceolate,
mostly 1.5-5 cm. long, acute or obtuse, rounded to acute at the base, the margins
inconspicuously dentate or crenate-dentate, 3-nerved, strigose or hirsute; pedicels 2-12
mm. long, the heads nutant; involucres commonly 4-6 mm. long; phyllaries oblong or
ovate-oblong, obtuse, hispid with hairs often more than 1 mm. long, and minutely
glandular-punctate; pales long-ciliate at the apex; flowers few, the corollas pale
yellow, the 5 lobes hirsute within on the margins; achenes 2-3 mm. long, more or less
tuberculate when mature; pappus cyathiform.
GALINSOGA Ruiz & Pavon
Annual herbs, usually branched, erect or decumbent; leaves opposite, usually
petiolate; inflorescences cymose; heads pedicellate, heterogamous, radiate; involucres
hemispheric or broadly campanulate; phyllaries biseriate, ovate, obtuse, membranous
but green, striate, subequal or the outer ones shorter; receptacle conic; pales thin,
subtending but not enfolding the disk flowers; ray flowers 4-5, pistillate, fertile, the
ligules small, tridentate, white, pink, or purplish red; disc flowers hermaphrodite,
NASH: FLORA OF GUATEMALA 243
fertile, the corollas 1-2 mm. long, 5-dentate; anthers minutely sagittate at the base;
style branches flattened, the appendages very short; achenes narrowly obpyramidal,
obscurely or conspicuously 4-5-angulate, or the outer ones compressed; pappus of the
ray achenes of few-several short, slender setae, or none, that of the disc achenes of
several laciniate or fimbriate scales, these short or long, sometimes aristate.
Three or four species, in the mountains of tropical America,
extending into the temperate regions of the United States, with
only one in Guatemala.
Some species are naturalized extensively as weeds in the United
States, where they have become abundant, especially in the waste
ground and vacant lots of large cities.
Galinsoga urticaefolia (HBK.) Benth. Bot. Voy. Sulphur:
119. 1844. Wiborgia urticaefolia HBK. Nov. Gen. 4: 257, t. 389.
1820. Sabazia urticaefolia DC. Prodr. 5: 497. 1836. G. parviflora
var. hispida DC. torn. cit. 677. Adventina ciliata Raf. New. Fl. 1: 67.
1836. G. hispida Benth. Bot. Voy. Sulphur: 119. 1844. G. aristulata
Bicknell, Bull. Torr. Bot. Club 43: 270. 1916. G. ciliata Blake,
Rhodora 24: 35. 1922. Hoja nueva (El Progreso); San Nicolas
(Sacatepequez). Figure 68.
Damp or wet fields or thickets, occasionally in open oak or oak-
pine forest, often a weed in waste or cultivated land, sometimes on
sandbars along streams, 250-3,800 m., most common at middle
elevations; Alta Verapaz; Chimaltenango; Chiquimula; Escuintla;
Guatemala; Huehuetenango; Jalapa; Jutiapa; El Progreso; Quezal-
tenango; El Quiche; Retalhuleu; Sacatepequez; Santa Rosa;
Suchitepequez; Totonicapan; Zacapa. Naturalized in the United
States; Mexico; El Salvador and Honduras to Panama; West
Indies; South America.
Erect annuals, usually much branched, mostly 15-50 cm. high, the stems more or
less pilose or hispid with spreading hairs; leaves short-petiolate, the blades ovate or
the uppermost ones lanceolate, mostly 2-7 cm. long, 1-4 cm. wide, acute, rounded or
acute at the base, the margins rather coarsely dentate, thinly pilose or hirsute;
inflorescences cymose, leafy, the pedicels numerous, slender, 0.5-3 cm. long, more or
less viscid-villous; heads 3-4 mm. high, 3-6 mm. broad; phyllaries green, 2-2.5 mm.
long, obtuse, striate, glabrate; pales lanceolate to linear-lanceolate, 2-3 mm. long, the
margins lacerate; ray flowers commonly 5, the ligules white or sometimes pink to
purplish-red, about 2 mm. long; disc flowers yellow; achenes about 1.5 mm. long,
those of the ray flowers somewhat flattened, usually glabrate, those of the disc
flowers black, 4-angulate, glabrate or hispidulous, the pappus scales 10-15, about as
long as the body of the achene, tapering and awnlike, the margins fimbriate.
GARCILASSA Poeppig
Erect annuals, the stems scabrous or hispidulous, branched; leaves mostly
244 FIELDIANA: BOTANY, VOLUME 24
opposite or ternate, petiolate, the blades membranaceous, 3-nerved, the margins
serrate; heads homogamous, discoid, 4-5-flowered, disposed in globose glomerules at
the ends of the branches; involucres very short; phyllaries few, strigillose; receptacle
small; pales membranaceous, embracing the flowers; corollas regular, the tube short,
the limb campanulate, 5-cleft; anthers entire at the base; style branches obscurely
appendaged, acute; achenes oblong, laterally compressed, contracted at the base,
pubescent, obscurely tuberculate; pappus reduced to a very short, inconspicuous,
ciliolate-fimbriate annulus.
The genus consists of a single species.
Garcilassa rivularis Poepp. & Endl. Nov. Gen. & Sp. 3: 46. t.
251. 1845. Figure 69.
Known in Guatemala from a single collection, damp thicket,
about 360 m., Alta Verapaz (near Pancajche, Standley 70609).
Nicaragua; Costa Rica; Panama; western South America.
Slender plants to about 1 m. high, the stems striate, sparsely strigose or glabrate;
lower leaves chiefly ternate, the upper ones opposite or subopposite, on long, slender
petioles, or the uppermost on short petioles, the blades very thin, lance-elliptic to
lanceolate, mostly 4-8 cm. long, acute or acuminate, cuneate to rounded at the base,
the margins crenate-serrate, sparsely scaberulous above, slightly paler and sparsely
strigose beneath; heads about 5 mm. high, short-pedicellate, in dense clusters;
phyllaries lanceolate, narrowly long- acuminate, strigillose, becoming yellowish in age;
pales mostly ovate, acute or acuminate, greenish, strigose; corollas greenish white,
pubescent; achenes scarcely 2 mm. long, obscurely tuberculate.
Collected only a few times in Central America, these plants
apparently are never plentiful. A single plant was found at the one
Guatemalan locality, the most northern station known for the
genus.
GOLDMANELLA Greenman
Sarmentose, perennial herbs, sparsely pilose or glabrate; leaves alternate, sessile
or short-petiolate, the blades very asymmetric, several-nerved from near the base,
membranaceous, the margins serrate; heads radiate, on very long, slender pedicels
disposed in an umbellate inflorescence at the ends of the branches or in their forks;
involucres campanulate; phyllaries 3-4-seriate, free; receptacle conic, paleaceous, the
pales membranous; ray flowers uniseriate, fertile, the ligules yellow; disc flowers
regular, fertile, the corolla yellow, the tube short, gradually ampliate above, the limb
5-dentate; anthers slightly sagittate at the base, apically appendaged; style branches
elongated, acute; achenes with subcoroniform pappus of 2-4 very short, stout, obtuse
awns.
A single species is known.
Goldmanella sarmentosa Greenm. Bot. Gaz. 45: 198. 1908.
Goldmania sarmentosa Greenm. Field Mus. Bot. 2: 271. 1907.
NASH: FLORA OF GUATEMALA 245
Caleopsis sarmentosa Fedde, Repert. Sp. Nov. 8: 326. 1910. Figure
70.
Open places, at or a little above sea level; British Honduras;
Mexico (Campeche).
Stems prostrate, rooting at the lower nodes, stout, glabrous below, pilose above
with ascending hairs; leaves sessile or subsessile, the blades ovate or rhombic-ovate,
mostly 3-6 cm. long, acute, very oblique at the base, rounded on one side, acute on
the other, the margins serrate, in age glabrous or nearly so; peduncles 5-10, very
slender, 2-8 cm. long, naked or with a few bractlets; involucres 6-8 mm. high;
phyllaries glabrous, yellowish with reddish brown lines, the outer ones short, broadly
ovate, acute, the inner ones oblong or oval, scarious-margined, rounded at the apex;
ray flowers 5-8, the ligules pale yellow, oblong-obovate, about 5 mm. long, 2-3-
dentate; disc flowers about 20; mature achenes oblong, 2.5-3 mm. long, glabrous but
slightly rugose, reddish brown; pappus subcoroniform of very short, stout, obtuse
awns.
The plants are conspicuous for their very asymmetric leaves, a
character unusual in the Compositae.
HELIOPSIS Persoon
Reference: T. R. Fisher, Taxonomy of the genus Heliopsis
(Compositae), Ohio Journ. Sci. 57: 171-191. 1957.
Perennial or annual herbs, usually with rough pubescence, sometimes glabrous;
leaves opposite or the uppermost alternate, petiolate, the margins dentate; heads
heterogamous, radiate, long-pedunculate, the peduncles often thickened at the apex;
involucres hemispheric or broadly campanulate; phyllaries 2-3-seriate, narrow,
subequal, the outer ones herbaceous; receptacle broadly convex; pales complicate,
embracing the disc flowers; ray flowers uniseriate, pistillate, usually fertile, the
corollas lacking a definite tube, sessile and persistent on the achene, the ligules bright
yellow, spreading, entire or inconspicuously bifid or tridentate; disc flowers
hermaphrodite, fertile, the corollas regular, the tube short, the limb cylindric, shortly
5-cleft; anthers entire at the base or minutely sagittate; style branches obtuse,
hirtous, short- appendaged; achenes glabrous, rather thick, obscurely 3-4-angulate or
subterete, sometimes becoming tuberculate; pappus none.
Of the 13 species, all American, in temperate regions of the
tropics and mostly in the mountains, only one if found in Central
America.
Heliopsis buphthalmoides (Jacq.) Dunal, Mem. Mus. Hist.
Nat. Paris 5: 57. 1819. Anthemis buphthalmoides Jacq. PL Hort.
Schoenbr. 2: 13, t. 151. 1797. Mabal (Huehuetenango). Figure 71.
Damp or wet thickets or oak forest, sometimes a weed in coffee
plantations, 750-3,000 m.; Huehuetenango; El Quiche; Sacatepe-
quez; Suchitepequez; Zacapa. Southern Mexico; Honduras; Costa
Rica; western South America.
246 FIELDIANA: BOTANY, VOLUME 24
Erect or decumbent perennials, the stems mostly simple below, 1 m. long or less,
sometimes rooting at the lower nodes, the internodes elongated, villous-hirsute or
glabrous; leaves petiolate, the blades membranaceous, ovate or broadly ovate, mostly
3-7.5 cm. long, acute or acuminate, truncate or rounded at the base and sometimes
abruptly decurrent on the petiole, triplinerved above the base, the margins irregularly
dentate, sparsely pilose on both surfaces with slender, soft, white hairs or almost
wholly glabrous; heads few, often only one, on very long, slender, naked, striate
peduncles, these slightly thickened at the apex, usually more or less hirsute, at least
near the apex; phyllaries biseriate, 6-7 mm. long, oblong or spathulate, obtuse or
rounded at the apex, ciliate, pubescent; ray flowers 8-10 (-14), the ligules yellow, 1-3
cm. long; pales rather rigid, linear-oblong, glabrous or nearly so, 7-8 mm. long,
obtuse; achenes glabrous; pappus none.
HETEROSPERMA Cavanilles
Branched annuals, erect or ascending to procumbent, pubescent or almost
glabrous; leaves opposite, the margins dentate or variously incised, often ternately or
pinnate ly dissected; inflorescences terminal and axillary; heads heterogamous,
radiate, pedunculate; involucres more or less campanulate; phyllaries few, biseriate,
the outer ones narrow, herbaceous, the inner ones subtending the ray flowers
membranaceous, striate, connate at the very base; receptacle flat; pales membrana-
ceous-scarious, almost flat, subtending the disc flowers; ray flowers pistillate, fertile,
the ligules spreading, usually inconspicuous, 2-3-dentate; disc flowers hermaphrodite,
fertile, the corollas yellow to orange, tubular, the limb little dilated, shallowly 5-cleft;
anthers obtuse at the base but very minutely sagittate; style branches of the disc
flowers short, acute, rather broad or subulate at the apex; achenes glabrous, dorsally
compressed, flat or concave, the outer ones rather broad and winged, without pappus,
the inner ones narrower, their wings narrower or none, attenuate into a short or long
beak, this commonly bearing at the apex 2 retrorse-aculeolate setae, or sometimes the
outermost disc achenes naked.
About eight species, all American, chiefly in the mountains of
tropical or warm-temperate regions. A single species is found in
Central America.
Heterosperma pinnatum Cav. Icon. 3: 34, t. 267. 1795. Figure
72.
Open, grassy or rocky slopes and meadows, sometimes in damp
thickets, pine-oak forest, waste or cultivated ground, often in sandy
soil along streams, 850-2,000 m.; Chimaltenango; Chiquimula;
Guatemala; Huehuetenango; Jalapa; Jutiapa; El Quiche; Sacatepe-
quez; Santa Rosa; Solola. Southwestern United States; Mexico;
Honduras.
Erect to procumbent annuals, commonly 20-50 cm. high, usually much branched,
the branches striate, hispidulous or almost glabrous; leaves petiolate or subsessile,
mostly 2-4 cm. long, divided into few or fairly numerous, linear segments, these
subacute, glabrous or nearly so, the petioles sparsely or densely long-ciliate;
peduncles numerous, often much longer than the leaves, sparsely pilose; involucres
NASH: FLORA OF GUATEMALA 247
broadly campanulate or hemispheric; outer phyllaries linear, about 5 mm. long,
green, ciliate, the inner ones about the same length, brownish or dark reddish,
conspicuously striate, broadly ovate, subacute or obtuse; ray flowers 3 or more, the
ligules pale yellow, inconspicuous, about 1.5 mm. long; disc flowers about 12, dull
yellow to orange; achenes black or dark brown with whitish wings or margins, those
of the ray flowers and outermost disc flowers 3-4 mm. long and 2-3 mm. broad, those
of the inner disc flowers 5-6 mm. long, with the beak 1-4 mm. long, scabrous on the
margins, the awns 2-4 mm. long.
HIDALGOA La Llave & Lexarza
Large, scandent, glabrous herbs with slender branches; leaves opposite, long-
petiolate, ternately or biternately divided, the leaflets broad, petiolulate or sessile, the
margins serrate; heads heterogamous, radiate, rather large, the long peduncles
solitary in the leaf axils; involucres open-campanulate; outermost phyllaries 3-5,
herbaceous, spreading, the inner 2 series membranaceous, erect; receptacle flat; pales
membranaceous, almost flat, subtending the ray flowers; ray flowers pistillate, fertile,
the style branches elongated, the ligules broad, spreading, orange or orange-red, 2-3-
dentate; disc flowers hermaphrodite, sterile, the corollas tubular, yellow, the limb 5-
dentate; styles apparently undivided but very minutely bifid at the apex; anthers
obtuse at the base, inconspicuously sagittate; ray achenes dorsally compressed, the
apex terminating in 2 inflexed, hornlike appendages.
The genus consists of four closely related species, with one in
Guatemala. Two are treated here, as the one in nearby Chiapas,
Mexico may be expected in Guatemala.
Outermost phyllaries commonly ovate; ray flowers 8-10 H. breedlovei.
Outermost phyllaries commonly linear to lanceolate; ray flowers about 5.
H. ternata.
Hidalgoa breedlovei Sherff, Sida 3: 261. 1966.
Not reported from Guatemala but may be expected, as it has
been collected several times in Chiapas, Mexico (damp forest, 1,600-
1,800 m.).
Slender, almost glabrous vines, often much branched; leaves long-petiolate, the
petioles often incurved at the base and functioning as tendrils, the 3 leaflets ovate to
triangular, mostly 3-5 cm. long, acute or short-acuminate, rounded to cuneate at the
base, the lateral ones asymmetrical, coarsely serrate; peduncles naked, very sparsely
hispid to almost glabrous, mostly 4-12 cm. long; phyllaries green, essentially glabrous
except for some indument at the base, the outer ones ovate to lanceolate, 0.3-0.6 cm.
long, the inner ones linear-lanceolate, about 1 cm. long, glabrous; ray flowers 8-10,
about 2 cm. long, orange to orange-red; pales linear-oblong, subacute; immature
achenes compressed, pubescent.
Hidalgoa ternata La Llave & Lex. Nov. Veg. Descr. 1: 15.
1824. H. steyermarkii Sherff, Field Mus. Bot. 23: 335. 1947 (type
248 FIELDIANA: BOTANY, VOLUME 24
from Huehuetenango, Steyermark 48814). Caballero (Huehue-
tenango). Figure 73.
Dense, wet thickets or forest, 350-1,650 m.; Alta Verapaz; Baja
Verapaz; Huehuetenango; Quezaltenango; San Marcos; Suchitepe-
quez. Southern Mexico; Costa Rica; Panama; South America.
Slender vines, climbing over large shrubs or good-sized trees, often much
branched and forming dense tangles, glabrous throughout or nearly so; petioles often
coiled or at least incurved at the base and functioning as tendrils, long and slender;
leaflets 3, ovate or triangular to elliptic, mostly 2-8 cm. long, acute or short-
acuminate, rounded to cuneate at the base, the lateral ones asymmetric, coarsely
crenate-serrate, thin; peduncles naked, long and slender; outer phyllaries green,
glabrous, linear to lanceolate, 6-9 mm. long, the inner ones about 8 mm. long, obtuse,
often pale-marginate; ray flowers about 5, the ligules bright orange-red, about 2 cm.
long; pales linear-oblong, subacute; achenes to about 8 mm. long.
HYMENOSTEPHIUM Bentham
Annual or perennial herbs or suffrutescent plants, commonly with rough
pubescence; leaves chiefly opposite, petiolate, the blades ovate to lanceolate,
generally 3-nerved or triplinerved, the margins crenate, serrate, or dentate;
inflorescences cymose and often corymbose-paniculate at the ends of branches; heads
heterogamous, radiate; involucres subcylindrical to broadly campanulate; phyllaries
biseriate, subequal or graduate, lanceolate or ovate, usually without conspicuously
herbaceous tips; receptacle flat or convex; pales concave or complicate, subtending or
embracing the disc flowers; ray flowers uniseriate, neutral, the ligules yellow, entire
or bidentate; disc flowers hermaphrodite, fertile, the corollas tubular, the limb little
dilated, 5-cleft; anthers rather obscurely sagittate at the base; style branches slightly
flattened, short-appendaged; achenes more or less obovoid, somewhat compressed,
pubescent or glabrous; pappus of few unequal, lacerate, fimbriate scales, or none.
About six species, in tropical America. Only the following three
are known from Central America.
Involucres in anthesis subcylindric, usually 2-3 mm. broad; phyllaries rigid, 1.5-2.5
mm. long; rays commonly about 5 mm. long H. microcephalum.
Involucres in anthesis campanulate, usually 4-7 mm. broad; phyllaries rigid or
reflexed, 4-6 mm. long; rays commonly 6-12 mm. long.
Indument of pedicels always of short, appressed hairs; indument of stems and
lower leaf surfaces sparse or dense, commonly appressed (that of veins
sometimes spreading) H. cordatum.
Indument of pedicels always of spreading, multiseptate hairs; indument of stems
and lower leaf surfaces always copious and spreading H. guatemalense.
Hymenostephium cordatum (Hook. & Arn.) Blake, Journ.
Bot. 53: 268. 1915. Wedelia cordata Hook. & Arn. Bot. Beechey
Voy. 435. 1841. W. subflexuosa Hook. & Arn. I.e. Gymnolomia
patens Gray, Proc. Amer. Acad. 5: 182. 1861. G. subflexuosa Benth.
& Hook, ex Hemsl. Biol. Centr. Am. Bot. 2: 163. 1881. Aspila
NASH: FLORA OF GUATEMALA 249
costaricensis Klatt, Bull. Soc. Belg. 31: 201. 1892. H. pilosulum
Blake, Journ. Bot. 53: 268. 1915. Sajan; sajan del rio (Guatemala
and Quezaltenango); singking (Chiquimula).
Damp or wet thickets and forest, or sometimes in rather dry
oak or oak-pine forest, or in abandoned fields, 300-3,000 m. (more
common between 1,000 and 2,200 m.); Alta Verapaz; Baja Verapaz;
Chimaltenango; Chiquimula; Escuintla; Guatemala; Huehue-
tenango; Jalapa; Quezaltenango; Retalhuleu; Sacatepequez; San
Marcos; Santa Rosa; Solola; Suchitepequez. Mexico; El Salvador
and Honduras to Panama; South America.
Erect or ascending herbs to 2 m. tall or sometimes reclining or more or less
scandent, the slender, terete stems then elongating to 3 or 4 m., strigose or strigillose
to puberulent or glabrate; leaves on petioles mostly 0.5-4 cm. long, the blades thin,
broadly ovate to lance-ovate, mostly 4-12 cm. long, triplinerved, acute or acuminate,
subcordate to cuneate at the base, the margins crenate to serrate, more or less
scabrous above, sparsely or densely pubescent beneath with mostly appressed hairs or
the hairs of costae and veins sometimes spreading; inflorescences usually lax and
open but the cymes occasionally congested; heads few or numerous on short or
elongated pedicels, these puberulent to strigillose with appressed hairs; involucres
broadly campanulate in anthesis; phyllaries lanceolate, acuminate, 4-6 mm. long,
appressed-strigillose or appressed-puberulent, more or less striate, often reflexed;
pales linear, acute, more or less fimbriate near the apex; ray flowers about 8, the
ligules bright yellow, commonly 6-11 mm. long; disc flowers yellow; achenes 2-2.8
mm. long, glabrous or densely appressed-pubescent; pappus none or of a few scales to
1 mm. long.
A weedy, highly variable complex with several intergrading
forms, including some with short rays, 4-5 mm. long, as in H.
micro cephalum (Less.) Blake. Plants with glabrous, epappose
achenes and those with pubescent, pappose achenes share the same
habitats and altitudes and are not distinct in any other feature.
It should be noted that the illustration of a Peruvian plant,
Gymnolomia rudbeckioides HBK. (Nov. Gen. & Sp. 4: 219, t. 374.
1820) appears very similar to the above species. However, according
to the description, the achenes of that plant are compressed, with
fimbriate margins, which would indicate another genus. Robinson
and Greenman (Proc. Boston Soc. Nat. Hist. 29: 95. 1899) stated
that they were "unable to match satisfactorily the original
description and figure of this species with any Mexican or Central
American plant."
Hymenostephium guatemalense (Robins. & Greenm.) Blake,
Contr. Gray Herb. 54: 8. 1918. Gymnolomia patens var. guatema-
lensis Robins. & Greenm. Proc. Boston Soc. Nat. Hist. 29: 94. 1899
250 FIELDIANA: BOTANY, VOLUME 24
(type from San Miguel Uspantan, El Quiche, Heyde & Lux 3370). G.
patens var. brachypoda Robins. & Greenm. torn. cit. 95 (type from
Coban, Alta Verapaz, Tuerckheim 852). G. guatemalensis Green-
man, Field Mus. Bot. 2: 347. 1912. Figure 74.
Thickets and forest, often in oak or oak-pine forest, 1,000-2,700
m.; Alta Verapaz; Baja Verapaz; Guatemala; Huehuetenango;
Jalapa; El Progreso; El Quiche; Sacatepequez; San Marcos; Solola.
Mexico; Honduras; Costa Rica; Panama.
Erect, ascending, or subscandent, herbaceous or sometimes suffrutescent plants
to 2 m. high, the stems sometimes elongating and reclining on other plants, terete,
usually densely villous-hirsute, sometimes hispid; leaves on petioles mostly 0.5-5 cm.
long, the blades ovate or broadly ovate, mostly 4-14 cm. long, acute or acuminate,
rounded or cuneate at the base, triplinerved, the margins crenate-serrate or serrate,
more or less scabrous on the upper surface, usually densely pilose or hirsute beneath
with spreading hairs; inflorescences usually lax and open but the cymes sometimes
congested; heads few or numerous on densely pilose, hispid, or hirsute pedicels, the
hairs multiseptate; involucres in anthesis broadly campanulate; phyllaries lanceolate,
acuminate, 4-6 mm. long, strigose or hispidulous, often striate, often reflexed; pales
linear, acute, usually more or less fimbriate near the apex; ray flowers about 8, the
ligules bright yellow, 6-12 mm. long; disc flowers yellow; achenes 2.5-3.2 mm. long,
glabrous or sparsely pilose; pappus none (in ours), or of a few unequal scales.
Common, weedy plants in many parts of Guatemala, sometimes
forming dense thickets.
Hymenostephium microcephalum (Less.) Blake, Contr. Gray
Herb. 54: 8. 1918. Gymnolomia microcephala Less. Linnaea 5: 153.
1830. Montanoa thomasii Klatt, Abh. Nat. Gesell. Halle 15: 328.
1882. G. patens var. abbreviata Robins. & Greenm., Proc. Amer.
Acad. 29: 387. 1894. G. patens var. macrophylla Robins. & Greenm.,
Proc. Boston Soc. Nat. Hist. 29: 95. 1899.
Damp thickets or forest, sometimes in pine forest, 1,000-2,000
m., Huehuetenango. Southern Mexico.
Erect, herbaceous or suffrutescent plants about 2 m. tall, the stems slender,
terete, striate, strigose or glabrate; leaves opposite or sometimes the uppermost ones
alternate, on petioles 0.5-3 cm. long, the blades lanceolate to ovate, mostly 3-10 cm.
long, 1.5-7 cm. wide, acuminate, cuneate to subcordate at the base, the margins
serrate, usually more or less scabrous above and strigillose or pilose beneath;
inflorescences cymose, usually becoming corymbose- paniculate; heads commonly
numerous, on pedicels mostly 0.5-2 cm. long, these a ppressed- pubescent; involucres in
anthesis subcylindric, 2-3 mm. broad; phyllaries rigid, lance-ovate, striate, appressed-
puberulent, mucronulate, 1.5-2.5 mm. long; pales 3-4 mm. long, striate, mucronulate;
ray flowers about 8, neutral, the ligules yellow, about 5 mm. long, minutely bifid at
the apex; disc flowers 20-35, the corollas about 3 mm. long, yellow, the lobes often
purplish; achenes about 2 mm. long, glabrous or sometimes sparsely pilosulous;
pappus none (in ours).
NASH: FLORA OF GUATEMALA 251
Blake considered H. mexicanum Benth. (1876) to be conspecific
with H. microcephalum. I have not seen authentic material of this
species, but according to the original description, H. mexicanum has
broadly campanulate involucres about 6 mm. long and achenes with
denticulate pappus scales.
IOSTEPHANE Bentham
Erect, scapose herbs or suffrutescent plants, the stems strigose and more or less
scabrous; basal leaves on winged petioles, the blades more or less trilobate to
pinnatifid, scabrous, the cauline leaves reduced to sessile bracts subtending the
inflorescences; heads pedunculate, solitary or 2-3, heterogamous, radiate; involucres
broadly campanulate to hemispheric; phyllaries 2-3-seriate, subherbaceous, green,
lanceolate, acuminate; receptacle convex to conical; pales concave, complicate, rigid,
membranaceous; ray flowers neutral, uniseriate, the ligules purplish or (in ours)
yellow; disc flowers numerous, hermaphrodite, fertile, regular, the tube short, the
limb cylindrical-campanulate, 5-cleft, yellow, more or less pubescent; style branches
acute; anthers sagittate at the base, the apical appendages lanceolate; disc achenes
obovate-oblong, the outer ones somewhat obcompressed, the inner ones more
compressed; pappus none.
This is a genus of two species, both Mexican. Neither has been
reported from Guatemala, but one is treated here as it occurs in
nearby Chiapas.
lostephane trilobata Hemsl. Biol. Centr. Am. Bot. 2: 169.
1882. Rudbeckia chrysantha Klatt, Leopoldina 23: 143. 1887.
Gymnolomia scaposa Brandg. Univ. Calif. Publ. Bot. 4: 93. 1910.
Figure 75.
Not reported from Guatemala but included here, as the type
(Ghiesbreght 101) was collected in nearby Chiapas, Mexico.
Erect, scapose herbs, the stems strigose, to about 40 cm. tall; basal leaves on
winged petioles 2-10 cm. long, the blades usually panduriform, sometimes ovate,
mostly 5-10 cm. long, indistinctly or deeply trilobate, the terminal lobes ovate, to
ovate-orbicular, acute or acuminate, rounded or truncate at the base and abruptly
decurrent on the petiole, the margins subentire to remotely serrate or crenate,
scabrous on both surfaces, the cauline leaves reduced to sessile bracts subtending the
inflorescence; heads solitary or 2-3 on peduncles 2-10 cm. long; involucres
hemispheric to broadly campanulate; phyllaries 2-3-seriate, 7-10 mm. long, lanceolate,
acuminate, strigose, ciliate; receptacle convex to conical; pales complicate, enfolding
the flowers, striate, puberulent, acuminate, mucronulate; ray flowers 6-8 (-13),
neutral, the tubes pubescent, the ligules yellow, mostly 9-15 mm. long; disc flowers
numerous, yellow, the corollas more or less pubescent, about 4 mm. long, the tube
less than 1 mm. long, more or less pubescent; outer achenes oblong-obovate, about 3
mm. long, slightly obcompressed, the inner ones more compressed, glabrous, striate,
black and shining when mature; pappus none.
252 FIELDIANA: BOTANY, VOLUME 24
JAEGERIA HBK.
Reference: Andrew M. Torres, Revision of Jaegeria
(Compositae-Heliantheae), Brittonia 20: 52-73. 1968.
Erect, spreading, or decumbent, branching annual or perennial herbs growing in
shallow water or wet or damp soil, the stems striate, pubescent or glabrate; leaves
opposite, sessile or petiolate, glabrous or pubescent, the margins dentate or entire;
heads heterogamous, radiate, borne on usually solitary, axillary peduncles; involucres
campanulate; phyllaries 1-2-seriate, equal or subequal, the lower inner portion
usually hyaline and enclosing the ray achenes; receptacle convex or conic; pales
scarious, ciliate, commonly embracing the hermaphrodite disc flowers; ray flowers
pistillate, fertile, the ligules white, yellow, pink, or purplish, up to 12 mm. long, the
apex entire or bi- or tridentate; style branches recurved; disc flowers numerous,
fertile, the corollas yellow or greenish, to 3 mm. long, the tube basally constricted and
pilose on the constriction; anthers sagittate at the base, apically appendaged; achenes
of both ray and disc flowers black, shining, glabrous; pappus none or reduced to a
minute annulus.
Eight species, all American, chiefly tropical, with only the
following in Central America.
Jaegeria hirta (Lag.) Less. Syn. Comp. 223. 1832. Acmella
hirta Lag. Gen. & Sp. PI. 30. 1815; 31. 1816. Mala hierba
(Huehuetenango). Figure 76.
Damp or wet pastures, cultivated fields, or banks, frequent on
sandbars along streams, sometimes in damp pine-oak woods, 250-
2,500 m. (most common in the mountains at middle elevations);
Alta Verapaz; Chimaltenango; Escuintla; Guatemala; Huehue-
tenango; Izabal; Jalapa; El Progreso; Quezaltenango; El Quiche;
Retalhuleu; Sacatepequez; Santa Rosa; Solola; Suchitepequez;
Totonicapan; Zacapa. Mexico; Honduras and El Salvador to
Panama; western South America.
Erect or spreading annuals, often much branched, commonly 10-40 cm. high, the
stems striate, very leafy, sparsely or very densely villous or hispid; lower leaves short -
petiolate, the upper ones sessile or nearly so, the blades ovate to ovate-oblong, mostly
1.5-5 cm. long, subacute to acuminate, cuneate to rounded at the base, thinly pilose
or hirsute, the margins undulate-dentate or subentire; heads numerous, on slender,
usually solitary peduncles (rarely 2, 3, or more in the leaf axils); involucres mostly 3-4
mm. wide; phyllaries uniseriate, lance-oblong to lance-linear, about 3.5 mm. long,
hispidulous or villous, the hyaline inner portion about 2 mm. long, erose; pales acute,
conspicuously ciliate; ray flowers inconspicuous, the ligules about 1.5 mm. long, pale
yellow to orange or often whitish in age or when dried, glabrous; achenes smooth,
black, little more than 1 mm. long, sometimes crowned by an inconspicuous annulus.
Very common and abundant weeds in cornfields of the central
mountains.
NASH: FLORA OF GUATEMALA 253
LAGASCEA Cavanilles
Reference: B. L. Robinson, Synopsis of the genus Nocca, Proc.
Amer. Acad. 36: 467-471. 1901.
Annual or perennial herbs or low shrubs, villous or scabrous; leaves opposite or
the uppermost alternate, the margins entire or dentate; heads discoid, with only one
or (rarely) two flowers, the heads aggregate in dense glomerules, these subtended by
herbaceous, ovate to linear bracts; involucres calyxlike, tubular, gamophyllous,
dentate; receptacle paleaceous; corolla tube narrow, the limb cylindric, 5-dentate or
5-lobate, yellow to white, pink, red, or reddish purple, exserted from the involucre;
anthers short-sagittate at the base; style branches elongated, acuminate; achenes
columnar or somewhat fusiform-falcate and attenuate toward the base; pappus of 2-
several, short, unequal scales or rudimentary.
About 10 species, in tropical America and mostly Mexican.
Only the following two are known from Central America.
Plants slender annuals 20-50 cm. tall; leaves petiolate, the blades mostly 2-5 cm. long;
corollas white (in ours), 3-4 mm. long L. mollis.
Plants stout perennials 1-2.5 m. tall; leaves sessile and usually amplexicaul, mostly 6-
20 cm. long; corollas red or pink (in ours), 12-14 mm. long L. helianthifolia.
Lagascea helianthifolia HBK. Nov. Gen. & Sp. 4: 25. 1820. L.
suaveolens HBK. I.e. Nocca helianthifolia Cass. Diet. Sci. Nat. 25:
104. 1822. N. helianthifolia var. suaveolens Robins. Proc. Amer.
Acad. 36: 468. 1901. L. helianthifolia var. suaveolens Robins, op. cit.
43: 38. 1907. Camelia (Escuintla); cardol bianco (Jutiapa). Figure
77.
Usually in rather dry places, brushy fields, slopes, rocky hills,
oak, pine, or mixed forest, 300-2,100 m.; Alta Verapaz; Chimal-
tenango; Chiquimula; Escuintla; Guatemala; Huehuetenango;
Jalapa; Jutiapa; El Quiche; Sacatepequez; Santa Rosa; Solola;
Zacapa. Central and southern Mexico; El Salvador; Honduras;
Nicaragua.
Erect, stout perennials, 1-2.5 m. tall, the stems solitary or several, stiff, leafy,
usually simple below the inflorescence, glandular-puberulent and hispid with long,
spreading, fragile hairs; leaves sessile, thick, obovate or oblong-ovate to pandurate,
mostly 6-20 cm. long, 2-9 cm. wide, acute or acuminate, rounded or shallowly cordate
at the base and usually amplexicaul, the margins coarsely and unequally serrate or
subentire, scabrous on the upper surface, densely velutinous-pilose beneath;
glomerules of heads large and suggesting a single large head, subtended by several
large, leaflike bracts; involucres sericeous-villous, about 1.5 cm. long, irregularly 4-5-
dentate, one lobe considerably longer than the others; corollas greenish white to
cream, long-exserted; achenes 6-8 mm. long, densely pilose, especially so near the
apex, enclosed in the rather persistent, calyxlike involucre; pappus rudimentary,
short, irregular.
254 FIELDIANA: BOTANY, VOLUME 24
Lagascea mollis Cav. Ann. Cienc. Nat. Madrid 6: 333, t. 44.
1803. Nocca mollis Jacq. Fragm. Bot. 58, t. 85. 1809.
Damp, grassy or brushy slopes, 600-900 m.; Chiquimula;
Jutiapa; El Progreso. Mexico; British Honduras; Honduras;
Nicaragua; West Indies; tropical South America; naturalized in
Africa, Asia, and India.
Erect, slender annuals, commonly 20-50 cm. tall, often much branched, the stems
pilose with soft, slender, spreading hairs; leaves petiolate, the blades thin, triangular-
ovate or rhombic-ovate to lance-ovate, mostly 2-5 cm. long, acute or acuminate,
rounded or acute at the base and often abruptly decurrent on the petiole, the margins
unequally dentate, rather densely short-pilose on both surfaces; glomerules of heads
about 1 cm. broad on slender, short or elongated peduncles, the bracts foliaceous,
oblanceolate-oblong, about 1 cm. long, obtuse, densely pilose; involucres about 6 mm.
long, densely pilose, the erect, subulate lobes about 2 mm. long; corollas pale purple;
achenes about 3 mm. long, glabrous or pilose near the apex; pappus reduced to an
inconspicuous crown or annulus.
LASIANTHAEA DC.
By KENNETH M. BECKER
Shrubs (in ours) or perennial herbs; leaves all opposite, lance-ovate to ovate (in
ours) or narrower, margins serrate to denticulate-mucronulate, triplinerved; heads
radiate; rays fertile, yellow (in ours), orange-red or purple, in terminal and axillary,
cymose, umbelliform clusters, these occasionally reduced to one head; involucres
campanulate to hemispherical (in ours) or narrower, 3-seriate, not or only slightly
graduate (in ours) to strongly graduate; phyllaries generally ovate, lance-ovate, or
ovate-oblong in outline, membranous (the outer often indurate) below, usually
herbaceous above; pales conduplicate; disc corollas narrow-campanulate to camp-
anulate, 5-lobate; anther thecae black (in ours) or brownish, auriculate; style
branches filiform, hispidulous dorsally; disc achenes cuneate, biconvex, flattened, the
adaxial margin narrowed out into a thin, sharp edge, no true wings present, awns 2,
stout, directly confluent with achenial margins; pappus of squamellae, these
sometimes reduced to a ciliate fringe, sometimes connate, faces not tuberculate; ray
achenes similar to disk achenes, triquetrous, 3-awned or toothed.
About 20 species, ranging from the southwestern United States
to Panama. Most are Mexican, only the following occuring in
Central America.
Lasianthaea fruticosa (L.) K. Becker, Phytologia 3: 297. 1975.
Bidens fruticosa L. Sp. PI. 833. 1753 (based on Bidens foliis ovatis
serratis petiolatis, caule fruticoso. Hort. Cliff. 399. 1737), not
Zexmenia fruticosa Rose, Contr. U. S. Nat. Herb. 1: 103. 1891.
Verbesina fruticosa L. Sp. PL ed. 2. 1271. 1763. Bidens frutescens
Miller, Gardn. Diet. ed. 8, no. 4. 1768 (Cartago, Costa Rica).
Zexmenia costaricensis Benth. in Oersted, Vidensk. Meddel. 1852:
BECKER: FLORA OF GUATEMALA 255
95. 1853 (type from Costa Rica). Z. nicaraguensis "Benth." ex C.
Muell. in Walp. Ann. Bot. 5: 226. 1858 (sphalm.) Z. villosa Polak.
Linnaea 41: 579. 1877 (type from Costa Rica). Z. elegans Sch. Bip.
ex W. W. Jones, Proc. Amer. Acad. 41: 157. 1905 (type from
Mirador, Veracruz, Mexico, Liebmann 378). Z. elegans Sch. Bip.
var. kellermannii Greenm. Field Mus. Bot. Ser. 2: 348. 1912 (type
from Los Amates, Izabal, W. A. Kellerman 7612). Narvalina
fruticosa (L.) Urb. Symb. Ant. 5: 265. 1907. Z. purpusii Brandg.
Univ. Calif. Publ. Bot. 6: 75. 1914 (type from Cerro del Boqueron,
Chiapas, Mexico, Purpus 6660). Z. costaricensis Benth. var. villosa
(Polak.) Blake, Journ. Bot. 53: 14. 1915. Z. frutescens Blake, Contr.
Gray Herb. 52: 50. 1917. Z. frutescens var. genuina Blake, torn. cit.
51. Z. frutescens var. villosa (Polak.) Blake, I.e. Z. macropoda
Blake, Contr. U. S. Nat. Herb. 22: 634. 1924. (type from Solola,
Holway 109). Vara colorada; sajdn; cambrillo; palo de escoba; flor
amarilla, taxixte (Izabal); taxiscon; tasiscobo Colorado; sos negro
(Peten, fide Lundell); sactah (Yucatan); faciscon; tisate. Figure 78.
Common in dry or moist thickets, in oak, pine or mixed forest,
often in clearings or other secondary situations, (85) 200-1,500
(2,100) m.; Alta Verapaz; Chimaltenango; Chiquimula; El Quiche;
Escuintla; Guatemala; Huehuetenango; Izabal; Jalapa; Jutiapa;
Peten; Quezaltenango; Retalhuleu; Sacatepequez;. Santa Rosa;
Solola; Zacapa. Southern Mexico to Panama (rarely adventive in
northern Venezuela).
Slender-stemmed, soft-wooded shrubs or small trees, sometimes straggling, to 3
(7) m. high, young stems varying from glabrous to densely sordid-pubescent, older
lenticellate, often light brown or whitish in color, glabrate; leaves opposite, on
petioles 0.5-4.5 cm. long, the blades sometimes rugose above, lance-ovate or more
commonly ovate, occasionally broadly ovate, 3-20 cm. long, 2-11 cm. wide, acute or
acuminate to caudate-acuminate at apex, cuneate at base, scabrous and hispid above,
in age sometimes lepidote-scabrous due to persistent hair bases, sparsely hispidulous
to pilose below, margins shallowly serrate or serrate, triplinerved; inflorescence of (1-)
3-9 headed, cymose, umbelliform clusters, terminal and from upper leaf axils,
peduncles 1-14 cm. long, slender, glabrous to hirsute or crisped- pubescent; involucres
commonly campanulate, varying to narrow-campanulate or hemispherical, 3-4-
seriate, not or only slightly graduate, 0.5-1.5 cm. high, 0.5-1.5 cm. wide; outer
phyllaries herbaceous, reticulate-veined and slightly ampliate above, indurate-
membranous below, ovate-oblong to lance-ovate in outline, acute, obtuse, or
occasionally rounded at apex, generally white-hirsute and puberulent, margins ciliate,
inner phyllaries membranous, slightly herbaceous above, ovate-oblong, acute to
rounded at apex; rays 8-14, yellow or bright yellow; disc corollas yellow; anther
thecae black; disc achenes compressed, the body 4-5 mm. high, 0.9-1.5 mm. wide,
glabrate or sparsely puberulent toward the apex, the awns (2-) 3-5 mm. long; pappus
squamellae minute, often reduced to a ciliate fringe; ray achenes with body 3-4 mm.
256 FIELDIANA: BOTANY, VOLUME 24
high, about 1.2 mm. wide, adaxial awn 2-3 mm. long, abaxial awns shorter; pappus
squamellae to 1 mm. long.
Widespread and common throughout Central America, this
species is quite variable in pubescence of stems, leaves, and
phyllaries. The stems are often used as broom handles, and
(according to Standley) the ashes from the wood are used to keep
the fingers smooth when spinning (El Salvador).
MELAMPODIUM Linneaus
References: B. L. Robinson, Synopsis of the genus Melampo-
dium, Proc. Amer. Acad. 36: 455-466. 1901; T. F. Stuessy, Revision
of the genus Melampodium (Compositae: Heliantheae), Rhodora 74:
1-70, 161-219. 1972.
Annual or perennial, herbaceous and/or suffruticose plants, glabrous, scabrous,
pilose, or sericeous, the stems decumbent to erect, dichotomously branched; leaves
opposite, decussate, the margins entire, dentate, or pinnatifid; heads solitary,
pedunculate at the ends of the branches, in the upper leaf axils, or in forks of
branches; receptacles paleaceous, convex; involucres biseriate; outer phyllaries 2-5,
often connate at the base, the inner ones indurate, each enclosing a ray achene,
closed or partially open at the apex, often extended into a hood or other appendage;
ray flowers 3-13, their spreading ligules bidentate or tridentate, yellow to creamy
white; style branches filiform, flattened, the tips obtuse, with stigmatic surfaces
throughout their length; hermaphrodite flowers 3-110, sterile, the corollas regular, the
limb 5(4)-lobate; styles undivided, or sometimes partly divided and then fused at the
apex; anthers entire at the base; ovaries capped by a disc; achenes glabrous, obovoid,
more or less laterally compressed; pappus absent.
A weedy genus of 37 species, with 12 in Guatemala.
Involucre with 2 outer phyllaries M. bibracteatum.
Involucre with 3-5 outer phyllaries.
Outer phyllaries commonly 3 (rarely some heads with 4 or 5).
Peduncles abundantly glandular- pubescent; heads 3-5.5 mm. in diameter; ray
flowers with ligules usually less than 2 mm. long M. paniculatum.
Peduncles sparsely to moderately glandular-pubescent; heads 5-15 mm. in
diameter; ray flowers with ligules usually more than 2 mm. long.
Principal leaves subauriculate to auriculate and clasping at the base; lower
leaves often hastate-pandurate M. gracile.
Principal leaves obtuse, cuneate, or abruptly attenuate to the base, neither
auriculate nor clasping; lower leaves never hastate-pandurate.
M. microcephalum.
Outer phyllaries always 5.
Leaves sericeous beneath.
Margins of outer phyllaries scarious M. linearilobum.
Margins of outer phyllaries herbaceous.
Heads 4-5 mm. high, 4-8 mm. broad; ray flowers 5-7, the ligules commonly
less than 1.5 mm. long; disc flowers 5-12 M. sericeum.
NASH: FLORA OF GUATEMALA 257
Heads 7-8 mm. high, 10-17 mm. broad; ray flowers 8-13, the ligules 4-6 mm.
long; disc flowers 50-100 M. americanum.
Leaves glabrous or more or less strigose beneath.
Leaf blades mostly 4-12 (-21) cm. long, the lower ones conspicuously perfoliate
at the base; involucres 15-32 mm. in diameter, spreading in age and
exposing the circle of large fruits, these 4-7 mm. long M. perfoliatum.
Leaf blades mostly 2-4 (-9) cm. long, the lower ones subauriculate at the base
or rather abruptly cuneate and then narrowing to a petiolar base;
involucres 5-14 mm. in diameter, spreading but not completely exposing all
of the fruits, these only 1.5-4 mm. long.
Fruits with apical appendage extended upward into a tapering, cirrhous awn
up to 8 mm. long M. longipilum.
Fruits without apical appendage.
Plants perennial, with fibrous roots and rhizomes; principal leaves sessile
or nearly so, oblong-ovate to elliptic, mostly 1.5-4 cm. long, obtuse at
the base or subauriculate; phyllaries eciliate M. montanum.
Plants annual, with tap roots; principal leaves more or less petiolate,
rarely sessile, rhombic or ovate-rhombic, usually rather abruptly
cuneate and then attenuate to the base to form a narrowly winged
petiole; phyllaries ciliate.
Ray flowers 8-13, the ligules 3.5-7 mm. long; disc flowers 40-70.
M. divarication.
Ray flowers 5-8, the ligules 1-2 mm. long; disc flowers 15-25.
M. costaricense.
Melamp odium americanum L. Sp. PL 2: 921. 1753. M.
heterophyllum Lag. Gen. & Sp. Nov. 33. 1816. M. sericeum HBK.
Nov. Gen. & Sp. 4: 272, t. 398. 1820, non Lag. 1816. M. angustifolium
DC. in DC. Prodr. 5: 519. 1836. M. nelsonii Greenm. Proc. Amer.
Acad. 41: 260. 1905.
Grassy, open places or in pine-oak forest, 1,300-1,400 m.; Baja
Verapaz; Huehuetenango; El Progreso. Southern Mexico.
Suffruticose perennials, usually branched below, the stems ascending or erect
(rarely decumbent), mostly white-pilose or strigose; leaves sessile, linear or linear-
lanceolate to lanceolate or oblong-elliptic, acute or obtuse, 3-6 cm. long, 0.2-2.7 cm.
wide, obtuse or attenuate to the base, the margins entire or sometimes 3-5-lobate, the
segments usually linear, scabrous above, usually densely white-sericeous beneath;
peduncles equalling or usually longer than the leaves; heads mostly 7-8 mm. high, 10-
17 mm. broad; involucres 7-10 mm. broad; outer phyllaries broadly ovate to rhombic,
acuminate (usually rather abruptly so), somewhat strigose near the apex, pilose near
the base; ray flowers conspicuous, 8-13, the ligules bright yellow to yellow-orange, 4-6
mm. long; disc flowers 50-100, the corollas yellow-orange; fruits 2-3 mm. long, usually
tuberculate on the angles, the hood apex usually mucronate, with a cupular, tapering
appendage to 3 mm. long.
Melampodium bibracteatum S. Wats. Proc. Amer. Acad. 26:
140. 1891.
258 FIELDIANA: BOTANY, VOLUME 24
In disturbed, wet, sandy soil, about 3,470 m., Huehuetenango,
Sierra de los Cuchumatanes, Beaman 3977. Mexico.
Annuals with ascending to decumbent, much branched stems 2-32 cm. high,
usually more or less pubescent above, glabrous below; leaves sessile, the blades
oblong to obovate or oblanceolate, mostly 1-4 cm. long, obtuse or subacute,
subauriculate to connate-clasping at the base, glabrous above, glabrous beneath or
with a few scattered hairs on the veins, the margins entire to irregularly lobate,
ciliate; heads 3.5-4.5 mm. high, 3.5-4.8 mm. broad, sessile or on peduncles to 3.5 mm.
long; involucres cupulate, outer phyllaries 2, ovate, acute, essentially glabrous,
ciliolate; ray flowers 3-6, the ligules yellow, about 1 mm. long; disc flowers 4, the
corollas yellow; fruits 2-3 mm. long, the lateral surfaces smooth, with several nerves.
Mel am podium costaricense Stuessy, Brittonia 22: 118, /. 7.
1970.
Not reported from Guatemala but collected in British
Honduras, El Cayo District, near sea level, Lundell 6107.
Erect annuals or the lateral stems sometimes decumbent, 15-35 cm. tall, glabrous
to tomentose; leaves on petioles 4-25 mm. long, the blades ovate-rhombic or the
younger ones lance-ovate, 2.5-7 cm. long, 1.5-5.5 cm. wide, acuminate or obtuse, at
the base rather abruptly cuneate and then attenuate to form a narrowly winged
petiole, sparsely strigose above, glabrous beneath or nearly so, or the sparse indument
confined to the costae and veins, the margins obscurely or coarsely serrate; heads 4-5
mm. high, on peduncles 0.5-7 cm. long; involucres cupulate; outer phyllaries 5,
connate for one-fourth to one-third their length, broadly ovate or rounded-ovate, 2.2-
5 mm. long, obtuse, glabrous, ciliate; ray flowers 5-8, the ligules yellow-orange,
elliptic, 1-2 mm. long; disc flowers 15-25, the corollas yellow; fruits about 3 mm. long,
the lateral surfaces with diagonal ridges and enlarged margins.
Melampodium divaricatum (L. Rich, ex Pers.) DC. in DC.
Prodr. 5: 520. 1836. Dysodium divaricatum L. Rich, ex Pers. Syn. PI.
2: 489. 1807, non Hort. ex DC. 1836. M. paludosum HBK. Nov. Gen.
6 Sp. 4: 237. 1820. Alcina minor Cass. Diet. Sci. Nat. 59: 243. 1829.
M. pumilum Benth. PL Hartweg. 64. 1840. M. tenellum Hook. &
Arn. var. flaccidum Benth. Bot. Voy. Sulphur 115. 1845. M.
divaricatum var. macranthum Regel in Schlecht. Linnaea 24: 198.
1851 (type grown in Bot. Gard. Turin, from seeds said to have been
collected in Guatemala by Warscewicz). M. flaccidum Benth. in
Oerst. Kjoeb. Vidensk. Meddel. 5-7: 86. 1852. M. copiosum Klatt,
Engl. Bot. Jahrb. 8: 41. 1887 (type from Alta Verapaz, Lehmann
1434). M. panamense Klatt, torn. cit. 42. Spilanthes guatemalensis
Vatke ex Donn.-Sm. Enum. PI. Guat. 1: 23. 1889, nom. nud.
Eleutheranthera divaricata Millsp. Field Mus. Bot. 1: 53. 1895.
Sacam (Huehuetenango); ik pirn (Quecchi, Alta Verapaz). Figure
79.
Damp or wet thickets, open fields or banks, village streets,
NASH: FLORA OF GUATEMALA 259
along streams in open forest, common weeds in waste or cultivated
ground, especially in coffee plantations, near sea level to 1,800 m.;
Alta Verapaz; Chimaltenango; Chiquimula; Escuintla; Guatemala;
Huehuetenango; Izabal; Jalapa; Jutiapa; Peten; El Progreso;
Quezaltenango; El Quiche; Retalhuleu; Santa Rosa; Solola;
Suchitepequez; Zacapa. Mexico; British Honduras to El Salvador
and Panama; West Indies; tropical South America.
Erect, usually rather stout annuals, often much branched, 15-100 cm. tall, the
lateral stems sometimes decumbent and rooting at the nodes, sparsely pilose or
villous or almost glabrous; leaves sessile or on narrowly winged petioles 2-20 mm.
long, the blades ovate to rhombic or the younger ones lance-ovate, mostly 2-12 cm.
long, acute, acuminate, or obtuse, usually rather abruptly cuneate at the base and
then attenuate to form a narrowly winged petiole, both surfaces sparsely strigillose or
the upper surface scabrous and the lower one hispidulous or almost glabrous,
triplinerved, the margins subentire to undulate-dentate or dentate-crenate; heads on
slender, naked peduncles 1.5-13 cm. long; involucres cupulate; outer phyllaries united
for one-third to one-half their length, in fruit forming a disc often more than 1 cm.
broad, usually tomentose near the base, the 5 lobes broadly obovate, rounded at the
apex, ciliate; ray flowers 8-13, the ligules yellow-orange, 3.5-7 mm. long, 1.6-3 mm.
wide; disc flowers 40-70, the corollas yellow-orange; fruits 3-4 mm. long, coarsely
reticulate-veined, the lateral surfaces diagonally striate and with enlarged margins.
Melampodium gracile Less. Linnaea 6: 407. 1831. M.
oblongi folium DC. in DC. Prodr. 5: 519. 1836. M. gracile var.
oblongifolium A. Gray, Proc. Amer. Acad. 5: 182. 1861. M.
microcarpum Blake, Contr. U. S. Nat. Herb. 22: 606. 1924.
Wet thickets, open, rocky ground, 350-2,000 m.; Alta Verapaz;
Huehuetenango; Peten. Mexico; British Honduras.
Erect annuals, 17-50 cm. tall, the stems usually branching sparsely to moderately
glandular-pubescent, the upper internodes often greatly elongated; leaves sessile,
mostly 5-10 cm. long, the upper and younger ones lanceolate, the principal ones
obovate-rhombic, the lower ones often hastate-pandurate, acute to acuminate, obtuse
to subamplexicaul at the base or obtuse and then abruptly narrowed into the petiolar
base, this dilated near the stem and subauriculate-amplexicaul, more or less strigillose
and usually scabrous above, scabrid beneath, the margins entire or obscurely crenate
to sinuate-dentate or irregularly cleft; heads 6-7 mm. high, on slender, naked
peduncles about equalling or longer than the leaves; involucres spreading or
shall owly cupulate, 7-10 mm. in diameter; outer phyllaries commonly 3 (rarely 4 or
5), slightly connate at the base, broadly ovate, acute to acuminate, more or less
strigose or hirsutulous; ray flowers 5-8, the ligules yellow-orange, conspicuous, ovate,
2-5 mm. long; disc flowers 30-45, the corollas yellow-orange; fruits 2.3-2.5 mm. long,
the lateral surfaces reticulate and striate, sometimes tuberculate on the angles.
Melampodium linearilobum DC. in DC. Prodr. 5: 518. 1836.
M. canescens Brandg. Zoe 5: 222. 1905. Posita (Zacapa).
Brushy plains, hillsides, sometimes in oak or pine forest, 200-
260 FIELDIANA: BOTANY, VOLUME 24
1,600 m.; Chiquimula; Huehuetenango; Jutiapa; El Quiche; Zacapa.
Mexico; El Salvador; Honduras; Nicaragua; Costa Rica.
Usually erect, rarely decumbent annuals to about 50 cm. tall, the stems slender,
usually much branched, white-pilose, strigose, or glabrate; leaves sessile, usually
deeply pinnatifid, 3-5.5 cm. long, the lobes mostly 3-7, linear, entire, obtuse, scabrous
above, usually densely white-sericeous beneath; heads 5-7 mm. high, on slender,
naked peduncles equalling or longer than the leaves; involucres about 6 (10) mm.
broad; outer phyllaries 5, slightly connate at the base, rounded-ovate to narrowly
ovate or somewhat obovate, acute or obtuse, dorsally pilose, the margins scarious;
ray flowers 7-8, conspicuous, the ligules bright yellow to yellow-orange, 2-4 mm. long;
disc flowers 45-75, the corollas yellow-orange; fruits about 3 mm. long, usually
tuberculate below, somewhat striate above, the hood apex with an appendage up to
2.5 mm. long.
Melampodium longipilum Robins. Proc. Amer. Acad. 27: 173.
1892. M. villicaule Greenm. Field Mus. Bot. 2: 345. 1912.
Dry, rocky slopes, 1,300-1,400 m.; Huehuetenango. Mexico.
Erect annuals, 7-30 cm. tall, stems branched, densely villous; leaves short-
petiolate or subsessile (rarely sessile), the blades ovate to oblong- elliptic or the
uppermost sometimes lanceolate, acute or subacute, 2-7.5 cm. long, rather abruptly
cuneate at the base, sometimes narrowing into a winged petiole, sometimes
subauriculate, appressed-pilose to tuberculate-hirsute on both surfaces, the margins
entire; heads 5-7.5 mm. high, on slender, naked peduncles 2-8 (-11.5) cm. long;
involucres cupulate, 6-9 mm. broad; outer phyllaries 5, slightly connate at the base,
broadly ovate, 3.5-5 mm. long, pilose with subappressed hairs; ray flowers 5-8, the
ligules bright yellow to yellow-orange, 3.5-5.5 mm. long; disc flowers 40-70, corollas
yellow-orange; fruits about 2.5 mm. long, the lateral surfaces tuberculate-rugose, the
apical appendage tapering, cirrhous, 1-8 mm. long.
Melampodium microcephalum Less. Linnaea 9: 268. 1834. M.
lanceolatum Sesse & Moc. Fl. Mex. ed. 2: 196. 1894, non DC. 1836.
In forest and fields, 100-2,000 m.; Huehuetenango; Peten.
Mexico.
Erect to decumbent annuals, the stems sometimes rooting at the nodes, villous
and sparsely glandular-pubescent; leaves sessile or short-petiolate, lance-ovate to
rhombic, 3-6.5 cm. long, acute or obtuse, obtuse at the base or abruptly cuneate and
then attenuate to the petiolar base, more or less strigose on both surfaces, the
margins entire to obscurely crenate or dentate; heads 3-4 mm. high, on peduncles
shorter or longer than the leaves; involucres spreading or shallowly cupulate, 5-8 (-10)
mm. in diameter, outer phyllaries 3 (rarely 4-5), slightly connate at the base, ovate,
acuminate or acute, more or less villous and glandular-pubescent; ray flowers 5-8, the
ligules yellow-orange, 2.5-4 mm. long; disc flowers 35-50, the corollas yellow-orange;
fruits about 2 mm. long, the lateral surfaces rugose-reticulate, tuberculate on the
angles.
Melampodium montanum Benth. PI. Hartweg. 64. 1840. M.
liebmanii Sch. Bip. ex Klatt, Leopoldina 23: 89. 1887. M. montanum
NASH: FLORA OF GUATEMALA 261
var. viridulum Stuessy, Rhodora 74: 191. 1972.
Open, rocky slopes in pine or pine-oak forest, 2,000-2,500 m.;
Huehuetenango. Mexico.
Erect, ascending, or decumbent perennials with fibrous roots and rhizomes, the
stems slender, rather stiff, reddish or purplish, 6-30 cm. long, villous-hirsute; leaves
sessile or subsessile, the blades ovate, oblong-ovate, or elliptic, 1.5-4 cm. long, obtuse
or rarely acute, obtuse or rarely cuneate at the base, often subauriculate; heads few,
on slender, naked, villous peduncles usually much exceeding the leaves; involucres
spreading or cupulate, outer phyllaries 5, connate for about one-fifth their length,
broadly ovate, obtuse or acute, eciliate, 4-5 mm. long; ray flowers 9-13, the ligules
yellow, the undersurfaces greenish or purplish at the apex and on the veins; disc
flowers 80-110, the corollas yellow-green; fruits 1.5-2 mm. long, the lateral surfaces
smooth to irregularly venose, sometimes tuberculate on the margins.
Stuessy separates his var. viridulum from the typical M.
montanum by the coloration of the undersurface of the ligules (light
green at apex and on veins instead of dark purple at apex and on
veins). This character, apparently conspicuous in fresh material, is
not evident in herbarium material.
Melampodium paniculatum Gardn. Hook. Lond. Journ. Bot.
7: 287. 1848. M. brachyglossum Donn.-Sm., Bot. Gaz. 13: 74. 1888
(type from Alta Verapaz, Tuerckheim 114).
Damp or wet meadows and thickets, sometimes in pine or oak-
pine forest, sometimes on sandbars along streams, common weeds of
cornfields, coffee plantations, and waste ground, 900-2,000 m.; Alta
Verapaz; Chimaltenango; Escuintla; Guatemala; Huehuetenango;
Jalapa; Quezaltenango; El Quiche; Retalhuleu; Sacatepequez; San
Marcos; Santa Rosa; Solola; Suchitepequez. Mexico; Honduras;
Costa Rica; South America.
Erect annuals, commonly 15-60 cm. tall, rarely as much as 1 m. tall, the stems
often much branched, hispid and glandular-pubescent, usually leafy; leaves sessile or
on broadly winged petioles, these sometimes dilated at the base, the blades ovate to
lanceolate, 3-9 cm. long, 1-5 cm. wide, acute or acuminate, cuneate to obtuse at the
base and then decurrent on the petiole to its often dilated or subauriculate base, the
margins subentire to coarsely and irregularly sinuate-dentate or serrate-crenate,
strigose on both surfaces, very scabrous above and somewhat scabrous beneath;
heads on glandular-pubescent peduncles 1-5 cm. long; involucres 3-7 mm. broad;
outer phyllaries 3 (4), distinct almost to the base, ovate, 2-4 mm. long, acute or
acuminate, more or less strigose and glandular; ray flowers 3-5, the ligules yellow,
ovate, inconspicuous, 1-2 mm. long; disc flowers 10-15, the corollas yellow; fruits
irregularly striate-rugose, 2-2.8 mm. long.
Melampodium perfoliatum (Cav.) HBK. Nov. Gen. & Sp. 4:
274. 1820. Alcina perfoliata Cav. Icon. PI. 1: 11, t. 15. 1791. Wedelia
perfoliata Willd. Sp. PI. 3 (3): 2335. 1803. Polymnia perfoliata
262 FIELDIANA: BOTANY, VOLUME 24
Poiret, Encycl. Method. Bot. 5: 506. 1804. Cayil (Huehuetenango).
Damp thickets and pine-oak forest, 800-1,800 m.; Guatemala;
Huehuetenango; El Quiche. Mexico; Costa Rica. Introduced into
Cuba and in the vicinity of Los Angeles, California (where it may
not have survived).
Erect annuals, the stems stout, 20-150 cm. tall, branching dichotomously,
glabrous or nearly so; leaves sessile, rhombic or rhombic-ovate, mostly 4-10 (-21) cm.
long, acute, rather abruptly cuneate and narrowing toward the dilated and
auriculate-amplexicaul base, strigose-scabrous on both surfaces or almost glabrate
beneath, the margins obscurely to conspicuously, irregularly serrate; heads 5-7 mm.
high, on peduncles 0.6-11 cm. long; involucres 15-32 mm. in diameter; outer phyllaries
5, oblong-elliptic, obtuse, connate near the base, glabrous or more or less strigose,
spreading in age, exposing the circle of large, laterally compressed fruits; ray flowers
8-13, the ligules yellow-orange, 2.5-4 mm. long; disc flowers 30-45, corollas yellow;
fruits 4-7 mm. long, the lateral surfaces smooth or more or less striate, often with a
few inconspicuous tubercles, or one or more minute, short spines near the apex.
The fruiting heads with their large, sepal-like, green phyllaries
and the whorl of fruits are suggestive of the seed pods of some
Anoda species.
Melampodium sericeum Lag. Elench. Hort. Madrid 1805;
Gen. & Sp. Nov. 32. 1816, non HBK. 1820. M. hispidum HBK. Nov.
Gen. & Sp. 4: 273, t. 399. 1820. M. sericeum var. brevipes A. Gray,
Proc. Amer. Acad. 22: 423. 1887. M. sericeum var. exappendiculatum
Robins, op. cit. 36: 459. 1901. Hierba de toro.
Usually in dry or damp fields, often a weed in cultivated
ground, rarely in open pine-oak forest, 1,100-2,000 m.; Baja Verapaz;
Escuintla; Guatemala; Huehuetenango; El Quiche. Mexico; El
Salvador.
Erect, ascending, or sometimes spreading and prostrate annuals, the stems much
branched, sparsely hirsute to pilose; leaves sessile, oblong-linear to oblanceolate,
mostly 2-5 cm. long, entire or sometimes pinnatifid with few large, broad lobes, the
unlobed leaves and the lobate divisions entire or nearly so, irregularly ciliate, strigose
on the upper surface, usually densely sericeous beneath; heads 4-5 mm. high, 4-8 mm.
broad, short-pedunculate; involucres cupulate; outer phyllaries 5, slightly connate at
the base, broadly obovate to narrowly rhombic, acute or obtuse, pilose to hispid; ray
flowers 5-7, inconspicuous, the ligules commonly less than 1.5 mm. long, yellow; disc
flowers 5-12, corollas yellow; fruits with lateral surfaces smooth and irregularly
striate or sometimes strongly verrucate, the hood apex muticous or cirrhous, the
appendage tapering, to 2 mm. long.
MELANTHERA Rohr
Reference: James C. Parks, A revision of North American and
Caribbean Melanthera (Compositae), Rhodora 75: 169-210. 1973.
NASH: FLORA OF GUATEMALA 263
Herbaceous or suffrutescent perennials, the branching stems usually erect or
ascending, sometimes weak and reclining on other plants, commonly more or less
scabrous; leaves opposite, petiolate or subsessile, the blades broadly ovate or
triangular-ovate to narrowly linear-lanceolate, triplinerved or penninerved, the
margins dentate or serrate, sometimes hastate-lobate, usually more or less scabrous;
heads homogamous and discoid, long-pedunculate, solitary or disposed in lax cymes;
involucres hemispheric; phyllaries 2-3-seriate, ovate to lanceolate, dry or the apices
herbaceous; receptacle convex or in age conic; pales rigid, striate, strongly carinate,
sharply acute to acuminate, aristate or caudate; flowers hermaphrodite, the corollas
regular, tubular, the limb white, 5-cleft; anthers black, truncate or sagittate, the
apical appendages white; style branches elongated, the appendages narrow, hirtellous;
achenes thick, indurate, obpyramidal, quadrangular to triangular, almost glabrous or
pubescent, 2-3 mm. long; pappus bristles 2-several, unequal, delicate, caducous,
arising from the center of the achene, around the narrow corolla tube.
Perhaps five or six valid species, apparently confined to the
western hemisphere, with two in Guatemala and British Honduras.
Leaf blades linear to linear-oblong; heads commonly less than 1 cm. broad.
M. angustifolia.
Leaf blades ovate to triangular, sometimes hastate-lobate; heads commonly 1-1.5 (-2)
cm. broad M. nivea.
Melanthera angustifolia A. Rich, in La Sagra, Fl. Cubana
Fan. 2: 54. 1840. M. lanceolata Benth. ex Oerst. Vid. Meddel. Kjoeb.
88. 1852. M. microphylla Steetz, Bot. Voy. Herald 156. 1854. M.
linearis Blake, Contr. U. S. Nat. Herb. 24: 30. 1922 (type from
Cristina, Izabal, Blake 7601). M. purpurascens Blake, op. cit. 22:
629. 1924. M. angustifolia var. subhastata O. E. Schulz, Rep. Sp.
Nov. 26: 109. 1929.
Wet to rather dry savannas or open pine forest, near sea level
to 700 m.; Alta Verapaz; Izabal; Peten. Florida; southern Mexico;
West Indies; Costa Rica; Panama.
Erect, ascending, or sometimes procumbent, rather sparsely branched herbs or
suffrutescent plants, to about 1 m. high, the several stems quadrangular, often
reddish or purplish, scabrous or almost glabrous; leaves subsessile or short-petiolate,
the blades linear to linear-oblong, mostly 3-9 cm. long, 0.5-1.5 cm. wide, acute to
obtuse, attenuate to the base, sometimes obscurely hastate near the base, the margins
irregularly denticulate, scabrous; heads few, commonly less than 1 cm. broad, solitary
on long, slender, naked peduncles; phyllaries ovate, 3-5 mm. long, green-tipped, at
least when young, strigose; pales oblanceolate, 4-5 mm. long, acute, the spinose tip
about 0.5 mm. long; flowers about 6.5 mm. long; achenes obpyramidal, about 2 mm.
long; pappus bristles 2-several, caducous.
Melanthera nivea (L.) Small, Fl. Southeast. U. S. 1251, 1340.
1903. Bidens nivea L. Sp. PL 833. 1753. Calea aspera Jacq. Coll.
Bot. 2: 290. 1788. Athanasia hastata Walt. Fl. Carol. 201. 1788.
Melananthera hastata Michx. Fl. Bor. Amer. 2: 107. 1803. M.
264 FIELDIANA: BOTANY, VOLUME 24
deltoidea Michx. I.e. Melanthera oxylepis DC. Prodr. 5: 545. 1836.
M. aspera (Jacq.) Small, Bull. Torr. Bot. Club 36: 164. 1909. M.
hastifolia Blake, Contr. U. S. Nat. Herb. 24: 29, /. 3. 1922 (type from
Cristina, Izabal, Blake 7601-A). M. oxycarpha Blake, op. cit. 22:
628. 1924. M. parviceps Blake, Journ. Wash. Acad. Sci. 22: 384. 1932
(type from British Honduras, El Cayo District, Bartlett 11882).
Boton bianco (Chimaltenango); Spanish needle (British Honduras).
Figure 80.
Wet to dry thickets or forest, often in second growth or a weed
in cultivated and waste ground, sea level to 2,100 m.; Alta Verapaz;
Chimaltenango; Chiquimula; Guatemala; Huehuetenango; Izabal;
Jalapa; Peten; Quezaltenango; El Quiche; Retalhuleu; Sacatepe-
quez; San Marcos; Santa Rosa; Suchitepequez; Zacapa. South-
eastern United States; Mexico; British Honduras to El Salvador
and Panama; West Indies; tropical South America.
Coarse, usually erect, perennial herbs or suffrutescent plants, 0.5-2 m. tall, rarely
more elongated and subscandent or reclining on other plants, usually much branched,
the stems more or less quadrangular, stramineous or sometimes reddish or maroon,
more or less scabrous; leaves petiolate, the blades broadly ovate to triangular-ovate
or rather narrowly triangular, mostly 5-12 cm. long, 2-8 cm. wide, acute or acuminate,
truncate to cuneate at the base or sometimes cordate, sometimes hastate-lobate near
the base, the margins serrate, triplinerved, strigose to hispidulous on both surfaces
and more or less scabrous, especially on the upper surface; heads 1-2 cm. broad, on
short or elongated, naked peduncles; phyllaries ovate to oblong-ovate or lanceolate,
acute, green-tipped or greenish for about half their length, at least at first, strigillose,
3-5 mm. long; pales oblanceolate, acuminate and narrowing into short or elongated,
spinose tips, these sometimes more or less recurved, pubescent or puberulent on the
keel and near the apex, usually conspicuously striate; corollas white, 5-8 mm. long;
anthers black, conspicuous, about 2 mm. long; achenes obpyramidal, 2-3 mm. long;
pappus bristles 2-several, caducous.
Widely distributed, abundant, weedy plants, often covering
large areas of abandoned land, said to supply good forage.
I have been unable to distinguish specimens annotated by
Parks as M. aspera from those he has determined to be M. nivea.
According to Parks, "In southeastern Mexico and adjacent
Guatemala, Melanthera nivea may intergrade with M. aspera var.
aspera but in the United States it is distinct in morphology and in
geographic range . . .". My study of Mexican, Guatemalan, and
other Central American specimens confirms the fact of the
intergradation, to such an extent that I find no constant,
distinguishing characters. By following Parks' key, one can
sometimes separate specimens from our area, but not species. He
has pointed out that in the United States, M. nivea inhabits "fairly
NASH: FLORA OF GUATEMALA 265
moist forests and borders" and that M. aspera is "often weedy,
preferring dry, rocky places . . . ." However, label data on Central
American collections does not support a presumed difference in
habitat or geographical range. The conspicuous difference in
specimens appears to be that seen in many plants: those from damp
thickets tend to have larger, thinner leaves, while those from drier,
more open locations have smaller, more scabrous leaves.
MILLERIA Linneaus
Tall, branching, viscid annuals; leaves opposite, on winged petioles, the blades
thin, the margins subentire to dentate; heads small, heterogamous, radiate, disposed
in dichotomous cymes, these becoming laxly paniculate; involucres obliquely
depressed-globose, accrescent in age and closed above the achene; phyllaries few, one
of them broad, concave, subcucullate, another smaller one opposite it, the inner 2 or
3 small, subhyaline; receptacles small, naked; ray flower 1, pistillate, fertile, the ligule
bright yellow, the limb trifid; disc flowers 4 or fewer, hermaphrodite, sterile, the
corollas tubular, regular, the broad limb very shortly 5-lobate; anthers sagittate, with
minute basal auricles, the apical appendage broadly ovate or triangular; style of the
hermaphrodite flowers undivided; achenes obovate, compressed, more or less curved,
naked, enclosed in the somewhat fleshy, persistent, green involucre.
The genus consists of a single species.
Milleria quinqueflora L. Sp. PL 1301. 1753. Olla vieja (Santa
Rosa); quesillo (Chimaltenango); pega chivo, sajan (Guatemala).
Figure 81.
Damp or wet thickets, often in second growth, rarely in forest,
common weeds in cultivated fields or waste ground, 200-1,800 m.;
Chimaltenango; Chiquimula; Escuintla; Guatemala; Jalapa; Ju-
tiapa; Peten; Santa Rosa; Zacapa. Mexico; British Honduras to El
Salvador and Panama; Cuba; northwestern South America.
Erect annual herbs, often 1 m. high, openly branched, the stems slender, brittle,
glandular-pilose; leaves very thin, the blades broadly ovate or rhombic-ovate to ovate
or the uppermost ones elliptic, mostly 5-20 cm. long and 3-15 cm. wide, sometimes as
broad as long, acute or acuminate, contracted at the broad base and decurrent to the
base of the petiole, triplinerved, thinly hirsute with whitish hairs, usually scabrous
above, the margins subentire to serrate-dentate; heads pedicellate, disposed in
dichotomous cymes, these on long, glandular-pilose peduncles, forming large, lax,
almost naked panicles, the pedicels usually nutant in fruit; involucres 3-4 mm. long;
phyllaries greenish, lance-ovate, acute or acuminate, ciliate, pilosulous; ray flowers
bright yellow, 3-4 mm. long; fruits about 5 mm. long, very irregular in shape, glabrous
or nearly so.
Called "chinchingua" in Chiapas, and the Maya name of
Yucatan is reported as "xentoloc." The plants are very viscid, the
266 FIELDIANA: BOTANY, VOLUME 24
leaves, stems, and flower heads adhering to clothing and to the hair
and feathers of animals and birds.
MONTANOA Cervantes
References: B. L. Robinson and J. M. Greenman, Montanoa,
Proc. Amer. Acad. 34: 508-521. 1899; S. F. Blake, Montanoa, in
Standl. Contr. U. S. Nat. Herb. 23: 1529-1536. 1926.
Large herbs, erect or subscandent shrubs, or trees, usually pubescent; leaves
opposite, mostly petiolate, the blades often lobate, the margins serrate, dentate, or
subentire; inflorescences cymose-paniculate; heads heterogamous and radiate;
involucres subcylindric to hemispheric; phyllaries 1-2-seriate, linear or lance-oblong to
ovate or rarely spathulate; pales carinate, enfolding the achenes, papyraceous or
subcartilaginous, acuminate or abruptly narrowed to an acute, often spinescent tip,
persistent, accrescent, and more or less squarrose in fruit, villous or glabrate;
receptacle conic; ray flowers uniseriate, neutral, the ligules white (in ours), spreading,
usually oblong and emarginate; disc flowers few-many, perfect, or the inner ones
sometimes sterile, the corollas regular, tubular, the limb 5-dentate; style branches
thickened upward, the appendage short or elongated; achenes without pappus, those
of the ray flowers abortive, those of the disc flowers rather thick, laterally
compressed, short, more or less obovoid.
Probably less than 40 species, in tropical America, with seven in
Guatemala. The genus was dedicated by Cevantes to Luis Montana,
native of Puebla, Mexico, a distinguished physician and naturalist
of his day.
Petioles broadly winged to the base, there dilated and conspicuously cordate-
amplexicaul M. pteropoda.
Petioles not winged, or if the leaf blade sometimes decurrent on the petiole, never to
the base.
Pales densely covered with conspicuous, long, silky hairs M. xanthiifolia.
Pales glabrate or variously pubescent but never with long, silky hairs.
Leaf blades deeply, palmately lobate M. hibiscifoUa.
Leaf blades not or only shallowly lobate.
Pales gradually tapering to the long, spinose tip, recurved or reflexed in fruit.
Heads disposed in large, leafy panicles; disc at anthesis about 1 cm. broad, or
less M. samalensis.
Heads ternate at the ends of branches or 1-3 in leaf axils; disc at anthesis
1.5-2 cm. broad M. echinacea.
Pales abruptly contracted into the short, spinose tip, not reflexed in fruit.
Leaves more or less angulate or very shallowly lobate, the upper surface
smooth to the touch or only slightly scabrous; disc at anthesis
commonly about 1 cm. broad; ligules of ray flowers 1.5-2.3 cm. long.
M. guatemalensis.
Leaves ovate, not angulate nor lobate, the upper surface conspicuously
scabrous; disc at anthesis commonly about 0.5 cm. broad; ligules of ray
flowers about 1.5 cm. long M. pauciflora.
NASH: FLORA OF GUATEMALA 267
Montanoa echinacea Blake, Brittonia 2: 345. 1937. Flor de
bolas.
Damp thickets or open slopes, 2,000-3,000 m., Huehuetenango
(type from southern slope of Sierra de los Cuchumatanes, above
Chiantla, Skutch 1276).
Slender, erect shrubs 1-3 m. high, the branches densely and minutely glandular
with sessile or short -stipitate glands and sparsely hirsute with multiseptate hairs;
leaves on slender, naked petioles mostly 2-4 cm. long, the blades ovate, mostly 8-14
cm. long and 4-7 cm. wide, falcate- acuminate, shallowly cordate at the base, the
margins crenate-serrate, triplinerved, pilosulous above, the lower surface glandular
and softly and densely pilose with spreading hairs; heads on peduncles 3-10 cm. long,
ternate at the ends of the branches and often 1-3 in the upper leaf axils; involucres
12-18 mm. high; phyllaries 3-seriate, densely pilosulous, subherbaceous, blackish
green, soon reflexed; disc in anthesis about 1 cm. high and 2 cm. broad; ray flowers
about 12, the ligules white, 10-18 mm. long; disc corollas white, hispidulous, about 6
mm. long; pales pilosulous dorsally except toward the apex and finely glandular,
about 1 cm. long, rather gradually narrowed into the strongly recurved or reflexed,
spinescent apex; achenes obovoid, obtusely quadrangular, blackish, glabrous, about 3
mm. long.
A very distinct species, remarkable for the few large heads on
long peduncles.
Montanoa guatemalensis Robins. & Greenm. Proc. Amer.
Acad. 34: 514. 1899. M. hexagona Robins. & Greenm. I.e. Palo santo
(San Marcos); sacapoc (Quezaltenango). Figure 82.
Usually in damp or wet thickets or forest, sometimes on open,
stony ledges, 1,300-3,300 m.; Alta Verapaz; Guatemala; Huehue-
tenango; Jalapa; Quezaltenango; El Quiche; Sacatepequez; San
Marcos; Santa Rosa (type from Jumaitepeque, Heyde & Lux 4216);
Solola. Mexico; Honduras; El Salvador.
Shrubs of 1-2 m. or sometimes trees more than 15 m. high, the stems and
branches subterete and striate to more or less angular, densely fulvous-tomentose
when young, in age glabrate; leaves on long, naked petioles, the blades thin, the
upper ones usually ovate to rounded ovate but the larger ones obtusely angulate or
very shallowly lobate, mostly 8-20 cm. long, acute or acuminate, the margins almost
entire or more or less serrate or crenate-dentate, obtuse to truncate at the base and
often abruptly contracted into a short, appendagelike portion which is united with
the blades, the upper surface smooth or slightly scabrous, almost glabrous or sparsely
pilose or pilosulous, the lower surface densely sordid- to men tose to thinly pilosulous
or glabrate; heads usually numerous, long-pedicellate, disposed in large, broad
panicles, the pedicels often recurving in fruit; phyllaries biseriate, 3-4 mm. long,
oblong to elliptic-lanceolate, acute or subacute, spreading, pilosulous; disc in anthesis
about 1 cm. broad, in fruit to 2.5 cm. broad; ray flowers 8-10, the ligules white,
commonly 1.5-2.5 cm. long; disc flowers numerous, the corollas yellow and viscid-
tomentulose; pales in anthesis pilosulous, later glabrate, more or less obovate, thin,
pale, with a very small spinose tip at the broad apex; achenes 2-3 mm. long.
268 FIELDIANA: BOTANY, VOLUME 24
The material referred here is extremely variable in the amount
of indument, but mature heads of all forms display no differences.
When in full flower at the beginning of the dry season, trees of
this species are handsome and showy from a long distance,
appearing almost covered with great masses of white heads. In this
species (and in M. pteropoda Blake), the disc flowers are sometimes
enormously distorted, developing into elongated, slender, linear or
clavate growths as much as 10 cm. long, giving the fruiting panicles
an extraordinary appearance. Dissection proved these growths to be
filled with spores, probably of one of the rust fungi.
Montanoa hibiscifolia (Benth.) Sch. Bip. ex C. Koch,
Wochenschr. Gaertn. 7: 407. 1864. Montagnaea hibiscifolia Benth.
in Oerst. Vid. Medd. Kjoeb. 1852: 89. 1852. Cajete (Escuintla); cana
rancho, vara de jaula (Guatemala); toquillo (Jalapa); xixil, quil
(Quecchi, Alta Verapaz).
Damp or wet thickets or open forest, frequently on steep, rocky
slopes, common in second growth, 400-2,500 m.; Alta Verapaz;
Chimaltenango; Chiquimula; Escuintla; Guatemala; Jalapa; Ju-
tiapa; Quezaltenango; Retalhuleu; Sacatepequez; Santa Rosa;
Solola; Suchitepequez. Mexico; Honduras and El Salvador to Costa
Rica.
Shrubs or trees 2-16 m. high, or sometimes apparently herbaceous throughout,
sparsely or much branched, the branches tomentulose or glabrate; leaves long-
petiolate, the petioles usually with 2 small appendagelike auricles at the apex, these
sometimes remote from the base of the leaf, the blades mostly 10-30 cm. long, broadly
rhombic to lance-ovate, palmately lobate, often to the middle, with usually 5 long-
acuminate lobes, commonly shallowly cordate or truncate at the base, sometimes
contracted and somewhat decurrent on the petiole, the margins crenate or dentate,
the upper surface puberulent or scaberulous, densely sordid-tomentose to puberulent
or glabrate beneath; heads numerous or few, long-pedicellate, disposed in panicles,
the disc at anthesis 7-9 mm. broad, in fruit about 2 cm. broad; phyllaries oblong-
lanceolate, subacute, tomentose; ray flowers 8-10, the ligules white, 1-1.5 cm. long,
spreading; disc flowers yellow; pales about 1 cm. long, glabrate, with a spinose tip at
the broad, emarginate apex; achenes deltoid or cuneate, rugose, about 2 mm. long,
and almost as broad at the top.
Montanoa pauciflora Klatt, Leopoldina 23: 90. 1887. Flor de
Conception (Sacatepequez); flor de pascua (Santa Rosa).
Damp or wet thickets, often in second growth, near sea level to
1,800 m.; Chimaltenango; Chiquimula; Escuintla; Guatemala;
Jutiapa; Peten; El Progreso; Sacatepequez; San Marcos; Santa
Rosa; Suchitepequez. Southern Mexico; British Honduras;
Honduras; El Salvador; Nicaragua.
NASH: FLORA OF GUATEMALA 269
Erect, arching, or scandent shrubs, often climbing over fairly large trees, the
branches slender, terete, puberulent to tomentulose or glabrate; leaves on long,
slender, naked petioles, the blades thin, more or less ovate, mostly 5-10 cm. long,
acute or acuminate, rounded or truncate at the base and often abruptly contracted
into a short, appendagelike portion, the margins serrate, not lobate, very scabrous on
the upper surface, sparsely or densely pilose beneath with mostly appressed hairs;
heads numerous, long-pedicellate, disposed in open, leafy panicles at the ends of
branches, the disc at anthesis about 0.5 mm. broad; phyllaries about 4 mm. long,
oblong-lanceolate or linear-lanceolate, pubescent; ray flowers about 8, the ligules
white, about 1.5 cm. long; disc flowers yellow; pales broad, accrescent, abruptly
contracted into a short, spinose tip; achenes about 2 mm. long, black, glabrous.
Montanoa pteropoda Blake, Proc. Biol. Soc. Wash. 37: 56.
1924. Chinchin redondo, sacapoc (Quezaltenango).
Wet to dry thickets or open forest, frequently in sandy soil in
pine-oak, oak, or Alnus forest, 1,600-3,800 m.; Chimaltenango;
Guatemala; Huehuetenango (type collected between San Martin
and Todos Santos, Nelson 3616); Quezaltenango; El Quiche;
Sacatepequez; San Marcos; Solola; Totonicapan.
Usually erect or arching shrubs, rarely weak trees, sometimes merely
suffrutescent, 1.5-4 m. tall, the stems terete, sordid-tomentose or glabrate; leaves
opposite, on broadly winged petioles mostly 1-6 cm. long, these cordate-amplexicaul
at the base, the blades rhombic-ovate to angular-ovate or triangular-ovate,
acuminate, mostly 7-15 cm. long, shallowly or moderately angular-lobate or
sometimes many of the leaves, especially the upper ones, not at all lobate, abruptly
contracted at the base and decurrent to the very base of the petiole, there dilated and
clasping, the margins usually crenate-serrate or serrate, rarely subentire, hispidulous-
puberulent and more or less scabrous on the upper surface, densely or thinly
hispidulous or pilosulous beneath; panicles lax, leafy, the heads usually numerous, on
long, slender pedicels; phyllaries oblong or oblong-ovate, about 4 mm. long, obtuse,
usually apiculate, sordid-pilosulous; pales in age thin and greatly accrescent, glabrate,
cuneate-obovate, mostly 6-10 mm. long, truncate and spinose-mucronate at the apex;
ray flowers commonly 8-10, the ligules white, 8-12 mm. long; disc flowers yellow;
achenes more or less obovoid, usually 4-angulate, 2-3 mm. long, glabrous.
Common in the mountains of central and western Guatemala,
often a weed in second growth, and so plentiful in the most
frequented parts of the country that it is difficult to understand
why it was not collected and named a century ago.
Montanoa samalensis Coulter, Bot. Gaz. 20: 49. 1895.
Damp thickets or forest, 500-1,800 m.; Retalhuleu (type from
Rio Samala, J. D. Smith 2858); Sacatepequez. Reported from
Zacatecas, Mexico, the record questionable.
Shrubs or trees, sometimes as much as 18 m. high, the young branches sordid-
tomentulose, soon glabrate, terete; leaves on slender petioles, the blades oblong-ovate
or lance-oblong, 8-24 cm. long, long-acuminate, cuneate-attenuate to the base and
270 FIELDIANA: BOTANY, VOLUME 24
often long-decurrent on the petiole, puberulent and scaberulous above, grayish
tomentulose beneath or in age glabrate or scaberulous, triplinerved, not lobate or the
lower leaves obscurely and very shallowly lobate near the base, the margins crenate-
serrate or subentire; heads few or numerous, on long, slender pedicels, disposed in
large, leafy panicles; phyllaries oblong to oblong-ovate, subacute, tomentulose; pales
greatly accrescent, mostly 10-15 mm. long, pale, puberulent or glabrate, acuminate,
the tips often recurved, with a short, spinose apex; ray flowers about 10, the ligules
white, spreading, 12-15 mm. long; disc flowers numerous, the corollas yellow, the disc
at anthesis about 1 cm. broad, but the fruiting heads 2.5-3 cm. broad; achenes about
3 mm. long, 4-angulate, glabrous.
The type of this species is a rather poor specimen, possibly
atypical, not matched by any recent collections from the Samala
Valley, where this genus is very well represented. Two specimens
from Sacatepequez, both in advanced fruit, probably represent the
same species, which must be very rare along the Samala.
Montanoa xanthiifolia Sch. Bip. ex C. Koch, Wochenschr.
Gaertn. 7: 406. 1864. M. seleriana Robins. & Greenm. Proc. Amer.
Acad. 34: 510. 1899. M. myriocephala Robins. & Greenm. torn. cit.
511. M. subglabra Blake, Contr. U. S. Nat. Herb. 22: 611. 1924 (type
from Huehuetenango, near Nenton, Nelson 3536). Tatascamite
bianco (Jutiapa).
Damp or wet thickets or open forest, often on rocky slopes, 300-
1,400 m.; Chimaltenango; Chiquimula; Escuintla; Guatemala;
Huehuetenango; Jutiapa; El Quiche; Zacapa. Southern Mexico;
Honduras; Nicaragua; Costa Rica.
Shrubs or small trees, 2-6 m. high, the branches terete, densely puberulent or
tomentose or soon glabrate; leaves on slender, naked petioles, the blades rather thin,
rhombic or rhombic-ovate and often shallowly trilobate or somewhat hastate-lobate,
acute or acuminate, cuneate at the base and sometimes short-decurrent on the
petiole, the margins more or less serrate, the upper surface hispidulous or pilosulous,
often very scabrous, grayish tomentulose beneath, often densely so, in age sometimes
glabrate; panicles small or large, lax or dense, the heads small, very numerous, on
slender pedicels; disc at anthesis 4-6 mm. broad; outer phyllaries commonly 5-7,
ovate or lance-ovate, obtuse or acute, 2-3 mm. long, pilosulous; ray flowers
commonly 3-5, the ligules white, mostly 4-5 mm. long; disc corollas cream to pale
yellow; pales very densely covered with long, silky hairs, the apex abruptly
contracted into a stramineous, lanceolate, acuminate, erect or recurved, spinose tip;
achenes dark brown, glabrous, shining, scarcely 3 mm. long.
NEUROLAENA R. Brown
References: P. A. Rydberg, N. Amer. Fl. 34: 306-308. 1927; H.
Robinson and R. D. Brettell, Tribal revisions in the Asteraceae. IV,
The relationships of Neurolaena, Schistocarpha and Alepidocline,
Phytologia 25: 439-445. 1973.
NASH: FLORA OF GUATEMALA 271
Erect, coarse herbs or shrubs, usually with rough pubescence; leaves alternate,
large, short-petiolate or sessile, the margins dentate, serrate, or denticulate and
sometimes trilobate; heads discoid or radiate, disposed in large, corymbiform panicles;
involucres campanulate; phyllaries 3-4-seriate, graduated, membranous, usually
conspicuously striate, obtuse or acute; receptacle flat, paleaceous; pales membranous,
unicostate, caducous; ray flowers (when present) pistillate and fertile, the tubes long
and slender, the ligules short, commonly less than 5 mm. long; disc flowers
hermaphrodite, fertile, their corollas regular, yellow, the tubes slender, the limb
elongated, scarcely ampliate, shortly 5-cleft; anthers blackish, minutely sagittate at
the base, apically appendaged; style branches slender, subacute, papillose or minutely
hirtellous; achenes oblong-turbinate, sometimes obscurely 4-5-costate, glabrous or
pubescent; pappus bristles numerous, 1-2-seriate, subequal, persistent.
Six species, all in tropical America, with five in Guatemala.
Heads radiate N. cobanensis.
Heads discoid.
Phyllaries acuminate, acute, or subacute.
Pedicels sordid-pilosulous with erect hairs; heads 8-10 mm. high, about 20-
flowered, 5-8 mm. across; pappus about 4 mm. long N. macrophylla.
Pedicels appressed-pubescent; heads about 13 mm. high, 60- 100-f lowered, 10-20
mm. across; pappus about 7 mm. long N. scfuppii.
Phyllaries rounded at the apex.
Leaf blades, at least the lower ones, trilobate; phyllaries more or less puberulent,
often densely so N. lobata.
Leaf blades undivided; phyllaries glabrous N. intermedia.
Neurolaena cobanensis Greenm. in Donn.-Sm. Bot. Gaz. 37:
418. 1904.
Dense, wet, mixed mountain forest, 1,300-1,600 m.; Alta
Verapaz (type from Coban, Tuerckheim 8414). Nicaragua.
Coarse perennials, 2-3 m. high, the stems striate, short-pubescent; leaves
narrowly oblanceolate, 10-30 cm. long, 1-4 cm. wide, commonly undivided, rarely the
lower ones trilobate, acuminate, attenuate to the base, the margins denticulate,
sparsely puberulent above or glabrate, minutely pubescent beneath, especially on the
veins; inflorescences corymbose-paniculate, the branches long and slender; heads
numerous, pedicellate, radiate; involucres 5-7 mm. high; phyllanes about 4-seriate,
oblong-lanceolate, acute or obtuse, puberulent; pales lance-linear; ray flowers 8-10,
the lingules 3-5 mm. long; disc flowers 20-25, the corollas about 6 mm. long; achenes
glabrous, 1.5 mm. long; pappus bristles 5 mm. long.
Apparently rare plants, represented in Guatemala by only one
collection other than the type, and in Nicaragua by only one
collection, Williams, Molina & Williams 24921, Dept. Matagalpa,
1,300-1,500 m., cloud forest area north of Sta. Maria de Ostuma,
Cordillera Central de Nicaragua.
Neurolaena intermedia Rydb. N. Amer. Fl. 34: 307. 1927.
Damp or wet, mixed forest, 1,200-1,350 m.; endemic; Alta
272 FIELDIANA: BOTANY, VOLUME 24
Verapaz (type from Coban, Tuerckheim II. 2105).
Tall perennials to 3 m. or more, the stems striate, puberulent; leaves petiolate or
the petioles often marginate to the base, the upper leaf blades lanceolate, the others
broadly lanceolate or elliptic-lanceolate, mostly 10-20 cm. long, 4-7 cm. wide, long-
acuminate, long-attenuate to the base, penninerved, the margins finely and closely
serrate, sparsely hispidulous-scabrous above, sparsely hirsutulous beneath;
inflorescences large, corymbose-paniculate, the heads numerous, the pedicels slender,
puberulent; involucres 7-9 mm. high; phyllaries 4-5-seriate, broadly oblong, 3-5-
striate, glabrous, rounded at the apex; pales linear; flowers 12-16, regular, the corollas
5-6 mm. long, the limb longer than the tube; achenes sparsely puberulent; pappus
bristles about 30, 5-6 mm. long.
Tuerckheim stated in 1908 on the type collection label, that the
species was rare, and it has been collected only once since then
(Steyermark 43915, 1942).
Neurolaena lobata (L.) R. Br. Trans. Linn. Soc. 12: 120. 1817.
Conyza lobata L. Sp. PL 862. 1753. N. lobata var. indivisa Donn.-
Sm. Bot. Gaz. 14: 27. 1889 (type from Pansamala, Alta Verapaz,
Tuerckheim 1223). Tres puntas (Izabal); mano de lagarto (Peten).
Damp or wet thickets or sometimes in oak forest, commonly in
second growth, often in cultivated fields, along brushy stream banks
and open hillsides or roadsides, sea level to 1,400 m.; Alta Verapaz;
Chiquimula; Escuintla; Izabal; Peten; El Progreso; Quezaltenango;
Retalhuleu; San Marcos; Santa Rosa; Suchitepequez. Southern
Mexico; British Honduras to El Salvador and Panama; West
Indies; northern and northwestern South America.
Erect, coarse herbs, 1-4 m. tall, usually sparsely branched, the stems striate,
sulcate, densely pubescent when young; leaves short-petiolate or almost sessile, the
almost glabrous beneath; inflorescences corymbose-paniculate; heads usually numer-
ous, discoid, about 20-flowered; involucres 5-6 mm. high; phyllaries 4-5-seriate,
acuminate, acute or cuneate at the base, often contracted and decurrent on the
petiole, the margins dentate or serrate, scabrous-hirsutulous above, densely short-
pilose beneath and often velutinous; inflorescences corymbose-paniculate, the heads
numerous, pedicellate, discoid; involucres 5-6 mm. high; phyllaries about 4-seriate,
oblong, rounded at the apex, 1-3-nerved, more or less puberulent; pales linear, obtuse,
4-5 mm. long; corollas yellow to orange-yellow, about 4 mm. long; achenes black,
essentially glabrous, about 1.5 mm. long; pappus uniseriate, the bristles 30 or more,
about 4 mm. long, yellowish white.
Common weeds of banana plantations and often abundant on
steep roadside banks, these plants are well known to the country
people in Central America as an esteemed local "remedy" for
malaria.
Neurolaena macrophylla Greenm. Proc. Amer. Acad. 39: 118.
1903. Arnica. Figure 83.
NASH: FLORA OF GUATEMALA 273
Dense, wet, mixed forest or in dense thickets, 900-1,500 m.;
Quezaltenango; San Marcos; Suchitepequez. Mexico (Chiapas).
Erect, coarse, suffrutescent plants usually 2-4 m. high, but also reported as "a
shrub or small tree" of 7.5 m., the stems pilose with short, mostly appressed hairs;
leaves short-petiolate, the blades thin, oblanceolate, mostly 15-40 cm. long and 4-10
cm. wide, long-acuminate, attenuate to the base and decurrent almost or quite to the
base of the petiole, the margins serrate, penninerved, sparsely short-pilose above,
almost glabrous beneath; inflorescences corymbose-paniculate; heads usually
numerous, discoid, about 20-f lowered; involucres 5-6 mm. high; phyllaries 4-5-seriate,
lanceolate or the outer ones ovate, acute, puberulent; corollas yellow, 6-7 mm. long;
achenes slightly puberulent above, about 2 mm. long; pappus bristles about 4 mm.
long, dirty white.
Not a common species but rather widely scattered along the
lowlands of the west coast.
Neurolaena schippii B. L. Robinson, Rhodora 37: 62. 1935.
Known only from the type collection, Camp 32 of British
Honduras-Guatemala Survey, alt. about 730 m., British Honduras,
Schipp S-735.
Erect, coarse, suffrutescent plants, 2-3 m. tall, the stems sordid-puberulent or
pilosulous; leaves short-petiolate, the blades rather thin, oblanceolate, mostly 15-45
cm. long and 4-10 cm. wide, acuminate to long-acuminate, attenuate to the base,
penninerved, the margins rather obscurely denticulate-serrate, scabrous above,
sparsely short-pilose beneath; inflorescences corymbose-paniculate; heads numerous,
discoid, 60-100-flowered; involucres 8-10 mm. high; phyllaries 4-5-seriate, the outer
ones ovate, the inner ones lanceolate to linear-oblong, acute to acuminate, more or
less puberulent; pales about 10 mm. long; corollas yellow, 6-7 mm. long; achenes
(immature) about 1 mm. long; pappus bristles brownish white, about 7 mm. long.
NOTOPTERA Urban
Reference: S. F. Blake, Notoptera, in A revision of Salmea and
some allied genera, Journ. Bot. 53: 202, 225-229. 1915.
Erect or scandent shrubs, more or less pubescent; leaves opposite, petiolate, the
blades oval, ovate or lanceolate, the margins entire or dentate; heads disposed in
terminal or axillary, cymose panicles, homogamous or heterogamous (discoid in
Guatemalan species); outermost flowers sometimes ligulate and sterile, those of the
disc hermaphrodite, fertile; involucres 2-6-seriate; phyllaries imbricate, appressed,
oblong-lanceolate to ovate, obtuse or subacute, somewhat indurate and striate;
receptacle convex; pales rather stiff, persistent, acute or obtuse, embracing the
achenes; ray flowers (when present) styliferous but sterile, the ligules oblong, yellow;
disc corollas yellow or white, straight or abruptly curved, the tube slender, the limb
funnelform, 5-dentate; style branches subobtuse or acute, papillose-hirtellous toward
the apex; anthers sagittate at the base, with ovate apical appendages; achenes oblong
to obovate, strongly compressed, glabrous or nearly so, often striate, narrowly winged
or wingless on the outer side, there with a naked or winged, short awn, on the inner
274 FIELDIANA: BOTANY, VOLUME 24
side broadly and obliquely winged above, the wing adnate to the longer pappus awn;
squamellae none.
About eight species, in Mexico, Central America, and Jamaica.
Only the following three are known in Central America.
Involucres subcylindric to narrowly campanulate; heads with about 15 flowers.
N. guatemalensis .
Involucres broadly campanulate to turbinate or hemispheric; heads with 40 or more
flowers.
Heads on pedicels mostly 4-10 mm. long (sea level to 1,200 m.) N. scabridula.
Heads sessile or on pedicels to about 3 mm. long (1,000-2,000 m.) N. brevipes.
Notoptera brevipes (Robins.) Blake, Journ. Bot. 53: 227. 1915.
Otopappus brevipes Robinson, Proc. Amer. Acad. 44: 621. 1909
(type from Chiapas).
Damp or wet, mountain thickets or oak forest, 1,100-2,000 m.;
Alta Verapaz; Baja Verapaz; Chimaltenango; Guatemala; Jalapa;
Sacatepequez; San Marcos; Solola. Mexico (Chiapas); Honduras;
Nicaragua.
Rather large woody vines or shrubs about 1.5 m. high, with arching branches, the
stems brownish, somewhat tomentulose when young; leaves on petioles 1-2 cm. long,
the blades lanceolate or lance-ovate, mostly 7-16 cm. long, long-acuminate to acute,
cuneate or rounded-cuneate at the base, rugose and with conspicuously elevated
venation, usually 3-nerved or triplinerved, sometimes penninerved on the same
branch, very scabrous on the upper surface, densely pilose beneath with short,
spreading hairs, the margins serrate or serrulate; heads usually numerous, discoid,
sessile or on pedicels to about 3 mm. long, disposed in axillary and terminal, cymose-
racemose panicles; involucres about 3.5 mm. high; phyllaries about 3-seriate,
unicostate, subobtuse or subacute, the outer ones ovate or ovate-oblong, the inner
ones oblong or lance-oblong; pales indurate, sparsely ciliate near the apex, acute or
acuminate, 3-4 mm. long; flowers white or greenish white, at least the outer ones with
corollas abruptly reflexed below the middle; achenes brownish, glabrous, about 2 mm.
long, tristriate on each side; pappus awns 2, very unequal, the inner one with a broad
wing decurrent on the achene.
This species has been reported from Guatemala as Otopappus
curviflorus Hemsl.
Notoptera guatemalensis Urban, Symb. Ant. 2: 465. 1901.
Salmea gaumeri Greenman, Field Mus. Bot. 3: 124. 1904. N.
gaumeri Greenman, op. cit. 2: 269. 1907. N. leptocephala Blake,
Proc. Biol. Soc. Wash. 34: 46. 1921.
The type, collected by Friedrichsthal, is said to have come from
Guatemala but probably is from Costa Rica. Two collections have
been made in Peten (Tun Ortiz 2204 and Contreras 5431). A
NASH: FLORA OF GUATEMALA 275
photograph of the type, formerly in the Berlin herbarium, is in the
herbarium of Field Museum. Mexico (Yucatan).
Scandent or arching shrubs, the stems striate, hispidulous to tomentose; leaves
on petioles mostly 0.5-1.5 cm. long, the blades subcoriaceous, ovate to oblong-ovate or
oblong, mostly 5-8 cm. long and 2.5-4 cm. wide, rounded at the apex or sometimes
acute to acuminate, rounded to subcordate at the base or sometimes cuneate, the
margins denticulate or subentire, very scabrous on the upper surface, densely
tomentose beneath with short, appressed or subappressed hairs; inflorescences
cymose-paniculate, the heads sessile and crowded or short -pedicellate, 7-9 mm. high,
about 15-flowered; involucres subcylindric to narrowly campanulate; phyllaries 2-3-
seriate, the outermost ones ovate, the inner ones oblong, obtuse or subacute,
puberulent or tomentulose; pales subacute, erose to erose-denticulate near the apex;
corollas erect or reflexed; achenes brown, cuneate to oblanceolate, 2.5-3.5 mm. long,
obliquely truncate, glabrous or nearly so; pappus awns unequal, the outer one 2-2.5
mm. long, winged, the wing decurrent on the achene.
The label of the type specimen of N. guatemalensis does not
bear the exact locality data, and the plant has not appeared in
Guatemalan collections until the two from Peten were made in
1972. However, the type-photograph appears to agree in detail with
N. gaumeri; Blake considered them to be probable synonyms and I
see no reason to continue to treat them separately.
Notoptera scabridula Blake, Journ. Bot. 52: 226. 1915. Figure
84.
Damp or wet thickets, often in wet pine forest, sometimes on
rather dry, brushy, rocky slopes, sea level to 1,200 m.; Chiquimula;
Guatemala; Izabal; Jalapa; Peten; Zacapa. Southern Mexico;
British Honduras; Honduras.
Suberect or arching shrubs about 2.5 m. high or often the stems elongating and
somewhat scandent, the young branches sordid-pilosulous and more or less viscid;
leaves on petioles 0.5-1.5 cm. long, the blades membranaceous or thicker, ovate to
lanceolate, mostly 5-9 cm. long and 2-6 cm. wide, acuminate, rounded at the base,
scaberulous above, densely pilose beneath with short, sordid, chiefly appressed hairs,
and somewhat glandular, the margins serrulate or subentire; inflorescences cymose-
paniculate at the ends of the branches; heads discoid, on pedicels 4-10 cm. long, these
stout, rigid, more or less pilose with brown indument; involucres turbinate-
hemispheric, about 2.5 mm. high; phyllaries 3-seriate, appressed, indurate, the outer
ones ovate-oblong, the inner oblong or oblong-lanceolate, obtuse or subacute,
ciliolate; pales indurate, carinate, acuminate, puberulent above, spreading or reflexed,
about 3 mm. long; corollas white, at least the outer ones abruptly reflexed above the
middle; achenes blackish brown, oblong-obovate, about 2 mm. long, striate, glabrous;
pappus awns very unequal, 0.7-1.5 mm. long, the inner one with a broad wing
decurrent on the achene.
The Maya name of Yucatan is "sojbac-che."
276 FIELDIANA: BOTANY, VOLUME 24
OTEIZA La Llave
By JOHN J. FAY
Herbs and shrubs, erect or sprawling and vinelike; leaves opposite, subsessile or
petiolate, the blades tri- or triplinerved, the margins more or less serrate, acuminate;
inflorescence a small or large, simple or compound, terminal cyme; heads radiate;
involucres about 3-seriate; phyllaries evenly imbricate, usually ciliate; ray flowers
commonly 8, fertile, corolla white (or perhaps sometimes yellow), the ligule tridentate
or trilobate; disc flowers 20-75, corolla yellow, lobes triangular, erect or reflexed,
throat cylindric, tube more narrowly cylindric and usually pubescent, styles swollen
near base, at mouth of nectary, branches with flattened, triangular appendages;
receptacle convex or conic; pales hyaline, flat or folded and enclosing the disc
achenes, simple or 2-3-lobate, acute and often lacerate at apex; achenes columnar or
more or less obovoid, smooth and shiny, black, glabrous, sometimes obscurely 3-5-
angled; pappus of numerous, slender, easily deciduous awns.
Three species, in Mexico and Guatemala, with only one in
Guatemala.
Oteiza ruacophlla (Donn.-Sm.) Fay, Phytologia 31: 16. 1975.
Perymenium ruacophilum Donn.-Sm., Bot. Gaz. 55: 437. 1913. Calea
insignis Blake, Contr. Gray Herb. 52: 56. 1917 (type from
Quezaltenango, Holway 817). Flor de sajoc (Quezaltenango). Figure
85.
Damp or wet, mixed mountain forest, 2,300-3,100 m.; Chimal-
tenango; Quezaltenango (type from Volcan Santa Maria, Nelson
3727); Sacatepequez; San Marcos; Totonicapan.
Erect or scrambling and vinelike shrubs to about 5 m. tall; leaves on petioles 2-3
cm. long, the blades ovate, acuminate, 9-13 cm. long, 5-9 cm. wide, cuneate to
truncate at base, strigillose-scabrid above, hirsute, strigillose or essentially glabrous
below, margins serrate, with slender, remote teeth; stems glabrous or with scattered,
spreading hairs; inflorescences terminal and from upper axils, simple or compound
cymes; peduncles glabrous; involucres cylindro-campanulate or -ovoid, 6-8 mm. high,
4.5-6 mm. wide; phyllaries firmly chartaceous, oblong, ciliate, with rounded, lacerate,
dark brown apices; ray flowers 5-8, the corollas white, with glabrous or pubescent
tube, the ligule oblong, shortly tridentate, 10-15 mm. long, slender; disc flowers 20-30,
corolla 5-6.5 mm. long, with tube 1.5-2.5 mm. long and triangular, often recurved,
lobes about 0.5 mm. long; anthers pale brown, 2-2.5 mm. long, including the oval
appendage; style branches 1-1.5 mm. long; receptacle conic; pales 6-7.5 mm. long,
unlobed; achenes 2-2.5 mm. high, about 0.5 mm. wide.
This distinctive species has been usually maintained in
Perymenium, solely it seems, because of its pappus of deciduous
awns. In sharp contrast to Perymenium, however, are its white rays
which lack conspicuous abaxial vascular bundles, the swelling near
the base of the style, shiny, striate, uncompressed achenes and pales
NASH: FLORA OF GUATEMALA 277
without keels. Although undoubtedly helianthoid, Oteiza seems
more at home in the subtribe Galinsoginae than in the Verbesi-
ninae, with Perymenium.
OTOPAPPUS Bentham
Reference: S. F. Blake, Otopappus, in A revision of Salmea and
some allied genera, Journ. Bot. 53: 229-235. 1915.
Shrubs, often scandent; leaves opposite, petiolate, the blades 3-nerved,
triplinerved, or penninerved, the margins usually serrate; heads homogamous or
heterogamous, radiate or discoid, disposed in usually small, axillary and terminal,
cymose panicles; involucres campanulate, usually with a few foliaceous bracteoles at
the base; phyllaries 3-6-seriate, the outer ones often herbaceous or herbaceous-tipped,
the inner ones indurate, obtuse or acute, commonly strigillose; receptacles almost flat
or slightly convex; pales rather narrow, persistent, enfolding the achenes; ray flowers,
when present, pistillate and fertile, the ligules yellow, 2-3-denticulate; disc flowers
hermaphrodite, fertile, tubular, yellow or orange, their style branches elongated,
subacute, papillose-hirtellous; ray achenes often trigonous and 3-winged, disc achenes
strongly compressed and with 1 or 2 wings, the wings obliquely united with the 2
awns, these unequal and more or less united with the paleaceous corona, this
composed of irregular, denticulate or lacerate, often more or less united squamellae.
The genera Salmea, Otopappus, and Notoptera are artificial
ones and it seems evident that the species now referred to them are
not well grouped and should be rearranged.
Although only four species have been collected in Guatemala, a
fifth, O. trinervis Blake, is included here, as it occurs in nearby
Chiapas, Mexico.
Heads radiate.
Leaves mostly 5-7 (-10) cm. long, acute or acuminate, very scabrous, at least on
the upper surface, usually on both surfaces; ray flowers commonly 9-17
(-28) O. scaber.
Leaves mostly 8-18 cm. long, long-acuminate (usually narrowly so), only slightly
scabrous on the upper surface, more or less strigose beneath; ray flowers
commonly 6-12.
Pedicels 10-30 mm. long; ligules mostly 5-10 mm. long O. verbesinoides.
Pedicels 2-8 mm. long; ligules mostly 2-4 mm. long O. trinervis.
Heads discoid.
Leaves penninerved; heads sessile and glomerate O. syncephalus.
Leaves mostly triplinerved or quintuplinerved above the base; heads usually
short-pedicellate (rarely some sessile), not glomerate O. glabratus.
Otopappus glabratus (Coult.) Blake, Journ. Bot. 53: 234.
1915. O. curviflorus var. glabratus Coulter, Bot. Gaz. 20: 50 1895.
Salmea curviflora var. glabrata Greenm. Field Mus. Bot. 3: 124.
1904. O. breuipes var. glabratus Robinson, Proc. Amer. Acad. 44:
622. 1909. Figure 86.
278 FIELDIANA: BOTANY, VOLUME 24
Dry to wet, often rocky thickets on plains or hillsides,
sometimes in roadside hedges, 250-1,800 m.; Chiquimula; Escuintla;
Jutiapa; Retalhuleu; Santa Rosa (type from Volcan de Jumaite-
peque, Heyde & Lux 4235). El Salvador; Honduras.
Arching shrubs, 2-3 m. high, or the branches elongating and scandent, strigose or
almost glabrous; leaves on slender petioles 1-2 cm. long, the blades usually narrowly
lanceolate, sometimes ovate-lanceolate, mostly 7-15 cm. long, 1.5-5 cm. wide,
narrowly long-acuminate, usually cuneate at the base or oblique, but sometimes
rounded, the margins serrulate, triplinerved or quintuplinerved above the base, the
upper surfaces strigillose or almost glabrous, the lower surfaces glabrous, strigose, or
sometimes short-pilose, especially on the veins; heads numerous, discoid, on pedicels
mostly 2-5 mm. long (rarely 2 or more sessile or subsessile on a pedicel), disposed in
axillary and terminal cymose-racemose panicles; involucres 2.5-3 mm. high; phyllaries
3-4-seriate, oval to oblong, striate, obtuse or rounded at the apex, ciliate; corollas
tubular, strongly reflexed; pales glabrous or nearly so, narrow, acuminate, about 5
mm. long; achenes glabrous, about 2 mm. long, rather broadly winged on only one
angle, with an awn about 2 mm. long adnate to it, the awn on the other other angle
less than 1 mm. long; corona of about 4 unequal, lacerate squamellae.
Otopappus scaber Blake, Contr. U.S. Nat. Herb. 22: 636. 1924.
In forest, often on limestone, sea level to 150 m.; Peten. British
Honduras; Mexico (Campeche and Chiapas).
Woody vines, sometimes 15 m. long, the branches densely strigillose when young;
leaves on petioles to 7 mm. long, the blades thick, oblong-ovate or elliptic- ovate,
mostly 5-10 cm. long, acute or short-acuminate, rounded at the base, the margins
serrate or serrulate, more or less lustrous above, densely tuberculate-hispidulous and
extremely scabrous, rather densely rough-hispidulous beneath; heads disposed in
terminal cymes of 3-5, the peduncles 1-4 cm. long; involucres 5-6 mm. high; phyllaries
about 5-seriate, the outermost ones spathulate, herbaceous, spreading or reflexed,
rounded or apiculate at the apex, the middle phyllaries oblong or oblong-ovate,
strigillose; ray flowers 8-16, the ligules yellow, 6-11 mm. long; disc corollas 6-7 mm.
long; pales narrow, acuminate, 8-10 mm. long, hispidulous; ray achenes trigonous,
narrowly 3-winged, 3.5 mm. long; ray pappus a cup of united squamellae; disc
achenes 2-winged, at least above; disc pappus of 2 unequal awns 1-2.5 mm. long and
of several fimbriate squamellae.
Otopappus syncephalus Donn.-Sm. Bot. Gaz. 40: 6. 1905.
Known only from the type, Cubilgiiitz, Alta Verapaz, 350 m.,
Tuerckheim 8694.
A shrub with stout, puberulent branches; leaves on petioles 1-2 cm. long, the
blades oblong-ovate, penninerved, mostly 12-17 cm. long, 4.5-6 cm. wide, short-
acuminate to acuminate, subtruncate or rounded at the base, the margins subentire
(very minutely and remotely serrate-denticulate), scabrous on both surfaces;
inflorescence cymose-paniculate; heads numerous, sessile in clusters of 3-6 and
glomerate, discoid, 5-6 mm. high, 1 5- 20-f lowered; involucres about 3 mm. high;
phyllaries 4-seriate, oval, obtuse; pales oblong, acuminate, sparsely puberulent;
NASH: FLORA OF GUATEMALA 279
corollas tubular, about 3 mm. long; achenes winged on only one angle; pappus of
irregular squamellae.
Otopappus trinervis Blake, Journ. Bot. 53: 231. 1915.
Not reported from Guatemala but to be expected there, as it
occurs in nearby Chiapas, Mexico.
Branching shrubs, the stems subterete, striate, the branches strigillose or
glabrate; leaves short-petiolate or subsessile, the blades oblong-ovate to ovate-
lanceolate, mostly 6-14 cm. long, long- acuminate, rounded at the base, the margins
more or less serrate, sparsely strigillose above or glabrate, sparsely strigose beneath;
inflorescences axillary and terminal, cymose, the heads in small clusters on densely
strigillose pedicels 2-8 mm. long; involucres 4-5 mm. high; phyllaries about 5-seriate,
ovate to oblong, the outer ones with spathulate tips; ray flowers inconspicuous, the
ligules yellow, mostly 2-4 mm. long; disc corollas 3-4 mm. long; pales narrow, acute,
about 6 mm. long; ray achenes narrowly 3-winged; disc achenes 3-5 mm. long,
broadly winged on the inner angle; coronas of lacerate, irregular squamellae.
Otopappus verbesinoides Benth. in Hook. Icon. 12: 47. t.
1153. 1873.
Damp or wet thickets or forest; often in second growth, 150-
2,300 m.; Alta Verapaz; Escuintla; Jalapa; Jutiapa; Quezaltenango;
El Quiche; Sacatepequez; San Marcos; Santa Rosa; Suchitepequez.
Suchitepequez. Southern Mexico; El Salvador and Honduras to
Costa Rica.
Woody vines or sometimes arching shrubs of 2-3 m., the branches strigose or
glabrate; leaves on petioles to 1 cm. long, the blades oblong-ovate to linear-
lanceolate, mostly 8-18 cm. long and 2-6.5 cm. wide, narrowly long-acuminate,
cuneate to narrowly rounded at the base, often lustrous above and sparsely
strigillose, sparsely strigose beneath, the margins more or less serrate or inconspic-
uously denticulate; heads disposed in terminal and axillary cymes, the 1-5 pedicels
mostly 1-3 cm. long; involucres 5-6 mm. high; phyllaries 5-6-seriate, the outer ones
ovate to oblong, strigillose, with spathulate or linear, herbaceous tips, the inner ones
4-5 mm. long; ray flowers 6-12, the ligules yellow, mostly 5-10 mm. long; disc corollas
reflexed at maturity, about 5 mm. long; pales narrow, acute, glabrous, about 7.5 mm.
long; ray achenes 3-4 mm. long, narrowly 3-winged; disc achenes very narrowly or
scarcely winged on the outer angle, broadly and obliquely winged above on the inner
angle; pappus squamellae lacerate.
PARTHENIUM Linneaus
Erect, branching annual or perennial herbs or shrubs, tomentose or scabrous;
leaves alternate, the margins entire, dentate, or pinnately dissected; heads small,
heterogamous, radiate, disposed in terminal, usually somewhat corymbiform
panicles; involucres broadly campanulate or hemispheric; phyllaries biseriate or few-
seriate, broad, appressed, dry, subequal or the outer ones gradually shorter;
receptacle small, convex or conic; pales membranaceous, subtending the hermaphro-
dite flowers; ray flowers uniseriate, pistillate, fertile, the ligules commonly white,
280 FIELDIANA: BOTANY, VOLUME 24
short, broad, bidentate or bifid; disc flowers hermaphrodite, sterile, the corolla
regular, tubular, the limb little ampliate, 4-5-dentate; anthers entire at the base; style
of the hermaphrodite flowers undivided; ray achenes dorsally compressed, carinate on
the inner side, bearing at the apex 2-3 short or elongated awns.
Perhaps a dozen species, all North American except the
following one, and mostly in Mexico, two species extending into
temperate North America. One member of the genus, P. argentatum
Gray, "guayule," native of northwestern Mexico and western Texas,
was once exploited as a source of commerical rubber and was
cultivated for that purpose in the southwestern United States.
Parthenium hysterophorus L. Sp. PL 988. 1753. Figure 87.
Weeds about dwellings or cultivated ground, sea level to 200
m.; Peten. Southern United States; common in many parts of
Mexico; British Honduras; West Indies; South America.
Erect annuals 30-75 cm. high, often much branched, strigose and often scabrous
throughout, usually grayish; leaves petiolate, the blades ovate to oblong in outline,
pinnatifid or bipinnatifid, the segments linear or lanceolate, dentate or lobate; heads
numerous, slender-pedicellate, 3-6 mm. broad; outer phyllaries broadly rhombic,
whitish with greenish tips, puberulent about the apex, the inner ones pale, scarious,
papillate on the inside and more or less ciliolate, subtending and at first enfolding the
ray achenes, later separating into 3 parts to release the achene; pales cuneate; ray
flowers few, inconspicuous, about 0.5 mm. long and less than 1 mm. broad; achenes
obovate, about 1 mm. long, black, glabrous or nearly so, crowned by the persistent
ray corolla and 2 short, flattened, obtuse awns.
The Maya name has been reported as "hauay." Common weeds
in many regions of Mexico and the West Indies, but rare in Central
America.
PERYMENIUM Schrader
By JOHN J. FAY
Reference: John J. Fay, Revision of Perymenium (Asteraceae -
Heliantheae) in Mexico and Central America, unpublished doctoral
dissertation submitted to the graduate faculty in Biology, City
University of New York. 1973.
Trees, shrubs, scrambling woody vines or herbs from 1 dm. to nearly 20 m. high;
leaves opposite, subsessile or petiolate, usually trinerved or triplinerved and serrate,
rarely penniveined or subentire; inflorescence a simple or compound terminal cyme or
a single head terminating a leafy branch; heads radiate; involucre 2-4-seriate;
phyllaries evenly or unevenly imbricate, inner ones with a spreading, ciliate apex or
apex appressed and transitional to pales; ray flowers 5-9 (-12), corolla yellow; disc
flowers 9-65, corolla yellow, lobes triangular, erect or reflexed, throat broadly or
narrowly cylindric or cylindro-campanulate, tube narrowly cylindric; receptacle
flattish or more or less convex; pales distinctly keeled along abaxial surface, enfolding
FAY: FLORA OF GUATEMALA 281
achenes; disc achenes biconvex or compressed-tetragonal, adaxial and abaxial angles
sometimes bearing wings; awnlike outgrowths sometimes present on upper adaxial
and abaxial angles (marginal awns); pappus of 15-35 deciduous awns inserted on a
more or less distinct beak or neck at apex of achene, adaxial and abaxial awns
usually longest and stoutest; ray achenes usually triquetrous, otherwise similar to
those of disc.
About 40 species from northern Mexico to Andean Peru.
Achenes winged nearly or quite to the base; the wings developed upward into awnlike
extensions (marginal awns).
Involucre 4.5-13.5 mm. high, 5-10 mm. wide.
Inner phyllaries erect or nearly so, without conspicuously spreading apex,
involucre smaller, 4.5-6 mm. high, 5-6.5 mm. wide; pales 5-6 mm. long.
P. grande var. grande.
Inner phyllaries with blunt, spreading apex, involucre larger, 6.5-13.5 mm. high,
6-10 mm. wide; pales 6.5-8.8 mm. long P. grande var. nelsonii.
Involucre 3.5-4.5 mm. high, 3-5 mm. wide.
Erect shrub or small tree with straight, ascending branches; leaves sparsely
strigillose, coriaceous P. nicaraguense.
Erect or, more frequently, sprawling and vinelike shrub with strongly divaricate,
upcurving branches; leaves strigillose, hispidulous, or hispid-pilose, herba-
ceous P. gymnolomoides.
Achenes unwinged or winged only on the upper angles, with or without distinct or
inconspicuous marginal awns.
Inner phyllaries with spreading or subspreading, yellowish, ciliate apex.
Disc corollas 5-8.5 mm. long; disc achenes 3-4 mm. long; pales 5-8 mm. long.
P. jalapanum.
Disc corollas 3-4.5 mm. long; disc achenes 2-2.5 mm. long; pales 2.5-4.5 mm. long.
P. gracile.
Inner phyllaries generally transitional to pales, without spreading, yellowish, ciliate
apex.
Phyllaries erect, acute, in 3 series, those of inner 2 series approximately equal in
length, distinctly oblong, those of outer series shorter, lance-ovate.
P. chloroleucum.
Phyllaries usually recurved and attenuate, in 3-4 more or less evenly imbricate
series, varying gradually from ovate in outer series to oblongish in inner
series P. ghiesbreghtii.
Perymenium chloroleucum Blake, Brittonia 2: 349. 1932.
Mixed forest at 1,700-2,500 m., blooming September through
December; Huehuetenango (type from Chiantla, in a deep ravine,
Skutch 1966); El Quiche. Mexico (Chiapas).
Slender-stemmed, freely-branched shrubs up to about 1.5 m. high; leaves 1.5-4
cm. long, 0.5-2.7 cm. wide, ovate or oval-ovate, acute, with cuneate or rounded to
truncate base, subentire or with about 5-8 shallow teeth on a side, sparsely to
moderately strigillose above, moderately to very densely silvery-strigillose and
distinctly lighter beneath, tri- or triplinerved, the petiole 2-11 mm. long; stem
moderately strigillose, with appressed or subspreading hairs, peduncles more densely
so; heads disposed in small (3-5-headed) cymes, usually raised on elongate internodes
282 FIELDIANA: BOTANY, VOLUME 24
terminally and from the upper axils; involucre 4.5-6.5 mm. high, 3.5-5.5 mm. wide,
campanulate or cylindro-campanulate, 3-seriate; outer phyllaries lance-ovate, acute,
moderately to rather densely strigillose and obscurely ciliate, inner two series longer,
equal or nearly so, distinctly oblong, acute or bluntish and often less densely
pubescent than outer series; ray flowers 7-9, lamina 4.5-8.5 mm. long, tube 0.9-1.2
mm. long; disc flowers 15-32, corolla 3.1-3.7 mm. long, with a narrow tube 0.6-1.1 mm.
long, cylindric or somewhat funnelform throat and deltoid, commonly reflexed lobes
0.4-0.7 mm. long; anthers with blackish thecae 1.2-1.4 mm. long and ovate
appendages 0.3-0.5 mm. long; style branches ca. 1 mm. long, including the short,
acute, indistinct appendage; receptacle flat or obscurely convex; pales 3.8-5.5 mm.
long, with erect, acute apex; disc achenes purplish, about 2.5-3 mm. high and 1-1.5
mm. wide, biconvex, sparingly tufted-strigillose; pappus of about 20 awns, the adaxial
one up to 2 mm. long; ray achenes slightly shorter and broader, otherwise similar to
those of disc.
This is apparently a rather rare species, known from fewer than
a dozen collections. It is, however, an altogether inconspicuous and
undistinguished plant in the field and its rarity in herbaria may be
due more to this than any real scarcity in nature.
Perymenium ghiesbreghtii Robins. & Greenm. Proc. Amer.
Acad. 34: 525. 1899. P. chalarolepis Robins. & Greenm. I.e. P.
purpusii Brandeg. Univ. Calif. Publ. Bot. 3: 395. 1914. P.
leptopodum Blake, Proc. Amer. Acad. 51: 523. 1916. P. inamoenum
Standl. & Steyerm. Field Mus. Bot. 23: 143. 1944.
Forests and roadsides at 650-3,800 m.; blooming year-round,
with a lull from March through June; Alta Verapaz; Chiquimula;
Guatemala; Huehuetenango; Jalapa; Quezaltenango; Solola;
Totonicapan. Mexico (Chiapas); British Honduras.
Sprawling or erect, suffrutescent plants to about 3 m. high; stems of the season
strongly 4 -grooved, frequently purplish and green-spotted, sparsely to densely strigose
or hispid; leaves 2.5-11.5 cm. long, 1-3 (-5.5) cm. wide, lanceolate or occasionally
lance-ovate, acute to attenuate, with cuneate to rounded or tuncate base, obscurely
to strongly serrate, sparsely strigillose or hispidulous above, sparsely to moderately so
below or, rarely, hispid-pilose; petiole slender, 0.5-2 cm. long; inflorescence terminal,
usually ample, freely branched, occasionally reduced to a single small cyme;
peduncles moderately to densely strigillose or hispidulous; involucre 3-seriate, 3-5
mm. high, 4.5-8 mm. wide; outer phyllaries ovate, obtuse to acute or, most frequently,
attenuate and recurved, sparsely to moderately strigillose and more or less ciliate,
inner phyllaries progressively more oblong and less pubescent, but usually retaining
recurved, attenuate apex; ray flowers 6-9 (-12), lamina 3.5-8 mm. long, tube 0.7-1.5
mm. long; disc flowers 13-36 (-49), corolla 3-5.3 mm. long, with cylindric throat,
narrow tube 0.5-1.8 mm. long and deltoid lobes 0.4-0.9 mm. long; anthers with
blackish or, rarely, light brown thecae 1-1.8 mm. long and ovate appendages 0.3-0.6
mm. long; style branches 0.7-1.5 mm. long, including the short, acute, hispidulous
appendage; receptacle obscurely to strongly convex; pales 3.8-7.5 mm. long, at least
the inner ones sharply acute, ciliate on the keel; disc achenes 1.8-3 mm. high, 0.9-1.5
mm. wide, biconvex or obscurely triquetrous, more or less ciliate on the angles, nearly
FAY: FLORA OF GUATEMALA 283
glabrous or with tufted, ascending hairs on the faces, strongly hispidulous near the
apex; pappus of about 15-35 awns, the adaxial one up to 4 mm. long; ray achenes 1.5-
3 mm. high, 0.8-1.4 mm. wide, obcompressed-triquetrous, otherwise similar to those of
disc flowers; chromosome number from meiotic material: n = 15, n = about 15.
This is an extremely variable species which has had several
names applied to it. Most frequently, it has passed as P. purpusii. A
comparison of types of all the names involved, as well as the
voluminous additional material now available in herbaria, however,
fails to demonstrate any reasonable basis for the recognition of
more than a single taxon.
P. ghiesbreghtii is probably the commonest member of the
genus in Guatemala and the only one that could be called weedy.
Perymenium gracile Hemsl. Biol. Cent. Am. Bot. 2: 181. 1881.
P. microcephalum Sch. Bip. in Klatt, Leopoldina 23: 143. 1887.
At 900-2,400 m., blooming July through October; known in
Guatemala from a single collection, Suchitepequez, Finca Moca,
Skutch 1470. Mexico.
Shrubs or possibly partially woody herbs, sparsely and minutely strigillose
throughout, height unknown; leaves ovate or lance-ovate, 3-8 cm. long, 1.5-3.5 cm.
wide, acute or attenuate, with a rounded base, shallowly serrate, with about 6-7 teeth
on a side, tri- or triplinerved, petiole 8-17 mm. long; stem distinctly 4-grooved;
inflorescence a leafy, terminal, compound cyme containing about 20-100 heads;
involucre 3-seriate, cylindro-campanulate or turbinate, 4.5-5.7 mm. high, 2.7-3.8 mm.
wide; outer phyllaries ovate, acute, ciliate at least distally, inner phyllaries oblong,
with a lax, yellowish, minutely ciliate, blunt apex; rays 5-8, lamina 6-9 mm. long,
tube 1-1.4 mm. long; disc flowers 9-13 (-19), corolla 3.8-4.2 mm. long, with a narrow
tube 0.9-1.5 mm. long, cylindric throat and deltoid, frequently reflexed lobes 0.5 mm.
long; anthers with brownish or blackish thecae 1.4-1.6 mm. long and ovate
appendages 0.4-0.5 mm. long; style branches 0.7-1.2 mm. long, including the short,
acute, barely hispidulous appendage; receptacle flat; pales 4.2-4.5 mm. long, narrowly
keeled, with an acute, erect, or incurved apex; disc achenes 2-2.5 mm. high and 1 mm.
wide, biconvex, with slender, marginal awns about 0.5 mm. long, essentially glabrous
on faces, very finely puberulent near apex; pappus of about 20 awns; ray achenes
obcompressed-triquetrous, slightly shorter and broader than, but otherwise similar to,
those of disc.
This species is quite distinctive morphologically, but its ecology
and geographical range are virtually unknown. It is one of only
three species in the genus which occur both east and west of the
Isthmus of Tehuantepec. The others are P. klattianum, which does
not reach Guatemala and whose range east of the isthmus needs
reconfirmation, and P. gymnolomoides, which is apparently restrict-
ed to moist lowlands along the Atlantic coast. It would be
interesting to know whether the single Guatemalan collection of P.
284 FIELDIANA: BOTANY, VOLUME 24
gracile, which is the only record of its occurrence east of the
isthmus, represents an isolated and disjunct population or part of a
more extensive, undocumented range.
Perymenium grande Hemsl. Biol. Cent. Am. Bot. 2: 181. 1881.
var. grande. P. turckheimii Klatt, Leopoldina 20: 95. 1884. P.
grande Hemsl. var. strigillosa Robins. & Greenm. Proc. Amer. Acad.
34: 529. 1899. P. strigillosum (Robins & Greenm.) Greenm. Field
Mus. Bot. 2: 348. 1912. 0a'ax (Quecchi, Alta Verapaz). Figure 88.
Forest at 900-3,000 m., blooming September through April; Alta
Verapaz; Chiquimula; Guatemala; Huehuetenango; Jalapa; El
Progreso; Sacatepequez; Solola. Honduras; El Salvador.
Shrubs or trees to about 20 m. high, usually smaller; leaves ovate or lance-ovate,
10-26 cm. long, 3.5-7.5 cm. wide, acute or, more often, acuminate-attenuate, base
cuneate to truncate, sparsely to moderately strigillose-scabrid above, sparsely to
densely hispidulous to hispid-pilose below, evenly serrate, triplinerved, with 2-several
pairs of prominent, parallel secondaries below the major pair, petiole 1.5-7 cm. long;
stem strongly 4 -grooved when young, very sparsely to densely strigillose to hispid-
pilose; inflorescence large and freely branched, terminal, of many small (about 3-
headed) cymes, usually about equalling the upper leaves; peduncles sparsely to
densely strigillose to hispid-pilose; involucre campanulate to hemispheric, 3-seriate,
4.5-6 mm. high, 5-6.5 mm. wide; outer phyllaries ovate to lance-ovate, sharply acute,
sparsely to moderately strigillose, with appressed to subspreading hairs, inner
phyllaries progressively less pubescent and more oblong, the innermost frequently
transitional to the pales, all phyllaries usually ciliate; ray flowers 7-8, lamina 8-13
mm. long, tube 1.6-2.3 mm. long; disc flowers about 25, corolla 4-4.5 mm. long, with
cylindric throat, slender tube 1-1.5 mm. long and deltoid lobes 0.6-0.7 mm. long;
anthers with blackish thecae 1.4-1.7 mm. long and ovate appendages 0.4-0.6 mm.
long; style branches 1-1.5 mm. long, with short, acute hispidulous appendages; pales
5-6 mm. long, with an acute to obtuse or rounded, usually erect, apex and ciliate
keel; disc achenes strongly compressed-biconvex, winged (wings 0.5-1 mm. wide) and
with well-developed marginal awns up to 2 mm. long; wings and marginal awns
frequently ciliate-lacerate, body 2.8-3.5 mm. high, excluding marginal awns, 1.5-2.6
mm. wide, including wings, faces smooth and glabrous or tuberculate and tufted-
strigillose; pappus of about 15 awns, the adaxial one longest, rarely fused to the
adaxial marginal awn, up to 2.8 mm. long; ray achenes similar to those of disc,
triquetrous or obcompressed-triquetrous, winged on all three angles, or the inner one
occasionally wingless, 2.5-3 mm. high.
This variety is restricted to a relatively small area at relatively
higher elevations than var. nelsonii. Although altitudinal limits
overlap broadly between 1,000-2,000 m., only var. nelsonii is
commonly found below 1,000 m. and only var. grande is commonly
found above 2,000 m. Seasonal separation is similarly incomplete,
with var. nelsonii reaching its peak of bloom from August to
November and var. grande peaking from November through
February.
FAY: FLORA OF GUATEMALA 285
Perymenium grande Hemsl. var. nelsonii (Robins. &
Greenm.) Fay, Phytologia 31: 16. 1975. P. nelsonii Robins. &
Greenm. Proc. Amer. Acad. 34: 529. 1899. P. latisquamum Blake,
Contr. U.S. Nat. Herb. 22: 626. 1926.
Forest at 250-2,200 m., blooming May through December;
Huehuetenango; Jalapa; Zacapa. Mexico; Honduras; El Salvador;
Nicaragua; Costa Rica.
Shrubs or small trees to about 8 m. high; leaves 8-18 cm. long, 3.5-9 cm. wide,
broadly ovate to lance-ovate, acute or, more often, acuminate-attenuate, base
cuneate to truncate or subcordate, evenly serrate, moderately to densely strigillose-
scabrid to hispidulous above, very sparsely strigillose to densely hispid-pilose or
tomentellous beneath, triplinerved, with 2-several pairs of prominent, parallel
secondaries below the major pair, petiole stout, 1-4 cm. long; stem strongly 4-grooved
when young, moderately to densely strigillose to pilose or hispid-pilose; heads in an
ample, terminal, cymose inflorescence, equalling or slightly exceeding the upper
leaves, rarely only 3-5 and subsessile at the end of a leafy branch; peduncles densely
strigillose to pilose; involucre 3-seriate, broadly campanula te to hemispheric, 6.5-13.5
mm. high, 6-10 mm. wide at anthesis; outer phyllaries ovate or lance-ovate, acute,
sparsely to moderately appressed or subspreading-strigillose, inner phyllaries
progressively less pubescent and more oblong, with strongly spreading, obtuse or
rounded, membranous apices; rays 5-10 (-12), lamina 9.5-17 mm. long, tube 1.5-3 mm.
long; disc flowers 25-65, corolla 5-8.5 mm. long, with cylindric throat, slender tube
1.5-3 mm. long and deltoid lobes 0.8-1 mm. long; anthers with blackish thecae 1.7-2.7
mm. long and ovate or lanceo-ovate appendage 0.6-0.7 mm. long; style branches 1.2-
1.8 mm. long with short, acute, hispidulous appendages; disc achenes strongly
compressed-biconvex, winged (wings 0.5-1 mm. wide) and with well-developed
marginal awns up to 1 mm. long, wings and marginal awns frequently ciliate-lacerate,
body 3-4.5 mm. high, excluding marginal awns, 1-3.5 mm. wide, including wings,
smooth and glabrous or tuberculate and tufted-strigillose on faces; pappus of about
15 awns, the adaxial one up to 4 mm. long; ray achenes similar to those of disc,
triquetrous or obcompressed-triquetrous, winged on all three angles, or the inner one
occasionally wingless, 2.5-4.5 mm. high, 1.2-3 mm. wide; receptacle slightly to strongly
convex; pales 6.5-8.8 mm. long, with acute to obtuse, usually erect apex and ciliate
keel; chromosome number from meiotic material: n = about 86; from mitotic
material: n = about 45.
This variety is far more widespread than var. grande. Its longer
and laxer inner phyllaries ordinarily mark it off well from the
typical variety, but certain Guatemalan collections display inter-
mediate morphology.
Perymenium gymnolomoides (Lessing) DC. Prodr. 5: 609.
1835. Lipotriche gymnolomoides Lessing, Linnaea 6: 408. 1831. P.
goldmanii Greenm. Field Mus. Bot. 2: 269. 1907. P. peckii Robins.
Proc. Amer. Acad. 47: 211. 1911.
Moist forests and swamps, 100-1,000 m., blooming December
through May; Alta Verapaz; Peten. Mexico; British Honduras.
286 FIELDIANA: BOTANY, VOLUME 24
Erect or sprawling and vinelike shrubs to about 2 m. in height; leaves lanceolate
or lance-ovate, acute or attenuate, with cuneate to rounded or truncate base, serrate,
with 10-20 shallow teeth on a side, tri- or triplinerved, moderately to rather densely
strigillose or hispidulous above, with curved hairs, sparsely to densely hispidulous or
hispid-pilose below, petiole 4-11 mm. long; stem 4-grooved, sparsely to densely
strigillose or hispid-pilose; inflorescence composed of small (about 3-7 headed) cymes
arising terminally and at the ends of spreading lateral branches, these departing at
nearly right angles to the main stem from the upper axils and curving upward;
pubescence of peduncles similar to that of stem, usually somewhat denser; involucre
campanulate or somewhat turbinate, (2-) 3-seriate, 3.5-4 mm. high, 3-4.5 mm. wide;
outer phyllaries ovate or oblongish, acute to acuminate or occasionally blunt,
sparsely to rather densely strigillose or hispid-pilose and distinctly ciliate, often
conspicuously resin-dotted, inner phyllaries more distinctly oblong, longer and
usually less densely pubescent; ray flowers 5-7, lamina 3.5-5 mm. long, tube 1.6-2.1
mm. long; disc flowers 20-28, corolla 2.7-4 mm. long with a narrow tube 1.1-2 mm.
long, the throat cylindric or cylindro-campanulate and deltoid, the lobes spreading or
reflexed 0.4-0.7 mm. long; corollas of peripheral disc flowers somewhat recurved and
spreading; anthers with blackish thecae 1.1-1.5 mm. long and ovate appendages 0.3-
0.5 mm. long; style branches 1-1.5 mm. long, including the short, acute, minutely
hispidulous appendage; receptacle low-convex; pales 3.2-5.2 mm. long, with ciliate
keel, apex acute and crurrpled, or blunt and erect; disc achenes biconvex, strongly
winged, wings 0.5-1 mm. wide, extending upwardly into distinct marginal awns, the
achenes 2-3 mm. high, exclusive of marginal awns, 0.9-2 mm. wide, including wings,
usually narrowed to a stipitiform base, smoothish and glabrous on the faces, minutely
puberulent near the apex, wings ciliate; pappus of about 15-25 awns, the adaxial one
up to about 2 mm. long; ray achenes obcompressed-triquetrous, winged on all three
angles, the adaxial wing frequently narrower than the other two, 1.7-2.5 mm. high,
0.9-3 mm. wide.
This species exhibits peculiarities of morphology and geographi-
cal distribution which set is apart from all other North American
members of the genus. Its peculiar branching pattern, vinelike
habit, achenes with a stalklike base, and recurved peripheral disc
flowers combine to give it an extremely distinctive appearance. It is
the only member of the genus with a considerable documented
distribution extending both east and west of the Isthmus of
Tehuantepec and one of few taxa occurring at lower elevations.
Perymenium jalapanum Standl. & Steyerm. Field Mus. Bot.
23: 144. 1944.
In and near cloud forests at 2,000-2,300 m., known to bloom
December through May; Guatemala; Jalapa (type from near
Miramundo, Steyermark 32685).
Shrubs, or perhaps largely herbaceous above, to about 2 m. high; leaves
lanceolate, 4-11.5 cm. long, 1.2-3.4 cm. wide, acute or attenuate, with a rounded-
cuneate base, serrate, with about 6-9 teeth on a side, sparsely strigillose on both
surfaces, triplinerved, petiole 6-15 mm. long; stem strongly 4-grooved, very sparsely to
moderately strigillose; heads borne terminally and from upper axils in cymes of 3-10;
FAY: FLORA OF GUATEMALA 287
involucre campanulate, 6.5-8.5 mm. high, 5-6 mm. wide, 3-seriate; outer phyllaries
ovate, acute, essentially glabrous on the back and ciliate, progressively more oblong
and rounded or blunt inward, the innermost with a lax, yellowish, ciliate apex; ray
flowers 8, lamina about 7.5 mm. long, tube about 2.5 mm. long; disc flowers about 25,
corolla 5-6.5 mm. long, with a narrow tube 1.2-2 mm. long, cylindric throat and
narrowly deltoid lobes about 1 mm. long; anthers with blackish thecae 2-2.5 mm. long
and acute appendages 0.5-0.6 mm. long; style branches 1.5-2.3 mm. long, including the
short, acute appendage; receptacle flat; pales 7-7.5 mm. long, narrowly keeled, with
an acute, often inflexed apex; disc achenes 3-3.5 mm. high, 1.1-1.3 mm. wide,
biconvex, slightly tuberculate and tufted-strigillose on the faces, more densely so near
the apex; pappus of about 15 awns, the adaxial one up to 2 mm. long; ray achenes
triquetrous, slightly shorter and wider, otherwise similar to those of disc.
This is a very obscure species, known only from three
collections. The occurrence in and near cloud forest is quite unusual
for the genus.
Perymenium nicaraguense Blake, Contr. U. S. Nat. Herb. 22:
624. 1924.
Forests at 600-2,000 m., blooming all year; Chiquimula;
Guatemala; Jalapa; Zacapa. Honduras; Nicaragua.
Shrubs or small trees to about 5 m. high; leaves coriaceous, oval or ovate to
lance-oval or lanceolate, acute or obtuse, with a cuneate or truncate base, subentire
or shallowly serrate with about 6-20 teeth on a side, usually triplinerved, frequently
with 2-several pairs of prominent, parallel secondaries below the major pair, rarely
penniveined, both surfaces very sparsely fine-strigillose, with strongly appressed hairs
along the veins only, minutely bullate beneath, with impressed veinlets, petiole 2-20
mm. long; stem 4 -grooved, at least when young, very sparsely to moderately
strigillose; inflorescence of few to numerous, small (about 3-6-headed) cymes, borne
terminally and from the upper axils, exceeded by or slightly exceeding the upper
leaves, rarely very ample, with about 200 heads in a compound terminal cyme;
peduncles moderately strigillose, with appressed or subspreading hairs; involucre
turbinate-campanulate, 3.5-4.5 mm. high, 3-5 mm. wide, 3-4-seriate; outer phyllaries
ovate, acute, moderately strigillose, progressively less pubescent, more oblong and
blunter inward, obscurely ciliate toward the base, with hairs no longer than those on
the back of the phyllary; ray flowers 7-9, lamina 3.5-7.1 mm. long, tube 0.7-2 mm.
long; disc flowers 10-30, corolla 3.8-5.1 mm. long, with a narrow tube 1.1-1.7 mm.
long, cylindric throat, and broadly or narrowly deltoid lobes 0.5-0.9 mm. long; anthers
with blackish thecae 1-1.8 mm. long and ovate or lance-ovate appendages 0.3-0.5 mm.
long; style branches about 1 mm. long, including the indistinct, acute appendage;
receptacle low-convex; pales 4.5-6 (-7.3) mm. long, with an acute, erect or inflexed
apex; disc achenes biconvex, narrowly winged (wing to 0.3 mm. wide) or, rarely,
merely callose-margined on the angles, with distinct, flat, frequently dissected,
marginal awns up to 1 mm. long, 2-2.8 mm. high, exclusive of marginal awns, 0.9-1.5
mm. wide, including wings; pappus of about 15-20 awns, the adaxial one up to about
2 mm. long.
Collections of this species from Guatemala and Honduras have
generally been referred to either P. grande or P. purpusii (the latter
288 FIELDIANA: BOTANY, VOLUME 24
is here treated as a synonym of P. ghiesbreghtii). Nevertheless, P.
nicaraguense is a highly distinctive species and apparently not
uncommon.
PHILACTIS Schrader
Reference: Andrew M. Torres, Revision of the genus Philactis
(Compositae), Brittonia 21: 322-331. 1969.
Erect, branching shrubs with glabrous or pilose stems; leaves opposite, petiolate,
the blades triangular-ovate to ovate, glabrous or pubescent, the margins dentate;
inflorescences terminal, cymose; heads pedunculate, heterogamous, radiate; in-
volucres hemispherical; phyllaries 2-3-seriate, spreading in age; receptacle conical;
pales uncinate or acuminate at the apex; ray flowers pistillate, fertile, the ligules first
yellow, becoming reddish or maroon in age, sessile on the achenes, the apex 2-3-
lobate; style branches recurved; ray achenes trigonous with one pappus awn; disc
flowers numerous, fertile, yellow; anthers sagittate at the base, the apex appendaged;
disc achenes 2-4-angles, pappus of 1-4 unequal or sometimes subequal awns.
Three species, limited to southern Mexico and Guatemala. Only
one is known in Guatemala but a second, from nearby Chiapas, is
also treated here.
Leaves glabrous; pales exceeding the disc flowers by about 2 mm P. nelsonii.
Leaves sparsely strigose, especially on the veins; pales about equalling or scarcely
exceeding the disc flowers P. liebmannii.
Philactis liebmannii (Klatt) Blake, Contr. U. S. Nat. Herb.
26: 240. 1930. Zinnia liebmannii Klatt, Leopoldina 23: 89. 1887.
Sanvitaliopsis liebmannii Sch. Bip. ex Klatt, I.e., pro. syn.
Grypocarpha liebmannii Blake, Contr. Gray Herb. 52: 35. 1917.
Melanthera fruticosa Brandg. Univ. Calif. Publ. Bot. 10: 421. 1924.
Figure 89.
About 900 m.; Santa Rosa. Mexico.
Erect, branching shrubs to about 1 m. tall, the stems sparsely pubescent; leaves
petiolate, the blades ovate to triangular-ovate, mostly 6-12 cm. long, acuminate or
acute, almost truncate at the base and then abruptly cuneate, sparsely strigose above
and sometimes somewhat scabrous, strigose beneath, especially on the costae and
veins, the margins dentate; peduncles 2-7 cm. long, striate, more or less pubescent
near the apex; heads few; involucres about 1 cm. broad; phyllaries 2-3-seriate,
unequal, lance-ovate, subulate, the apices squarrose; pales acuminate, often
uncinate, scarcely exceeding the disc flowers; ray flowers 13-20, the ligules yellow; ray
achenes about 2 mm. long, with one awn about 2 mm. long; disc achenes about 2.4
mm. long, flattened or trigonous, sparsely pubescent, the 2 (-3) awns unequal or
sometimes subequal.
Philactis nelsonii (Greenm.) Blake, Contr. U. S. Nat. Herb.
26: 241. 1930. Gypocarpha nelsonii Greenm. in Sargent, Trees &
Shrubs 1: 145, t. 73. 1903. Sanvitaliopsis nelsonii Greenm. Proc.
Amer. Acad. 41: 261. 1905.
NASH: FLORA OF GUATEMALA 289
Shrubs 1: 145, t. 73. 1903. Sanvitaliopsis nelsonii Greenm. Proc.
Amer. Acad. 41: 261. 1905.
Not reported from Guatemala but included here as it occurs in
nearby Chiapas, Mexico.
Erect, branching shrubs to about 1 m. tall, the stems glabrous; leaves petiolate,
the blades triangular-ovate, acuminate or acute, mostly 4-10 cm. long, glabrous, the
margins serrate-dentate; peduncles striate, to 4 cm. long; involucres about 1 cm.
broad; phyllaries 2-3-seriate, lanceolate, subulate, the apices more or less squarrose;
pales exceeding the disc flowers by about 2 mm.; ray flowers 15-22, the ligules yellow,
about 9 mm. long; ray achenes about 4 mm. long, with one awn, about 3 mm. long;
disc corollas about 4 mm. long; disc achenes about 3 mm. long, ciliate on the angles,
with (1-) 2-4 unequal awns.
PODACHAENIUM Bentham
Shrubs or trees, the stems tomentose; leaves opposite, or the uppermost ones
sometimes alternate, long-petiolate, the blades large, membranaceous, the principal
ones more or less angulate-lobate, triplinerved, tomentose; inflorescences corymbose-
paniculate; heads radiate, appearing subglobose in fruit; involucres short; phyllaries
2-3-seriate, narrow, herbaceous, the outermost ones shortest; receptacle convex; pales
complicate, embracing the flowers; ray flowers uniseriate, pistillate, fertile, the ligules
white, spreading, the apex entire or 2-3-dentate; disc flowers numerous, hermaphro-
dite, fertile, the corollas tubular, yellow, the limb longer than the tube, with 5 short
lobes or teeth; anthers obtuse, obscurely sagittate at the base; style branches of the
hermaphrodite flowers complanate toward the apex, with very short appendages; ray
achenes dorsally compressed, disc achenes laterally compressed, the margins
sometimes essentially entire but often with rather narrow cartilaginous wings, these
entire or dentate or lacerate, more or less contracted at the base into a winged stipe;
pappus of 2-several, broad, lacerate squamellae, one or two of these sometimes
elongated and appearing awnlike.
A single species is known.
Podachaenium eminens (Lag.) Sch. Bip. Flora 44: 557. 1861.
Ferdinanda eminens Lag. Gen. & Sp. Nov. 31. 1816. P. paniculatum
Benth. in Oerst. Vid. Medd. Kjoeb. 1852; 99. 1852. Sacapoc
(Quezaltenango); tatascamite (Zacapa). Figure 90.
Damp or wet thickets or wooded ravines, sometimes in second
growth, 300-2,600 m. (most common at middle elevations); Alta
Verapaz; Chimaltenango; Guatemala; Jalapa; Peten; El Progreso;
Quezaltenango; Retalhuleu; Sacatepequez; San Marcos; Santa
Rosa; Solola; Zacapa. Mexico; British Honduras; Honduras; El
Salvador; Nicaragua; Costa Rica.
Coarse shrubs or small trees to about 8 m. high and with a trunk 10 cm. in
diameter, the branches stout, densely soft-tomentose; leaves on petioles mostly 4-14
cm. long, the blades thin, broadly ovate to suborbicular, often broader than long, the
upper ones commonly 6-14 cm. long and broad, the margins subentire, the lower ones
290 FIELDIANA: BOTANY, VOLUME 24
frequently as much as 30 cm. long and broad, the margins shallow ly angulate-lobate,
both surfaces tomentose, but more densely so beneath; inflorescences terminal,
broadly paniculate; heads numerous; phyllaries very unequal, lance-oblong, acute or
subacute; pales thin but stiff; acute; ray flowers commonly 12 or more, the ligules
white, spreading or drooping, 8-15 mm. long; disc corollas yellow, 2-3 mm. long;
achenes 1-1.5 mm. long; pappus 0.5-1 mm. long.
Handsome plants, in general appearance much like Montanoa,
but in the barrancos of the Occidente, where it is abundant, it
blooms after the Montanoas have ceased flowering.
POLYMNIA Linneaus
Reference: J. R. Wells, A taxonomic study of Polymnia
(Compositae), Brittonia 17: 144-159. 1965.
Perennial or annual herbs (usually large and coarse), shrubs, or trees, the stems
glabrous, scabrous, or villous, often viscid; leaves opposite or the uppermost
alternate, usually petiolate, rarely sessile, the blades usually large, the margins entire,
angulate, lobate, or pinnate-lobate, the petioles frequently winged, often dilated and
clasping at the base; heads solitary to many, heterogamous, radiate, often disposed in
corymbiform panicles; involucres hemispheric or spreading; phyllaries biseriate, the
outer ones spreading, foliaceous, the inner ones in age embracing the ray achenes;
receptacle flat or more or less convex; pales complicate, embracing the disc flowers;
ray flowers usually uniseriate, rarely biseriate, piltillate, fertile, the tube often
pubescent, sometimes densely so near the base, the ligules yellow, white, or orange-
purplish, funnelform, the limb ampliate, 5-dentate, sometimes contracted below the
teeth; styles bifid but the short branches erect and sometimes appearing undivided;
anthers minutely sagittate at the base, the apical appendage ovate; ray achenes
thick, glabrous, obovoid to spherical, sometimes slightly compressed laterally; pappus
wanting.
About 20 species, all but one American, ranging from southern
Canada to Argentina, mostly in temperate regions, with four in
Guatemala.
Principal leaves triangular to broadly ovate in outline or halberd-shaped, the petioles
(at least of the middle and lower leaves) winged to the base and there more or
less dilated and clasping; peduncles villous, hisped, or stipitate-glandular,
sometimes glandular and villous; mature achenes obovoid.
Leaves deeply pinnate-lobate with mostly 5 lobes, the sinuses at least halfway to
midvein; heads small, the disc 5-8 mm. in diameter, the outer phyllaries 5-8
mm. long; ligules or ray flowers commonly orange-red (rarely pale yellow or
reddish purple), about 6 mm. long P. oaxacana.
Leaves mostly palmate-lobate or angulate, if pinnate-lobate the sinuses relatively
shallow, not extending halfway to midvein; heads larger, the disc commonly 8-
15 mm. in diameter, the outer phyllaries 8-16 mm. long; ligules or ray flowers
bright yellow, mostly 10-15 mm. long.
Outer phyllaries eciliate, the inner ones with involute margins producing a
conspicuously elongated, aristate, stipitate-glandular point 0.5-1 cm. long
NASH: FLORA OF GUATEMALA 291
above the broad basal portion, this lower part also conspicuously stipitate-
glandular with glands up to 1 mm. long P. riparia.
Outer phyllaries ciliate, the inner ones not with involute margins, acuminate but
not to an aristate point, if the basal portion glandular, the glands sessile or
nearly so.
Peduncles densely villous to hispid and sometimes sparsely glandular.
P. maculata.
Peduncles densely stipitate-glandular P. maculata var. adenotricha.
Principal leaves narrowly lanceolate to lance-ovate, the petioles not winged (the base
of the leaf blade sometimes abruptly contracted and cuneate to short-decurrent
on the petiole but this never extending to the base); peduncles glabrous; mature
achenes spherical P. quichensis.
Polymnia maculata Cav. Icon. 3: 14. t. 227. 1795. P. maculata
var. vulgaris Blake, Contr. Gray Herb. 52: 33. 1917. P. maculata
var. hypomalaca Blake, I.e. Ax (Quecchi, Alta Verapaz); chocotorro
(Santa Rosa); mirasol (Quezaltenango, Retalhuleu). Figure 91.
Damp or wet thickets, mixed forest, sometimes in oak or pine
forest, often a weed in cultivated ground or second growth, 200-
3,000 m.; Alta Verapaz; Baja Verapaz; Chimaltenango; Chiquimula;
Escuintla; Guatemala; Jalapa; Jutiapa; Peten; El Progreso;
Quezaltenango; El Quiche; Retalhuleu; Sacatepequez; San Marcos;
Santa Rosa; Solola; Suchitepequez; Zacapa. Southern Mexico;
British Honduras to El Salvador and Panama.
Erect, coarse herbs, 1-3 (-5) m. tall, often much branched, the stems purple-
spotted, sparsely or densely short-villous and sparsely glandular; uppermost leaves
usually sessile or on winged petioles (rarely the petioles not or scarcely winged), the
principal and lower leaves on long, winged petioles, these commonly dilated and
clasping at the base, sometimes perfoliate, the blades triangular to broadly ovate or
halberd-shaped in outline, mostly 12-30 (-45) cm. long, often palmate-lobate or
angulate, the lobes acute or acuminate, the blades triplinerved, usually abruptly
contracted and decurrent to form the broadly winged petiole, the margins commonly
very coarsely and irregularly dentate, sometimes subentire, sparsely villous-
hispidulous above and scabrous, short-pilose to villosulous beneath and often
canescent; heads usually numerous and then forming large, leafy, corymbiform
panicles, the long peduncles and pedicels usually hispid-pilose or villous, rarely
glabrous; outer phyllaries 5 (6), broadly ovate to lanceolate, (5-) 8-16 mm. long and 5-
10 mm. wide, very obtuse to acute, often unequal, ciliate, essentially glabrous or more
or less villous; disc commonly (8-) 10-15 mm. in diameter; ray flowers 15-20, the
ligules yellow, 1-2.5 cm. long, tridentate; disc flowers numerous, the corollas yellow;
achenes black, obovoid, striate, 4-5 mm. long.
Polymnia maculata var. adenotricha Blake, Contr. Gray
Herb. 52: 34. 1917.
Damp or wet thickets, sometimes in pine-oak forest, 1,000-2,000
m.; Guatemala; Jalapa; Quezaltenango; Santa Rosa (type from
292 FIELDIANA: BOTANY, VOLUME 24
Chupadero, Heyde & Lux 3807); Totonicapan. Southern Mexico;
Honduras; Costa Rica.
In all respects like the typical variety except that the peduncles
and upper branches are covered with stipitate glands as well as
being villous or hispid-pilose. Ranging almost as widely as P.
maculata and apparently of scant taxonomic significance.
Polymnia oaxacana Sch. Bip. Leopoldina 23: 89. 1887. P.
nelsonii Greenm. Proc. Amer. Acad. 39: 99. 1903.
In forest and thickets, 800-1,700 m.; Alta Verapaz; Baja
Verapaz; Huehuetenango; El Quiche. Southern Mexico; Honduras.
Erect, sparsely branched herbs to 2 m. tall, the stems slender, terete, striate,
often purplish, densely glandular and sparsely or densely villous with multiseptate
hairs; uppermost leaves sometimes sessile or subsessile, the principal leaves on long or
short, broadly winged petioles, these dilated at the base, auriculate, amplexicaul,
often perfoliate, the blades mostly 7-20 cm. long, more or less triangular in outline,
deeply pinnate-lobate, with usually 5 lobes, the lobes or segments triangular to
lanceolate, acuminate, sinuate-dentate, the upper surface dark green and very
scabrous, the lower surface paler and hispidulous; heads few, on long, densely
stipitate-glandular peduncles; disc 5-8 mm. in diameter; outer phyllaries 5, ovate or
lance-ovate, mostly 6-11 mm. long and 4-6 mm. wide, acute to acuminate, green,
scabrous and villosulous, ciliate, spreading in age; ray flowers about 10, the ligules
commonly orange, sometimes yellow or reddish purple; mostly 8-10 mm. long,
tridentate; disc flowers numerous, yellow; achenes dark brown or black, obovoid,
striate, about 3 mm. long.
Polymnia quichensis Coulter, Bot. Gaz. 20: 48. 1895. P.
latisquama Blake, Journ. Wash. Acad. Sci. 16: 421. 1926. Arnica
(San Marcos); carricillo (Jalapa).
Dense, wet, mixed forest or thickets, 2,000-3,000 m.; Huehue-
tenango; Jalapa; El Quiche (type from Chiul, Heyde & Lux 3375);
San Marcos. Costa Rica.
Coarse herbs, 1-3 m. tall, erect or reclining, sparsely branched, the stems terete,
usually glabrous, sometimes sparsely puberulent; leaves on short petioles, these not
winged nor dilated at the base, the blades narrowly lanceolate to ovate-lanceolate,
mostly 9-30 cm. long, 3-10 cm. wide, acuminate to narrowly long-acuminate, usually
oblique at the base, rounded and then abruptly cuneate, briefly decurrent on the
petiole, mostly triplinerved, sometimes quinqueplinerved, conspicuously reticulate-
veined, appearing glabrous but more or less scabrous on costae and veins of both
surfaces and sometimes with scattered, very short hairs elsewhere, the margins
subentire to irregularly dentate, sometimes sinuate-lobate; heads very few, mostly
solitary, on long, slender, glabrous peduncles; disc 1-1.5 cm. broad; outer phyllaries
broadly ovate to broadly lanceolate, acute, 1.5-2.5 cm. long, mostly 1-1.7 cm. wide,
green, glabrous or nearly so, sometimes puberulent near the apex, the margins
obscurely and minutely ciliolate; ray flowers 7-15, the ligules tridentate, yellow, 1.5-
NASH: FLORA OF GUATEMALA 293
2.5 cm. long; disc flowers numerous, yellow; mature achenes spherical, black, striate,
about 5 mm. long and broad.
Polymnia riparia HBK. Nov. Gen. & Sp. 4: 282. 1820.
Wet or damp thickets and forest, 1,200-2,400 m.; Quezal-
tenango; San Marcos. South America.
Erect, perennial herbs or sometimes shrubby, the stems to 4 m. tall, glabrous to
villous, often purple-spotted; leaves sessile or on winged petioles, these narrowing
toward the stem but dilated at the base, the blades mostly 10-30 (-50) cm. long and 5-
30 cm. wide, the uppermost ones ovate to ovate-lanceolate or rhombic-ovate, the
principal (middle and lower) ones triangular to broadly ovate in outline, often lobate,
the lobes acute and the margins callose-dentate, the petiolar bases also often
irregularly lobate with shallow sinuses, scabrous to sparsely pilose above, usually
essentially glabrous beneath in ours, but said to be sometimes pilose or velutinous;
heads solitary or several; peduncles villous and/or often stipitate-glandular; outer
phyllaries (4) 5, broadly ovate to oblong-oval or oblanceolate, obtuse, mostly 8-15
mm. long and 6-10 mm. wide, usually glabrous, eciliate, rarely puberulous or
glandular-pilose, the inner phyllaries commonly 10-14, with involute margins
producing a conspicuously elongated, aristate, stipitate-glandular point, 0.5-1 cm.
long above the broad basal portion clasping the achenes, this lower portion also
conspicuously stipitate-glandular; disc commonly 8-10 mm. in diameter; ray flowers
uniseriate, commonly 12-20, the ligules yellow, about 10 mm. long; disc flowers about
30, yellow, or in age reddish brown to purplish; achenes obovoid, dark brown, striate,
3-4 mm. long.
RENSONIA Blake
Erect shrubs or small trees; leaves opposite, petiolate, scabrous, penninerved or
obscurely triplinerved, the margins serrate; heads small, pedicellate, heterogamous,
radiate, disposed in terminal, cymose panicles; involucres turbinate-campanulate,
biseriate; phyllaries 8-9, oblong-obovate, indurate below, thick-herbaceous above;
receptacles small, flat; outer pales almost flat, the inner ones narro , complicate; ray
flowers about 8, uniseriate, pistillate, fertile, the ligules bright yellow, bidentate; disc
flowers about 20, hermaphrodite, sterile, the corollas tubular, yellow, the slender tube
much longer than the funnelform limb, this 5-dentate; anthers obtuse at the base,
with an ovate apical appendage; style branches of the pistillate flowers elongated,
those of the hermaphrodite flowers short, thickened and hispidulous toward the apex;
ray achenes obovate, obcompressed, without pappus but 2-winged, the wings narrow,
entire or lacerate above, prolonged above the achene into 2 triangular, awnlike,
lacerate teeth; sterile disc achenes trigonous, wingless, their pappus a short, thick,
entire, hispidulous crown, with or without a single short, slender awn.
The genus contains a single species. It was named for Dr.
Carlos Renson, native of Belgium, who was associated with the
Department of Agriculture of El Salvador for almost 50 years, and
made the earliest large collection of Salvadorean plants.
Rensonia salvadorica Blake, Journ. Wash. Acad. Sci. 13: 145.
1923; op. cit. 17: 63. 1927. Figure 92.
294 FIELDIANA: BOTANY, VOLUME 24
Damp thickets and forested, rocky ravines, 400-2,000 m.;
Escuintla; Sacatepequez; Santa Rosa; Suchitepequez. Mexico
(Chiapas); El Salvador; Costa Rica.
Shrubs or small trees, 1.5-6 m. tall, much branched, the branches slender,
strigillose, striate; leaves on slender petioles 2.5-8.5 cm. long, the blades ovate, mostly
8-25 cm. long, 3-10 cm. wide, rather abruptly acuminate or long-acuminate, cuneate
at the base or often rounded and then abruptly cuneate and short-decurrent on the
petiole, commonly somewhat oblique at the base, the margins finely serrate- dentate,
very scabrous above, short-strigose and hispidulous beneath; heads often very
numerous, the pedicels to 2 cm. long; involucres 5-7 mm. high, strigillose; phyllaries
oblong-obovate, acute, 3-5-nerved; rays about equalling the involucre; disc corollas
about 5 mm. long; pales acuminate, strigillose, about 5 mm. long; ray achenes 5.5
mm. long, the body blackish, about 4 mm. long, winged, hispidulous toward the apex
on the outer side.
Known in El Salvador by the names "canilla," "tatascamillo,"
and "vara de zope."
ROJASIANTHE Standley & Steyermark
Tall, erect herbs, often very large, lignescent below and becoming treelike, the
stems terete, usually hollow, simple below, often much branched above; leaves
opposite, large, long-petiolate, the blades membranaceous, palmately 3-7-nerved,
generally lobate or angulate, the margins mucronate-dentate; heads heterogamous,
radiate, large and showy, few to several terminating the branches, the peduncles
slightly thickened at the apex; involucres broadly hemispheric; phyllaries 3-seriate,
the 6 outer ones firm, shorter and narrower than the inner ones (at least the 2
outermost ones ciliolate), then 2 orbicular to broadly ovate, eciliate ones, and finally
the inner, imbricate, 8-10 larger ones, these spreading in anthesis and with upcurved
margins; receptacle plano-convex, paleaceous; pales subtending and almost
completely enfolding both ray and disc flowers, accrescent in fruit, becoming
setulose-pectinate; ray flowers 12-15, uniseriate, neutral, the ligules spreading, white
or purplish white, entire, the pappus caducous, of several short, upwardly serrate
awns; disc flowers hermaphrodite, fertile, numerous (200 or more), the corollas
tubular-funnelform, the narrow tube puberulent, the throat and limb dark brown to
black, 5-cleft; style branches elongated, acute; anthers obtuse at the base, the apical
appendage ovate; achenes obovate, sparsely appressed-puberulent, the dorsal and
ventral surfaces carinate when mature; pappus caducous, composed of about 10
linear, subequal, upwardly serrate awns.
The genus consists of a single species.
Rojasianthe superba Standl. & Steyerm. Field Mus. Bot. 22:
315. /. 1, 2. 1940. Lamun, shil (Huehuetenango); poh, espina
(Quezaltenango); mac (San Marcos). Figure 93.
Usually in narrow, wet, wooded barrancos, in forest or on
exposed slopes, 2,200-3,400 m.; Huehuetenango; Quezaltenango; San
Marcos (type from Volcan de Tajumulco, between Las Canojas and
top of ridge, Steyermark 35835). Mexico (Chiapas).
NASH: FLORA OF GUATEMALA 295
Plants commonly 3-6 m. tall, lignescent below, the trunk sometimes as much as
10 cm. in diameter, the branches striate, hirtellous; leaves on petioles 2-10 cm. long,
the blades ovate or broadly triangular-ovate in outline, acuminate to long-acuminate,
truncate or cordate at the base and often with a small auricular appendage at the
junction with the petiole, mostly 10-22 cm. long, 8-22 cm. wide, 2-4-angulate on each
side, the lobes short, dentate and sometimes again lobate, minutely scaberulous on
the upper surface, scaberulous and viscid-glandular beneath, the margins mucronate-
crenate and denticulate; heads 3-6 at the end of a branch, the peduncles densely
hirtellous; phyllaries 3-seriate, obtuse, the outer ones 6, firm, relatively narrow, 8-11
mm. long, appressed-puberulent (at least the 2 outermost ones ciliolate), then a series
of 2 orbicular and 2 narrow, eciliate ones, and finally, 8-10 imbricate, larger ones of
more delicate texture, mostly 1.5-2 cm. long; disc about 1.5 cm. high, 2.5-3.5 cm.
broad; pales with spiny teeth 3-4 mm. long in fruit; ray flowers 12-15, the ligules
elliptic, acuminate, 2.5-4 cm. long, white or purplish white; ray pappus 1.5-2 mm.
long; disc flowers with whitish, densely puberulent tubes, the throat and limb dark
brown to purplish black; achenes about 6 mm. long, blackish, mottled with dull
brown; pappus 2-3 mm. long
Although the authors described the phyllaries as "distinctly 2-
seriate," my interpretation of three series is based on several
longitudinal and cross-section dissections, which also proved the
disc to be plano-convex rather than flat.
In some years only small plants develop, with very few heads,
but in other years, plants become arborescent; the smaller ones can
be pulled easily from the soil, revealing long, slender taproots.
RUMFORDIA DeCandolle
Reference: B. L. Robinson, A revision of the genus Rumfordia,
Proc. Amer. Acad. 44: 592-596. 1909.
Shrubs or perennial herbs, 1-3 m. tall, the stems often hollow; leaves opposite,
sessile or on winged petioles, sometimes connate and perfoliate at the base, the
margins serrate or dentate; heads heterogamous, radiate, usually disposed in
panicles; involucres hemispheric; phyllaries biseriate, the outer ones ovate to elliptic
or oblong-lanceolate, spreading, the inner ones much smaller, ovate, paleiform, erect,
cucullate, enfolding the achenes of the ray flowers; receptacle plano-convex,
paleaceous; ray flowers 6-20, fertile, the ligules elliptic or oblong, yellow or in age
beoming whitish; disc flowers 10 or more, hermaphrodite, fertile, the corollas tubular,
yellow, the narrow tube often equalling the ampliate limb in length, the limb 5-
dentate; anthers minutely sagittate at the base, the apical appendage ovate; achenes
obovoid, somewhat compressed, naked, glabrous; pappus wanting.
Twelve species, in Mexico and Central America, with three in
Guatemala. A fourth species from Chiapas, Mexico is also treated
here, as it may be expected in Guatemala.
Leaf blades elliptic-oblong, penninerved; outer phyllaries 4-6 mm. long.
R. penninervis.
296 FIELDIANA: BOTANY, VOLUME 24
Leaf blades ovate or triangular-ovate, triplinerved; outer phyllaries mostly 15-25 mm.
long.
Pedicels long-villous (at least above); outermost phyllaries more or less villous
dorsally with multiseptate hairs R. guatemalensis.
Pedicels hispid and conspicuously stipitate-glandular; outermost phyllaries minute-
ly viscid-puberulent.
Leaf blades mostly 10-17 cm. long; outer phyllaries 0.4-0.8 cm. wide, broadly
cuneate to subtruncate at the base. Mexico (Chiapas) R. media.
Leaf blades mostly 15-30 cm. long; outer phyllaries 1-2 cm. wide, broadly
rounded to cordate at the base. Guatemala (San Marcos) R. standleyi.
Rumfordia guatemalensis (Coulter) Blake, Journ. Wash.
Acad. Sci. 18: 25. 1928. Tetragonotheca guatemalensis Coulter, Bot.
Gaz. 16: 99. 1891 (type from Senahu, Alta Verapaz, J. D. Smith
1592). R. verapazensis Blake, Contr. U. S. Nat. Herb. 22: 609. 1924
(type from Finca Sepacuite, Alta Verapaz, Cook & Griggs 239).
Polymnia verapazensis Standl. & Steyerm. Field Mus. Bot. 23: 144.
1944 (type from Panzal, Baja Verapaz, Tuerckheim II. 1723).
Wet thickets or forest, often along streams, 600-1,500 m.; Alta
Verapaz; Baja Verapaz.
Erect branching herbs, 1-1.5 m. high, the branches densely villous or pilosulous
above with weak, multiseptate hairs, glabrous below; principal leaves petiolate, the
petioles broadly winged, 1-9 cm. long, not dilated at the base, the blades broadly
triangular-ovate or rounded-ovate, mostly 12-18 cm. long, 10-15 cm. wide, acuminate,
abruptly contracted and broadly rounded or cuneate-rounded at the base and long-
decurrent on the petiole, sometimes subhastate-lobate at the base, triplinerved above
the base, the margins closely denticulate, sparsely villous or glabrate above, thinly
tomentose or appressed-tomentose beneath or sometimes glabrate, gland-dotted, the
upper leaves much smaller, short- petiolate or subsessile, the blades ovate or ovate-
lanceolate, long-acuminate, not lobate, gradually narrowed to the base; heads on
long, slender pedicels, disposed in corymbiform panicles, the pedicels densely long-
villous, at least above; outer phyllaries 5, greenish, broadly ovate or rhombic-ovate or
sometimes narrower, mostly 12-20 mm. long, 6-10 mm. wide, acuminate or abruptly
long-acuminate, conspicuously 3-nerved, long-villous dorsally with multiseptate hairs,
the inner phyllaries oblong-ovate, acuminate, about 7.5 mm. long, stipitate-glandular
and /or hispid-pilose; ray flowers about 15, the ligules yellow, narrowly oblong or
linear-oblong, 10-15 mm. long; disc corollas about 7.5 mm. long, the tube pubescent to
stipitate-glandular, about as long as the slender, funnelform throat; pales obovate,
abruptly acuminate, about 5.5 mm. long, sometimes stipitate-glandular and
hispidulous, ciliate; achenes obovoid, about 2 mm. long, blackish, glabrous.
Rumfordia media Blake, Journ. Wash. Acad. Sci. 28: 490.
1938.
Not reported from Guatemala but to be expected there. Mexico
(Chiapas).
Erect, coarse herbs, the stems angulate. hollow, glabrate or sparsely short-
villosulous to puberulent; leaves opposite, on broadly winged petioles 3-9 cm. long.
NASH: FLORA OF GUATEMALA 297
these somewhat dilated and connate at the base, the leaf blades triangular-ovate,
mostly 10-17 cm. long, 6-14 cm. wide, acute or acuminate, abruptly narrowed into the
petiole, often subhastate at the base, triplinerved, the margins irregularly denticulate-
serrate, thinly villosulous above, scarcely rough to the touch, densely or thinly viscid-
tomentose beneath, leaves at base of panicles much reduced; heads few or numerous,
2-3 cm. wide when mature, pedicellate, disposed in terminal panicles, the branches
and pedicels usually densely covered with stipitate glands, the pedicels short or
elongated; outer phyllaries 5, spreading or reflexed at maturity, ovate, lance-ovate, or
elliptic-ovate, unequal in length, mostly 1.5-1.7 cm. long, 0.4-0.8 cm. wide, acute,
broadly cuneate to subtruncate at base, 3-5-nerved, densely but minutely glandular-
puberulent outside, the inner phyllaries (subtending the ray flowers) ovate to lance-
ovate, acuminate, usually densely stipitate-glandular; receptacle about 1 cm. broad;
ray flowers 10-13, fertile, the ligules yellow, 10-12 mm. long, trilobate at the apex;
disc corollas numerous, yellow, about 6.5 mm. long; achenes brownish black,
glabrous, cuneate-obovoid, about 2 mm. long.
Rumfordia penninervis Blake, Brittonia 2: 343. 1937. Te.
Damp, brushy hillsides, 1,800-3,500 m.; Quezaltenango (type
collected along road between Quezaltenango and Colomba, Skutch
1973); San Marcos (Volcan de Tajumulco).
Erect shrubs, 2-3 m. high, the branches glabrous; leaves sessile or nearly so,
elliptic-oblong, mostly 17-26 cm. long and 5-10 cm. wide, acuminate, gradually
attenuate to the base, the margins serrate, glabrous above, pilosulous on the costae
and veins beneath or almost glabrous, penninerved; panicles usually large, the
pedicellate heads numerous, the pedicels 1-2.5 cm. long, pilosulous and glandular-
pilose; disc conic, 6-8 mm. high; outer phyllaries 5, obovate, 4-6 mm. long, obtuse or
rounded at the apex, sometimes ciliolate, glabrous, the inner phyllaries 10, lanceolate,
acuminate; ray flowers 10, the ligules yellow, about 12 mm. long; disc corollas pilose
and stipitate-glandular on the tube, almost 5 mm. long; achenes obovoid, about 2
mm. long or slightly larger, blackish, striate.
Rumfordia standleyi (Steyerm.) Standl. & Steyerm. Field
Mus. Bot. 23: 265. 1947. Polymnia standleyi Steyerm. Field Mus.
Bot. 23: 106. 1944. Mac. Figure 94.
Damp or wet, mixed forest or thickets, 2,500-3,000 m.; San
Marcos (type collected between La Vega ridge and northeast slopes
of Volcan de Tacana, vicinity of San Rafael, Steyermark 36181 ; also
collected on Volcan Tajumulco).
Erect, coarse herbs, 2-5 m. high, the stems branching, angulate, hollow,
sometimes as much as 5 cm. in diameter, glabrate or sparsely short-villosulous or
puberulent; leaves opposite, on broadly winged petioles 3-15 cm. long, these dilated
and auriculate-clasping and connate at the base, the leaf blades triangular-ovate,
mostly 15-30 cm. long, 6-16 cm. wide, acute, abruptly narrowed into the petiole, often
subhastate at the base, triplinerved, the margins irregularly denticulate-serrate,
thinly villosulous above, scarcely rough to the touch, densely or thinly viscid-
tomentose beneath, leaves at base of panicles reduced; heads numerous, 2-3 cm. wide,
pedicellate, disposed in large fairly dense panicles, the branches and pedicels usually
298 FIELDIANA: BOTANY, VOLUME 24
densely covered with stipitate glands, the pedicels short or elongated; outer phyllaries
5, spreading or reflexed when mature, ovate or rounded-ovate, 1.5-2.5 cm. long, 1-2
cm. wide, acute, rounded to cordate at the base, 5-6-nerved, densely glandular-
puberulent outside; receptacle about 1 cm. broad; ray flowers 9-11, fertile, the ligules
yellow, 1.5-2.5 cm. long, trilobate at the apex; disc corollas numerous, yellow, 7-8 mm.
long; achenes dark brown, striate, cuneate-obovoid, about 2.5 mm. long.
SABAZIA Cassini
Reference: B. L. Robinson & J. M. Greenman, Revision of the
genus Sabazia, Proc. Amer. Acad. 40: 3-6. 1904; E. K. Longpre, The
systematics of the genera Sabazia, Selloa and Tricarpha
(Compositae), Mus. Mich. St. Univ. Publ. Biol. 4: 287-383. 1970.
Small annual or perennial herbs, the stems erect, decumbent, or procumbent,
glabrous to densely pilose or hirsute; leaves opposite, cauline, petiolate or sessile, the
blades ovate or lanceolate to elliptic or linear, mostly 3-nerved or triplinerved, the
margins more or less serrate or entire; inflorescences cymose with few heads or
usually the heads solitary on conspicuous peduncles; heads radiate; involucres
campanulate or hemispherical; phyllaries 2-3-seriate (in ours), imbricate, the outer
ones usually herbaceous on the margins, often carinate, subequal, the apices often
tinged with purple, the inner ones membranaceous, lacerate-ciliate above the middle;
pales persistent, scarious, elliptic-lanceolate to lanceolate, often bifid or irregularly
trilobate; receptacles conical or convex; ray flowers uniseriate, pistillate, fertile, the
tube usually pubescent, the ligule white on the upper surface, usually purplish or
pinkish beneath, the apex tridentate (rarely bifid); disc flowers hermaphrodite, fertile,
the corollas yellow, tubular, the tube usually pubescent, the throat narrowly
campanulate, the limb 5-cleft; style branches acute or obtuse; anthers minutely
sagittate at the base, apically appendaged; achenes obovate or turbinate, terete or
obscurely angulate, black, glabrous or pubescent, with or without pappus; pappus,
when present, composed of few-several fimbriate, awnlike squamellae.
Eleven species, generally confined to montane areas, ranging
from Mexico to Colombia, the majority in Mexico, with only two in
Guatemala.
Leaf blades mostly ovate to lance-ovate, acuminate, 2-6 cm. long, 0.8-6 cm. wide.
S. sarmentosa.
Leaf blades mostly rhombic-lanceolate, linear-lanceolate, or linear-elliptic, acute or
obtuse, 0.8-2.5 cm. long, usually less than 0.8 cm. wide S. pinetorum.
Sabazia pinetorum Blake, Brittonia 2: 347. 1937. S. pinetorum
var. dispar Blake, torn. cit. 348 (type from Huehuetenango, Sierra
de los Cuchumatanes, Skutch 1234 in part). Figure 95.
In open pine or juniper forest, often on limestone bluffs or
outcrop, 3,140-3,500 m., Huehuetenango (type from Sierra de los
Cuchumatanes, Skutch 1234 in part).
Decumbent perennials, the stems slender, to 63 cm. long, simple or sparsely
branched, sparsely pilose; leaves on hirsute petioles 2-6 mm. long, the blades mostly
NASH: FLORA OF GUATEMALA 299
0.8-2.5 cm. long, 0.2-1.2 cm. wide, the lower ones commonly rhombic-lanceolate to
lanceolate, the upper ones linear-lanceolate to linear-elliptic, acute or obtuse, cuneate
or obtuse at the base, the margins entire to shallowly crenate-mucronulate, sparsely
pilose to densely strigose on both surfaces, 3-nerved; peduncles 5-15 cm. long, pilose
and sometimes glandular, the hairs usually appressed below the head; heads solitary;
involucres 4-5 mm. high; phyllaries 2-3-seriate, the outer ones herbaceous, ovate or
elliptic-ovate, obtuse, 3.5-5 mm. long, entire, glabrous, lacerate or ciliate, striate,
often purplish on the margins, the inner ones a little longer, acute, the margins
scarious; receptacle conical; pales persistent, acuminate, sometimes with 1 or 2 lateral
lobes; ray flowers about 8, the ligules white above, pinkish or purplish below, 5-10
mm. long, tridentate; disc flowers about 60, yellow, about 3 mm. long; achenes black,
obconical, obscurely ridged, those of the ray flowers glabrous; ray pappus wanting;
achenes of the disc flowers glabrous or hirtellous; disc pappus wanting or sometimes
of 14-20 unequal, fimbriate, awnlike squamellae 1-2.2 mm. long.
Sabazia sarmentosa Less. Linnaea 5: 148. 1830. Allocarpus
sabazioides Schlecht. op. cit. 9: 590. 1834. Calea sabazioides Hemsl.
Biol. Centr. Am. Bot. 2: 206. 1881. Tridax ehrenbergii Sch. Bip. ex
Klatt, Leopoldina 23: 145. 1887. S. radicans Blake, Contr. U. S.
Nat. Herb. 22: 615. 1924.
Damp forest or thickets, sometimes in oak forest, on open,
brushy slopes, along stream beds, rarely on dry, rocky, open places,
sometimes a weed in cornfields, 2,900-3,800 m.; Chimaltenango;
Huehuetenango; El Progreso; Quezaltenango; El Quiche; Sacatepe-
quez; San Marcos; Solola. Southern Mexico.
Procumbent, decumbent, or ascending perennials (rarely erect), the stems to
about 90 cm. long, simple or branching, more or less strigose-pilose above, glabrous
below, sometimes rooting at the nodes; leaves short-petiolate, the blades ovate to
lance-ovate, mostly 2-6 cm. long, 0.8-6 cm. wide, acuminate, truncate or cuneate at
the base and then often decurrent on the petioles, the margins serrate-dentate,
sparsely to densely strigose; peduncles 5-20 cm. long, more or less appressed-
pubescent and sometimes glandular; heads 1-8, terminal or axillary, the inflorescence
sometimes subcymose; involucres 4-6 cm. high; phyllaries 2-3-seriate, the outer ones
herbaceous, elliptic-ovate to lance-ovate, obtuse, 3-6 mm. long, the inner ones slightly
longer, acute to acuminate, scarious or hyaline; ray flowers 5-11, the ligules white
above, white or pinkish beneath, 5-10 mm. long, tridentate or trilobate; disc flowers
30-65 (-100), yellow, the tube broad; achenes 1-2.5 mm. long, obscurely 3-4-ridged,
glabrous or strigose, especially along the margins, with or without pappus; pappus,
when present, consisting of 12-16 awnlike, fimbriate squamellae about 2 mm. long.
SALMEA DC.
Reference: S. F. Blake, A revision of Salmea and some allied
genera, Journ. Bot. 53: 196-201. 1915.
Shrubs, erect or scandent, the stems terete, striate; leaves all or mostly opposite,
petiolate or subsessile, the blades usually more or less coriaceous, triplinerved or
penninerved, glabrous or pubescent, the margins entire or dentate; inflorescences
300 FIELDIANA: BOTANY, VOLUME 24
cymose, often becoming paniculate; heads discoid, homogamous, pedicellate or sessile,
on axillary and terminal peduncles; involucres narrowly or broadly campanulate;
phyllaries 2-6-seriate, appressed, ovate or ovate-lanceolate to oval, chartaceous or
membranous-coriaceous, sometimes with herbaceous tips, often striate, often ciliate;
receptacle more or less conic, paleaceous; pales oblong, subacute or obtuse, persistent,
enfolding the achenes; flowers numerous, the corollas erect, white, the tubes short,
the throat broadly cylindrical, the limb 5-dentate; anthers minutely sagittate at the
base, the apical appendage triangular-ovate, obtuse; style branches papillose-
hirtellous toward the apex; achenes oblong-cuneate, strongly compressed, dark brown
to purplish black, with or without a paler border, the margins usually ciliate; pappus
of 2 (rarely 3) subequal, upwardly ciliate awns and sometimes a few, irregular, much
shorter squamellae also present on the crown.
About 10 species, all in tropical America, with only four known
from Central America.
Involucres narrow, the heads turbinate, mostly 7-9 mm. high, 3-4 mm. wide.
S. orthocephala.
Involucres broad, the heads broadly ovoid to subglobose, mostly 4.5-7 mm. high, 4-7
mm. wide.
Phyllaries all rounded at the apex; achenes more or less appressed-pubescent.
S. pubescens.
Phyllaries all acute to acuminate, or at least the outer ones acute; achenes
essentially glabrous except for the ciliate margins.
Stems and pedicels glabrous or nearly so; outer phyllaries acute, the others
subacute to obtuse; achenes with 2 awns S. scandens.
Stems and pedicels covered with dense, feltlike tomentum; all phyllaries acute to
acuminate; achenes with 2-3 principal awns and 1-6 shorter, but
conspicuous, irregular squamellae S. tomentosa.
Salmea orthocephala Standl. & Steyerm. Field Mus. Bot. 23:
145. 1944.
Damp or dry thickets, along streams or on rocky slopes, 300-450
m.; Baja Verapaz; Chiquimula; Guatemala (type from thicket near
Concua Bridge, over Rio Motagua, Standley 59321). Honduras.
Shrubs, 1.5-2.5 m. high, suberect or sarmentose or somewhat scandent, the
branches striate, glabrous or nearly so; leaves on petioles 0.3-1 cm. long, the blades
chartaceous, ovate to lance-ovate or triangular-ovate, mostly 8-13 cm. long, 3-6 cm.
wide, narrowly long-acuminate, rounded at the base and often abruptly contracted
and short-decurrent on the petiole, triplinerved, glabrous on both surfaces or
sometimes more or less pilose beneath on costae and veins, the margins remotely
serrate-dentate or subentire; inflorescences densely cymose-paniculate, the panicles
corymbiform, as much as 10 cm. broad; heads numerous, 7-9 mm. high, appearing
turbinate, sessile or on glabrous pedicels to 8 mm. long; involucres narrowly
campanulate, 4-6 mm. high; phyllaries 3-4 -seriate, ovate-orbicular or obovate-oval,
broadly rounded at the apex, ciliate, the margins pale, dorsally glabrous or nearly so;
pales hyaline, narrow, obtuse, puberulent near the apex; corollas white, glabrous,
about 3 mm. long; achenes blackish, 2-3 mm. long, sparsely appressed-pilose, the awns
1-1.5 mm. long, deciduous.
NASH: FLORA OF GUATEMALA 301
Salmea pubescens (Blake) Standl. & Steyerm. Field Mus. Bot.
23: 146. 1944. S. scandens var. pubescens Blake, Brittonia 2: 351.
1937 (type from Huehuetenango, Skutch 1644).
Damp or dry, wooded ravines and rocky hillsides, often in pine-
oak or oak forest, 1,100-2,400 m.; Alta Verapaz; Guatemala;
Huehuetenango; Jalapa; Zacapa. Mexico (Chiapas).
Sarmentose or scandent shrubs, the branches densely brownish pilose; leaves on
short, thick petioles or the uppermost ones subsessile, the blades subcoriaceous, often
lustrous, ovate to rounded-ovate, mostly 3-7 cm. long, 1-4 cm. wide, acute or
acuminate, broadly rounded at the base and sometimes contracted and short-
decurrent on the petiole, triplinerved, the upper surfaces puberulent along the veins,
the lower surfaces densely and softly pilose or tomentose, the margins remotely
denticulate or subentire; inflorescences corymbiform; heads 5-7 mm. high, on
pubescent pedicels; involucres campanulate; phyllaries usually densely pubescent, the
outer ones shortest, all rounded at the apex and more or less ciliolate; pales obtuse,
puberulent; corollas white; achenes about 2 mm. long, dark brown to blackish,
appressed-pubescent, ciliate on the margins; principal pappus awns 2, commonly
about 2 mm. long (rarely only 1-1.5 mm. long), sometimes one considerably longer
than the other, often 2 or 3 very small squamellae also present.
Salmea scandens (L.) DC. Cat. Hort. Monsp. 141. 1813.
Bidens scandens L. Sp. PI. 833. 1753. S. grandiceps Cass. Diet. Sci.
Nat. 47: 88. 1827. S. scandens var. obtusata Blake, Journ. Bot. 53:
197. 1915 (type from Alta Verapaz, Tuerckheim 392, in part).
Bajche, caycam, kekchicay (Alta Verapaz); vara de fuego (Quezal-
tenango); iclab (British Honduras, Maya). Figure 96.
Damp or wet thickets or forest, often in pine-oak forest,
frequently in second growth, sea level to 3,000 m. (most common
1,500-2,400 m.); Alta Verapaz; Chimaltenango; Chiquimula; Es-
cuintla; Guatemala; Huehuetenango; Izabal; Peten; Quezal-
tenango; El Quiche; Retalhuleu; Sacatepequez; San Marcos; Santa
Rosa; Solola; Suchitepequez. Southern Mexico; British Honduras to
El Salvador and Panama; West Indies; tropical South America.
Erect, arching or (more often) more or less scandent shrubs, the branches
glabrate; leaves short-petiolate, the blades usually more or less coriaceous, often
lustrous, broadly ovate to oblong-ovate or lanceolate, mostly 4-12 cm. long, acute or
acuminate, subcordate at the base or rounded and rather abruptly cuneate or short -
decurrent on the petiole, glabrous or nearly so, sometimes pilose beneath along the
costae and veins, the margins usually remotely repand-dentate to entire, rarely
serrate-dentate; inflorescences cymose, the small panicles sometimes corymbiform;
heads usually numerous, 4.5-7 mm. high, on glabrous pedicels; involucres camp-
anulate, 2.5-4 mm. high; phyllaries about 3-seriate, broadly ovate to ovate-lanceolate,
the outermost ones acute, the others acute, subacute, or obtuse, glabrous or
pubescent, ciliolate; pales narrow, obtuse, almost glabrous or somewhat puberulent,
often ciliolate; corollas white; achenes brown or blackish, about 2 mm. long, usually
glabrous but ciliate on the margins; pappus awns commonly about 1 mm. long.
302 FIELDIANA: BOTANY, VOLUME 24
In Guatemala (especially near Coban, Alta Verapaz) these
plants are much used in decoction for domestic medicine. In El
Salvador (where they are called "salta-afuera") and in British
Honduras, they are used as a fish poison, said to be so powerful that
the fish "jump out of the water."
Salmea tomentosa D. Nash, Phytologia 31: 361. 1975.
Damp forest, about 1,650 m.; Baja Verapaz (type from
mountain-side north of divide north of Santa Rosa, Standley 69898;
Zacapa (Sierra de Las Minas).
Scandent shrubs, the branches densely brown-tomentose; leaves on short, thick
petioles or the uppermost ones subsessile, the blades coriaceous, often lustrous above,
ovate to rounded-ovate, mostly 2-9 cm. long, 1.5-6 cm. wide, acuminate, broadly
rounded to subcordate at the base, triplinerved, the upper surfaces glabrous or
sometimes puberulent along costae and veins, the lower surfaces more or less pilose or
tomentose, the margins remotely denticulate or subentire; inflorescences cymose,
corymbiform, the peduncles and pedicels densely covered with brownish, feltlike
tomentum; heads 6-7 mm. high, 5-6 mm. broad, 1-3 on a pedicel; involucres
campanulate; phyllaries about 3-seriate, ovate to lanceolate, acute to acuminate,
densely pubescent, ciliolate; pales acute or obtuse, at least the outer ones usually
conspicuously carinate, pubescent to puberulent, at least near the apex, ciliolate;
corollas white, glabrous; achenes 2.5-3 mm. long, blackish, glabrous, minutely striate
when mature, the margins ciliate; pappus of 2 or sometimes 3 principal awns often 2
mm. long, and 1-6 shorter, but conspicuous, irregular squamellae.
SANVITALIA Lamarck
Reference: Andrew M. Torres, Revision of Sanvitalia
(Compositae - Heliantheae), Brittonia 16: 417-433. 1964.
Annual or perennial herbs or rarely low shrubs, branched, usually prostrate or
decumbent, the stems pubescent; leaves opposite, petiolate or sessile, the margins
entire or sometimes lobate; heads heterogamous, radiate, usually pedunculate,
involucres hemispherical or broadly campanulate; phyllaries 1-3-seriate, subequal, dry
or herbaceous at the apex or the outermost foliaceous; receptacle conical; pales
conduplicate, hyaline, subtending or enfolding the disc flowers; ray flowers 1-2-
seriate, persistent, pistillate, fertile, lacking a distinct tube, the ligules yellow,
spreading; disc flowers hermaphrodite, fertile, usually purplish, regular, tubular, the
limb little dilated, 5-dentate; anthers sagittate at the base, apically appendaged; style
branches truncate or with a short appendage; ray achenes thick, triquetrous,
glabrous, the angles produced into 3 awns; disc achenes commonly dimorphic,
narrow, the outer ones usually triquetrous, sometimes 4 -angled, the inner ones
compressed and often winged, the wings usually produced into short awns.
Seven species, in Mexico and southwestern United States with
one extending into Guatemala.
Sanvitalia procumbens Lam. Journ. Hist. Nat. Paris 2: 176.
NASH: FLORA OF GUATEMALA 303
t.33. 1792. S. villosa Cav. Ic. Descr. PL 4: 31. 1797. S. acinifolia DC.
Prodr. 5: 628. 1836. Figure 97.
Dry, rocky slopes, or a weed in cornfields, 1,100-1,350 m.;
Huehuetenango; El Progreso; El Quiche. Mexico.
Procumbent or prostrate annuals, much branched from the base, the stems
mostly 10-40 cm. long, sometimes forming dense mats, hirsute or hispidulous, very
leafy; leaves short-petiolate, the blades thick, broadly lanceolate to ovate or oblong,
1-3 (-5) cm. long, rounded or very obtuse at each end, strigose, the margins entire;
heads numerous, solitary, leafy-bracteate, on short or more or less elongated, hirsute
peduncles; involucres hemispherical, usually less than 1 cm. broad; phyllaries 4-5
mm. long, pale green, imbricate, broadly ovate to oval or almost rounded, very obtuse
or rounded at the apex, sometimes mucronulate, villous, densely ciliate with long,
white hairs; pales rather rigid, cuspidate; ray flowers about 12, sessile on the achenes,
the ligules yellow or orange, minutely bifid at the apex, glabrous, 4-9 mm. long; disc
flowers numerous, the corollas about 3 mm. long, yellow or purple; ray achenes with
3 awns; outer disc achenes winged or wingless with 1-2 short awns, the inner achenes
densely and coarsely tuberculate, obscurely or conspicuously 3-4 -angled, ciliate,
awnless or with 1-2 short awns.
Variable, weedy plants, abundant in many parts of Mexico but
rare in Guatemala.
SCHISTOCARPHA Lessing
References: P. A. Rydberg, Schistocarpha, N. Amer. Flora 34:
303-306. 1927; H. Robinson and R. D. Brettell, Tribal revisions in
the Asteraceae. IV, The Relationships of Neurolaena, Schisto-
carpha and Alepidocline, Phytologia 25: 439-445. 1973.
Erect, coarse, branched herbs; leaves opposite, petiolate, the petioles often
dilated at the base and sometimes clasping, the blades usually more or less decurrent
on the petiole, the margins dentate or serrate; inflorescences cymose-paniculate;
heads (in ours) radiate, usually numerous; involucres campanulate; phyllaries 3-4-
seriate, imbricate, graduate, membranous, striate; receptacle convex, paleaceous;
pales membranous, lacerate, mostly 3-cleft; ray flowers usually uniseriate, rarely in
several series, pistillate, fertile, the ligules none or often very small and
inconspicuous, white, 3-4-dentate; disc flowers numerous, hermaphrodite, fertile, their
corolla tubes slender, abruptly dilated into the limb, this 5-dentate; anthers blackish,
sagittate at the base, apically appendaged; style branches slender, papillose or
minutely hirtellous; achenes more or less oblong, attenuate to the base, glabrous,
usually minutely striate, callous at each end, black, lustrous; pappus uniseriate, the
bristles usually about 20.
About a dozen species, in tropical America, with six in
Guatemala. The plants look much like some species of Eupatorium,
and most of the species are monotonously alike in general
appearance. This perhaps accounts for so many incorrect determi-
nations; for instance, numerous Guatemalan collections that were
304 FIELDIANA: BOTANY, VOLUME 24
originally distributed as S. paniculata Klatt have now been
determined to be either S. seleri Rydb., S. platyphylla Greenm., or
S. oppositifolia (Kuntze) Rydb. S. paniculata, apparently limited to
Costa Rica, has glandular inflorescences, large heads with in-
volucres about 7 mm. high and 1 cm. wide, 20-30 ray flowers with
ligules 5-8 mm. long, and 50-70 disk flowers.
Ligules of pistillate flowers minute (0.3-2 mm. long) or none.
Pistillate flowers numerous in several series, their ligules mostly less than 0.5 mm.
long; styles at anthesis scarcely or only slightly exceeding the corolla tubes.
S. oppositifolia.
Pistillate flowers few in a single series, commonly without ligules (if present, 1-2
mm. long); styles conspicuously exceeding the corolla tubes, the exserted
portion often almost as long as the tube.
Stems usually glabrate or thinly pilose; phyllaries obtuse or subacute.
S. platyphylla
Stems usually villous-pilose; phyllaries acute to acuminate S. kellermanii.
Ligules of pistillate flowers conspicuous, 3-9 mm. long.
Pedicels essentially glabrous except for scattered, long-stipitate, gland-tipped hairs;
ray flowers with glabrous corolla tubes; achenes 2.5-3 mm. long.
S. steyermarkiana.
Pedicels hirtellous, with or without gland-tipped hairs; ray flowers with pubescent
corolla tubes; achenes 1-1.5 mm. long.
Involucres 5-6 mm. high; ray flowers with ligules 5-8 mm. long S. longiligula.
Involucres 4-5 mm. high; ray flowers with ligules commonly 3-4 mm. long (rarely
to 5 mm) S. seleri.
Schistocarpha kellermanii Rydb. N. Amer. Fl. 34: 306. 1927.
Chichavac (Chimaltenango); sajan (San Marcos).
Damp or wet thickets or mixed forest or sometimes in oak
forest, 1,500-3,200 m.; Chimaltenango; Escuintla; Quezaltenango;
Sacatepequez (type from Volcan de Agua, KeHerman 7293); San
Marcos; Solola. Mexico (Chiapas).
Erect, coarse, suffrutescent herbs, 1-3.5 m. high, the branches stout, striate,
villous- hirsute; leaves on petioles 5-11 cm. long, the leaf blades broadly ovate or
triangular-ovate to cordate, mostly 8-18 cm. long, acuminate, abruptly contracted at
the base and cuneate-decurrent on the petiole for one-half to two-thirds its length,
the margins closely dentate-serrate, thinly or densely villous-hirsute on both surfaces;
inflorescences dense, rounded panicles; heads numerous, on slender, hirtellous
pedicels; involucres 4-6 mm. high; phyllaries lanceolate or linear-lanceolate, acute to
acuminate, hirsute or glabrate, or the inner ones merely ciliate, somewhat fimbriate
at the apex; pales 5-7 mm. long, irregularly 3-cleft; pistillate flowers few, uniseriate,
usually without ligules (if ligules present, these minute), the tube much exceeded by
the style; disc corollas about 6 mm. long; achenes glabrous, about 1.5 mm. long;
pappus bristles about 5 mm. long.
Schistocarpha longiligula Rydb. N. Amer. Fl. 34: 305. 1927.
NASH: FLORA OF GUATEMALA 305
Wet, mixed forest, 1,800-2,600 m.; El Progreso; El Quiche (type
from San Miguel Uspantan, Heyde & Lux 3383).
Erect perennials, 2-3 m. high, the branches villous- pilose; leaves or petioles 3-8
cm. long; leaf blades broadly ovate to triangular-ovate, mostly 9-18 cm. long,
acuminate, abruptly contracted at the base and then partially decurrent on the
petiole, the margins finely dentate, more or less villous-pilose on both surfaces, more
densely so beneath; inflorescences paniculate; heads numerous, radiate, on slender
pedicels; involucres 5-6 mm. high; phyllaries 3-4-seriate, linear, obtuse to subacute,
glabrate; ray flowers 10-12, the ligules 5-8 mm. long; disc corollas yellow, about 4
mm. long; achenes glabrous, 1-1.5 mm. long; pappus bristles white, about 3 mm. long.
In the original description, the achenes are said to be 4 mm.
long, but in my examination of the holotype and two isotypes, I
found none longer than 1.5 mm.
In herbaria, a number of specimens of S. seleri Rydb. have been
misidentified as S. longiligula.
Schistocarpha oppositifolia (Kuntze) Rydb. N. Amer. Fl. 34:
306. 1927. Zycona oppositifolia Kuntze, Rev. Gen. 1: 373. 1891. S.
hoffmanii Kuntze, op. cit. 3: 170. 1898.
Wet thickets or open fields, often on sandbars along streams,
sometimes in pine groves, near sea level to 1,200 m.; Alta Verapaz;
Chiquimula; Izabal; Peten; Retalhuleu; Zacapa. Southern Mexico;
British Honduras; Honduras; Nicaragua; Costa Rica; Panama;
western South America.
Erect, coarse annual or perennial herbs, 0.5-2 m. high, the branches villous-
hirsute to glabrate; leaves on petioles 2-15 cm. long, more or less broadly winged near
the blade but only very narrowly winged near the base, the blades broadly ovate,
mostly 10-20 cm. long, acuminate or long-acuminate, abruptly contracted at the base,
then cuneate and decurrent on the petiole, the margins serrate-dentate, often
coarsely and remotely so, thinly hirsutulous on both surfaces; inflorescences small or
large, more or less rounded panicles, often leafy, the heads short-pedicellate;
involucres 6-8 mm. high, 7-10 mm. broad; phyllaries about 3-seriate, lance-linear to
oblong, obtuse or subacute, striate, ciliate; pales 6-7 mm. long, deeply 3-cleft; ray
flowers numerous in several series, their styles scarcely exceeding the corolla tubes,
the ligules minute (mostly less than 0.5 mm. long) or (rarely) none; disc flowers
numerous, the corollas yellow, about 5 mm. long; achenes about 1.5 mm. long,
glabrous; pappus bristles about 5 mm. long.
Schistocarpha platyphylla Greenm. Field Mus. Bot. 2: 274.
1907. Quesillo (Quezaltenango). Figure 98.
Damp or wet thickets or mixed forest, (650) 1,000-3,400 m.
(more common above 2,000 m.); Chimaltenango; Escuintla; Quezal-
tenango (type from Volcan de Santa Maria, Kellerman 5295);
306 FIELDIANA: BOTANY, VOLUME 24
Retalhuleu; Sacatepequez; San Marcos; Suchitepequez. Mexico
(Chiapas); El Salvador.
Erect, branched herbs, 1-3 m. high, the stems and branches thinly pilose or
glabrate; petioles mostly 2-10 cm. long, more or less winged, at least above and
sometimes almost to the base, the blades thin, ovate, rounded-ovate, triangular-
ovate, or almost cordate, mostly 10-25 cm. long, acuminate, rounded or acute or
subcordate at the base and usually abruptly contracted and then cuneate-decurrent
on the petiole, the margins closely dentate or serrate-dentate, sparsely pilose on both
surfaces; inflorescences often very large, corymbiform-paniculate; heads on slender
pedicels; involucres 4-6 mm. high; phyllaries about 3-seriate, lance-oblong, obtuse or
subacute, striate, glabrous or nearly so; pales about 3 mm. long, irregularly 3-5-cleft;
pistillate flowers about 13, commonly without ligules but when these present, only 1-
2 mm. long, their styles conspicuously exceeding the corolla tubes; disc flowers
yellow, about 5 mm. long, the tube puberulent; achenes about 1 mm. long, essentially
glabrous; pappus about 4 mm. long.
Schistocarpha seleri Rydb. N. Amer. Fl. 34: 305. 1927.
Damp or wet thickets or dense forest, 1,300-3,200 m.; Alta
Verapaz; Baja Verapaz; Chimaltenango; Quezaltenango; El Quiche;
San Marcos; Solola; Suchitepequez; Totonicapan. Mexico
(Chiapas); Honduras; Nicaragua; Costa Rica.
Erect, branched herbs, 1-2 m. tall, the branches usually softly and densely
hirsute or pilose; leaves on petioles 2-10 cm. long, the leaf blades thin, broadly ovate
to triangular-ovate, mostly 5-20 cm. long, acuminate, abruptly contracted at the base
and then cuneate-decurrent on the petiole for one-third to one-half its length, the
margins serrate, sparsely or densely short-hirsute on both surfaces; inflorescences
corymbose-paniculate; heads numerous, radiate, on hirtellous pedicels; involucres 4
(-5) mm. high; phyllaries 3-seriate, elliptic-oblong, rounded or obtuse, more or less
ciliate and somewhat fimbriate at the apex; pales lacerate; ray flowers 10-15, the
ligules white, 3-4 (-5) mm. long; disc corollas yellow, about 5 mm. long; achenes
black, 1-1.5 mm. long, glabrous; pappus bristles 3-4 mm. long.
Schistocarpha steyermarkiana H. Robinson, Phytologia 29:
248. 1974.
Known only from the type collection, Volcan Agua, alt. 10,000
ft., Sacatepequez, Kellerman 7223.
Erect, suffrutescent plants, 1-2 m. tall, sparsely branched, the stems sparsely
pilose to glabrate; leaves on petioles 1.5-4 cm. long, the blades broadly ovate, mostly
5-12 cm. long, acuminate, the base subtruncate or almost rounded and then abruptly
and often obliquely cuneate, the margins rather coarsely serrate, sparingly pilose on
both surfaces, paler beneath; inflorescence a lax panicle; heads radiate, on pedicels 4-
13 mm. long, these sparsely glandular with long-stipitate, gland-tipped hairs;
involucres 5-7 mm. high; phyllaries about 4-seriate, oblong-lanceolate, glabrous, the
apices obtuse to subacute and densely fimbriate; pales narrowly linear, acute,
fimbriate; ray flowers 8-10, the tube short, about 1.5 mm. long, indistinct, glabrous,
the ligules 7-9 mm. long; disc flowers 25-30, the corollas yellow, about 5.5 mm. long;
achenes 2.5-3 mm. long, glabrous; pappus soon deciduous, the bristles 3-4 mm. long.
NASH: FLORA OF GUATEMALA 307
SCLEROCARPUS Jacquin
Annual or perennial, branching herbs, strigose or villous; leaves petiolate, the
upper ones alternate, the lower ones opposite, the margins coarsely dentate; heads
heterogamous, radiate, pedunculate at the ends of the branches or in the forks of
branches, sometimes opposite the leaves; involucres campanulate; phyllaries few,
uniseriate or biseriate, herbaceous throughout or nearly so; receptacle convex or
conic; pales first enclosing the base of the disc flowers, in age finally closing above
the achene, becoming indurate, rugose or tuberculate, often contracted into an apical
beak, deciduous with the achene; ray flowers neutral, uniseriate, the ligules yellow or
orange-yellow, ovate to orbicular; disc flowers hermaphrodite, fertile, the corollas
regular, the limb and tube not or scarcely differentiated, the lobes bearded within;
anthers yellow, entire at the base or minutely sagittate; style branches of the
hermaphrodite flowers elongated and terete; achenes obovoid, thick, laterally
compressed, narrowed at the base, naked or with a short annulus at the apex.
Eight species, all American save one in tropical Africa, with
three in Guatemala.
Heads leafy-bracteate at base S. phyllocephalus.
Heads not leafy-bracteate at base.
Phyllaries linear elliptic to oblanceolate; fruits conspicuously beaked.
S. uniserialis var. frutescens.
Phyllaries broadly ovate to obovate; fruits not or scarcely beaked S. divaricatus.
Sclerocarpus divaricatus (Benth.) Benth. & Hook. f. ex
Hemsl. Biol. Cent. Am. Bot. 2: 164. 1881. Gymnopsis divaricata
Benth. Bot. Voy. Sulph. 116. 1844. S. orcuttii Greenm. Field Mus.
Bot. 2: 346. 1912. Figure 99.
Damp or dry thickets or weedy fields, near sea level to 900 m.;
Alta Verapaz; Escuintla; Quezaltenango; Retalhuleu; Santa Rosa;
Suchitepequez. Mexico; Honduras and El Salvador to Costa Rica;
Colombia.
Erect or ascending, often much branched annuals to about 1.5 m. tall, the stems
striate, more or less strigose; leaves petiolate or the uppermost sessile, the upper
leaves alternate, the lower ones opposite, the blades ovate to triangular or lance-
ovate, mostly 2-5 cm. long, acute or acuminate (rarely obtuse), cuneate at the base or
sometimes almost truncate and then abruptly short-cuneate, 3-nerved, the margins
coarsely dentate to subentire, sparsely or densely hirsute or strigose above, usually
densely strigose beneath; heads few or numerous, solitary on peduncles mostly 3-10
cm. long; phyllaries green, broadly ovate to obovate, 5-10 mm. long, acute, strigose-
hispid; pales 6-8 mm. long, pubescent, especially near the apex, turgid when mature
and coarsely tuberculate, closely investing the achene; ray flowers commonly 5-7, the
tube 4-5 mm. long, the ligules deep yellow to orange-yellow, tridentate, 5-10 mm.
long, appressed-pubescent; disc flowers 15-30, 8-10 mm. long, pubescent; fruits (black,
glabrous achene within the closely investing pale) about 3 mm. long, tuberculate;
pappus none.
308 FIELDIANA: BOTANY, VOLUME 24
Sclerocarpus phyllocephalus Blake, Contr. U. S. Nat. Herb.
24: 27, t. 8. 1922.
Damp thickets on plains or hillsides or in thinly wooded
ravines, often a weed in waste ground, near sea level to 900 m.;
Chiquimula; Izabal (type from Cristina, Blake 7648); El Progreso;
Zacapa. Honduras.
Erect or spreading herbs commonly 20-60 cm. high but sometimes attaining
almost 1 m. in height, often much branched, hispid-pilose or strigose with white hairs;
leaves petiolate, the upper ones alternate, the lower ones opposite, the blades ovate or
rhombic-ovate to lance-ovate, mostly 2-8 cm. long, acuminate or acute, cuneate at
the base, triplinerved, the margins coarsely serrate-dentate, thinly or very densely
hispid or strigose above, usually densely strigose beneath; heads few or numerous,
solitary at the ends of the branches, long-pedunculate, subtended at the base by 2, 3,
or more, large or small leaves or bracts; phyllaries uniseriate, very unequal, about 6,
spathulate to oblong-ovate, petiolulate, the margins entire or serrate, 6-20 mm. long;
pales more or less pubescent, densely so near the apex, when mature becoming
somewhat tuberculate and closely investing the achene; ray flowers 5-7, the tube
about 2 mm. long, more or less pubescent, the ligules orange-yellow, 4-7 mm. long,
suborbicular; disc corollas 7-9 mm. long, the limb hispid-pilose; fruits (blackish
achenes within the closely investing pale) rostrate, strigillose, about 6 mm. long,
curved, the body obovoid, 3-4 mm. long, costate and somewhat tuberculate, the beak
about 2 mm. long; pappus none.
Sclerocarpus uniserialis (Hook.) Benth. & Hook. f. ex Hemsl.
Biol. Cent. Am. Bot. 2: 164. 1881. Gymnopsis uniserialis Hook. Icon.
PL 2, t. 145. 1837. Aldama uniserialis (Hook.) A. Gray, Journ.
Boston Soc. Nat. Hist. 6: 228. 1850. Sclerocarpus major Small, Fl.
Southeastern U. S. 1250. 1903.
Erect, branching annuals mostly 0.3-2 m. tall, the stems more or less strigose;
leaves alternate above, opposite below, petiolate, the blades ovate-trullate to
rhombic, mostly 3-6 cm. long, acute or acuminate, more or less cuneate at the base,
the margins coarsely dentate, usually scabrous above, strigose beneath; heads few or
numerous, terminal, on peduncles mostly 3-15 cm. long; phyllaries uniseriate, 5-12,
spreading or reflexed in age, mostly 5-10 mm. long, linear-elliptic to oblanceolate; ray
flowers 5-9, neutral, the ligules orange-yellow, ovate-oblong to suborbicular, 6-20 mm.
long; disc flowers commonly 15-40, yellow or reddish, papillate to strigose; pales 6-10
mm. long, enclosing the corolla to about halfway, later closely investing and
surpassing the achene, narrowing to a beak, not or scarcely bent; achenes 3-6 mm.
long, narrowly fusiform; pappus scales less than 1 mm. long.
Only the following variety occurs in Guatemala.
Sclerocarpus uniserialis (Hook.) Benth. & Hook. f. var.
frutescens (Brandg.) Feddema, Phytologia 23: 206. 1972.
Shrubby slopes, 800-900 m., Huehuetenango. Mexico; British
Honduras.
NASH: FLORA OF GUATEMALA 309
Differs from the typical variety in its somewhat shorter pales,
mostly 5-8 mm. long, at least the outer ones with strongly bent,
narrow beaks, and in its achenes that are obliquely obovate.
SELLOA HBK.
Reference: E. K. Longpre, The systematics of the genera
Sabazia, Selloa and Tricarpha (Compositae), Publ. Mus. Mich. St.
Univ. Biol. 4: 287-383. 1970.
Erect, ascending, or decumbent perennials from short rootstocks, the stems
repent or stoloniferous, simple or branching from the base; leaves opposite, sessile or
nearly so, the basal ones broadly obovate or spathulate to spathulate-oblanceolate,
acute or obtuse, attenuate to the petioliform base, the margins entire or remotely
denticulate, glabrous or thinly pilose, cauline leaves similar but smaller, spathulate to
linear-oblong or linear-lanceolate; peduncles terminal or axillary, solitary or
subcymose; heads radiate; involucres hemispherical or broadly turbinate; phyllaries
2-3-seriate, the outer ones herbaceous, oval-ovate to broadly elliptic-ovate, obtuse,
dark purplish at least along the margins, the inner ones lance-ovate, to oblong-ovate,
acute or apiculate, usually broader than the outer ones; receptacle convex; pales
persistent, linear -setiform; ray flowers fertile, the ligules white on the upper surface,
white or lavender beneath, tridentate or trilobate at the apex; disc flowers numerous,
hermaphrodite, fertile, the corollas yellow or greenish yellow, tubular, shallowly 5-
cleft; style branches recurved; anthers rounded or minutely sagittate at the base,
apically appendaged; ray achenes turbinate or laterally compressed, obscurely 3-5-
angulate, glabrous, epappose or crowned with 5-10 caducous, awnlike squamellae;
disc achenes similar but more obovoid.
Three species, all restricted to high elevations, one in Mexico,
one in Costa Rica, and one in Guatemala.
Selloa obtusata (Blake) Longpre, Publ. Mus. Mich. St. Univ.
Biol. 4: 374. 1970. Sabazia obtusata Blake, Brittonia 2: 346. 1937.
Figure 100.
Rocky, alpine or subalpine meadows, sometimes in open pine or
juniper forest, 3,200-3,800 m., Huehuetenango (type from Sierra de
los Cuchumatanes, Skutch 1265).
Ascending or decumbent perennials 10-22 cm. high, the stems pilose and stipitate-
glandular, repent or stoloniferous, simple or sparsely branched from near the base;
leaves opposite, sessile or nearly so, the basal ones spathulate-oblanceolate or broadly
oblong-obovate, mostly 1-3.5 cm. long, 0.2-0.6 (-0.9) cm. wide, obtuse or rounded at
the apex, attenuate into a petioliform base, the margins entire or with a minute tooth
on each side toward the apex, ciliate, triplinerved, thinly pilose on both surfaces with
mostly gland-tipped hairs, the upper, cauline leaves obovate to spathulate or linear-
oblong, obtuse, mostly 0.5-3 cm. long, 0.2-1 cm. wide; peduncles terminal or axillary,
solitary, mostly 4-10 cm. long; heads radiate; involucres hemispherical, 4-6 mm. high;
phyllaries 2-3-seriate, equal, oval to ovate, obtuse or rounded, ciliate and sparsely
310 FIELDIANA: BOTANY, VOLUME 24
pilosulous with gland-tipped hairs; pales linear, acuminate, 1-3 mm. long, entire,
scarious; ray flowers 11-21, the ligules white, 4-5 mm. long, trilobate; disc flowers 30-
35, greenish yellow, 2-3 mm. long; achenes black, obovoid or turbinate, glabrous, 1-2
mm. long; pappus none.
SIGESBECKIA Linneaus
Reference: J. E. Humbles, Observations on the genus Siges-
beckia, Ciencia y Naturaleza 13: 2-19. 1972; R. McVaugh and C.
Anderson, North American counterparts of Sigesbeckia orientalis
(Compositae), Contr. Univ. Mich. Herb. 9: 485-493. 1972.
Erect, ascending, or creeping herbs, often much branched, glandular-pilose and
usually very viscid; leaves opposite, petiolate or sessile, the blades triplinerved;
inflorescences usually cymose, sometimes becoming paniculate; heads heterogamous,
radiate (in ours); involucres campanulate or hemispheric; phyllaries biseriate, the
outer ones usually 5, herbaceous, linear-oblanceolate to spathulate, stipitate-
glandular, spreading or reflexed, the inner ones erect, usually shorter, broader,
subtending the ray flowers and more or less enclosing the achenes; receptacle slightly
convex, the pales somewhat concave, membranaceous, subtending or enfolding the
disc flowers; ray flowers uniseriate, pistillate, fertile, the ligules white, yellow, or
maroon, usually small, broad, spreading, tridentate; disc flowers hermaphrodite, all
fertile or the inner ones sterile, the corollas regular, tubular, the limb campanulate, 3-
4-5-cleft, anthers obscurely sagittate at the base, apically appendaged; style branches
of the disc flowers short, flattened, papillose; achenes obovoid-oblong, turgid, often
incurved, glabrous; pappus none.
Nine species, in tropical regions of both hemispheres, with three
in Guatemala.
Stems scapelike from a basal rosette, naked or bearing a single pair of small, bractlike
leaves; ray flowers conspicuous, the ligules 5-10 mm. long S. nudicaulis.
Stems leafy, without basal rosette; ray flowers inconspicuous, the ligules 2-3 mm.
long.
Leaves commonly broadest near the middle; phyllaries mostly 3-6 mm. long (in
ours), with stipitate glands 0.2-0.3 mm. long; anthers yellow; denuded
receptacle longer than broad; achenes straight or nearly so S. agrestis.
Leaves commonly broadest near the base; phyllaries mostly 6-14 mm. long (in
ours), with stipitate glands 0.5-0.6 mm. long; anthers green; denuded
receptacles broader than long; achenes more or less curved S.jorullensis.
Sigesbeckia agrestis Poepp. & Endl. Nov. Gen. & Sp. 3: 45, t.
256. 1835. Ton-tzun (Huehuetenango). Figure 101.
Damp pine or pine-oak forest, fields, thickets, sometimes in
sand along streams, or a weed in coffee plantations, 1,350-2,000 m.;
Alta Verapaz; Chimaltenango; Guatemala; Huehuetenango; Sacate-
pequez; Solola. Mexico; El Salvador; Nicaragua; Costa Rica; Haiti;
Colombia; Peru.
NASH: FLORA OF GUATEMALA 311
Erect or ascending annuals, 1-2 m. high, the stems more or less glandular and
villous; upper leaves sessile, the blades lanceolate, lower leaves on winged petioles,
these dilated or auriculate and clasping at the base, the blades thin, lanceolate to
ovate, usually broadest near the middle, mostly 4-10 cm. long, acute or acuminate,
the margins serrate-dentate, truncate or subcordate at the base and abruptly
contracted into the winged petioles, thinly or rather densely long-pilose or villous on
both surfaces and glandular-punctate on the lower surface; heads numerous, 3-5 mm.
high, on glandular peduncles, disposed in irregular panicles; outer phyllaries 5, often
reflexed, spathulate, mostly 3-6 mm. long, with stipitate glands 0.2-0.3 mm. long,
sometimes with long, eglandular hairs also present, the 5-8 inner phyllaries shorter,
broader, also stipitate-glandular; pales obovate, membranous, the central ones often
persistent; ray flowers commonly 5-8, the ligules yellow or orange-yellow, 2-3 mm.
long, tridentate; disc flowers commonly 8-15, the corollas yellow, about 2 mm. long;
anthers yellow; achenes nearly straight, lineate, black, 4-angulate, 1.5-2.3 mm. long;
pappus none.
Sigesbeckia jorullensis HBK. Nov. Gen. & Sp. 4: 283. 1820. S.
cordifolia HBK. I.e. S. serrata DC. Prodr. 5: 496. 1836. Polymnia
odoratissima Sesse & Mocifio, PL Nov. Hisp. 148. 1890. Cua-
nahuatch (Quezaltenango); pega-pega (Jalapa).
Damp to very wet, mixed forest, or in pine, oak, or Cupressus
forest, sometimes in damp thickets or open, usually wet meadows,
1,400-4,000 m.; Chimaltenango; Huehuetenango; Jalapa; Quezal-
tenango; El Quiche; Sacatepequez; San Marcos; Solola; Totonica-
pan. Mexico; Costa Rica; Panama; Haiti; Dominican Republic;
South America.
Erect herbs to 1.5 m. high, the stems more or less glandular and villous; upper
leaves sessile, the blades lanceolate, lower leaves on winged petioles (these more or
less dilated and clasping at the base), the blades triangular- ovate to lanceolate,
usually broadest near the base, mostly 3-12 (-16) cm. long, acuminate or acute, the
margins serrate, truncate or subcordate at the base and abruptly contracted into the
winged petiole, thinly or rather densely pilose or villous on both surfaces and
sometimes inconspicuously glandular-punctate on the lower surface; heads few or
numerous, commonly 5-7 mm. high, on glandular peduncles, disposed in irregular
panicles; outer phyllaries 5, spreading, narrowly linear-spathulate, mostly 6-14 mm.
long, with stipitate glands 0.5-0.6 mm. long, the 5-8 inner phyllaries shorter, broader,
also stipitate-glandular; pales scarious, glandular only at apices; ray flowers 5-8, the
ligules yellow, sometimes drying reddish purple, 1.5-3 mm. long, tridentate; disc
flowers commonly 10-20 (-30), the corollas yellow, about 2 mm. long; anthers green;
achenes curved, black, striate, 2-2.7 mm. long; pappus none.
These plants are unpleasantly viscid, the heads, stems, and
leaves adhering to clothing and even to the hands.
Sigesbeckia nudicaulis Standl. & Steyerm. Fieldiana: Botany
23: 262. 1947.
Known only from the type, Huehuetenango, Sierra de los
312 FIELDIANA: BOTANY, VOLUME 24
Cuchumatanes, above San Juan Ixcoy, along trail to Tojquia, 2,800-
3,400 m., Steyermark 50114.
Erect perennials, 20-30 cm. tall, the stems solitary, scapelike, from a basal
rosette, purplish, densely glandular-pilose, naked or bearing a single pair of small,
sessile, bractlike leaves; basal leaves commonly about 8, on winged petioles, the
blades broadly ovate or rhombic-ovate to obovate, obtuse, 3-6 cm. long, rounded,
subtruncate, or cuneate at the base and usually abruptly contracted into the broadly
winged petiole, the margins undulate-serrate or subentire, densely villous above,
glabrous beneath or nearly so; heads 1-5, the disc about 6 mm. high, 9-12 mm. broad;
peduncles stipitate-glandular, mostly 5-15 cm. long; outer phyllaries 5-8, spreading or
reflexed, linear-oblong to spathulate, 6-12 mm. long, stipitate-glandular, the inner
phyllaries obovate-oblong, 3-5 mm. long, glandular; ray flowers 10-15, the ligules
yellow, 5-10 mm. long, tridentate; disc flowers numerous, the corollas about 2 mm.
long; achenes blackish, angular-obovoid, glabrous, about 2 mm. long; pappus none.
SIMSIA Persoon
References: S. F. Blake, A revision of Encelia and some related
genera, Proc. Amer. Acad. 49: 376-396. 1913; New and noteworthy
Compositae, chiefly Mexican, Contr. Gray Herb. 52: 16-59. 1917. H.
Robinson and R. D. Brettell, A survey of the Mexican and Central
American species of Simsia, Phytologia 24: 361-377. 1972.
Annual or perennial herbs, mostly with harsh pubescence, the stems erect,
branching, leafy; principal leaves opposite (the uppermost often alternate), sessile or
petiolate, the blades often trilobate or hastate, the petioles naked or winged and with
foliaceous bracts often present on stem nodes; inflorescences basically cymose, often
becoming paniculate; heads radiate (in ours); phyllaries 2-4-seriate, subequal or
unequal, mostly lance-ovate to lance-linear; receptacle slightly convex; pales
scarious, rigid, acute or acuminate and usually mucronate, enfolding the achenes and
persistent after their fall; ray flowers neutral, the ligules obscurely bidentate or
tridentate, usually yellow, rarely purple, pinkish, or white; disc flowers hermaphro-
dite, fertile, the corollas yellow or purplish, the tubes short, usually pubescent, the
limb cylindrical, 5-dentate; style branches acuminate or acute, hispidulous; achenes
strongly compressed, obovate or oblong, glabrous or appressed-pubescent, the edges
thin but not winged; pappus of 2 awns or wanting.
About 35 species, all American, in tropical or warm-temperate
regions, with nine in Guatemala.
Rays purple or deep pink (rarely white) S. sanguined.
Rays yellow.
Leaves densely silvery-sericeous beneath.
Achenes densely appressed-pilose, with 2 awns S. sericea.
Achenes glabrous, awnless S. ghiesbreghtii.
Leaves not sericeous beneath.
Heads subsessile or on pedicels about 1 mm. long S. steyermarkii.
Heads on pedicels 1-8 cm. long.
Phyllaries conspicuously unequal, the outermost ones ovate to lance-ovate.
NASH: FLORA OF GUATEMALA 313
Petiole bases sometimes dilated and /or small, rounded, foliar discs
sometimes forming there; disc flowers commonly 20-35 S. holwayi.
Petiole bases not dilated, lacking foliar discs; disc flowers commonly 8-10
(rarely 20) S. lagascaeformis.
Phyllaries subequal or slightly unequal, the outermost ones linear- lanceolate
to almost linear.
Lower leaf surfaces strigose with strongly appressed hairs S. amplexicaulis.
Lower leaf surfaces more or less hispid, setose, or hirsute with spreading
hairs.
Petiole bases of lower leaves sometimes dilated and/or small, foliar discs
sometimes formed at lower nodes; heads (excluding rays) 15-25 mm.
broad; achenes 5-7 mm. long S. grandiflora.
Petiole bases of lower leaves never dilated, lower nodes never with foliar
discs; heads (excluding rays) 7-12 mm. broad; achenes 4-5 mm. long.
S. foetida.
Simsia amplexicaulis (Cav.) Pers. Syn. PL 2: 478. 1807.
Coreopsis amplexicaulis Cav. Descr. PI. 226. 1802. Encelia
mexicana Mart, ex DC. Prodr. 5: 578. 1836, as syn. Cardillo
(Huehuetenango, fide Aguilar). Figure 102.
Damp or dry thickets, brushy plains, along sandy stream beds
in open forest, often a weed in gardens and cornfields, 1,200-3,000
m.; Chimaltenango; Guatemala; Huehuetenango; Jalapa; Quezal-
tenango; El Quiche; Sacatepequez; San Marcos; Solola. Mexico.
Erect annuals, 0.5-1 m. high, usually much branched, the stems often purplish,
glandular-puberulent and hispid; upper leaves frequently alternate, often sessile, the
lower ones opposite, petiolate, the blades mostly 2-14 cm. long, ovate or triangular,
often trilobate near the base, acute, the margins irregularly crenate-dentate, broadly
cuneate to cordate at the base, strigose with strongly appressed hairs, the petioles
often more or less winged and commonly dilated and amplexicaul at the base but not
usually conjoined; inflorescences cymose-paniculate; heads usually numerous, about
1 cm. high, the disc 7-12 mm. broad, on pedicels 1-6 cm. long, the peduncles often
much longer, these glandular-hispid; phyllaries biseriate, subequal or slightly unequal,
5-9 mm. long, lance-linear, acuminate, herbaceous almost throughout, glandular-
hispid, the inner ones often purplish; pales 8-9 mm. long, glandular near the apex,
purplish; ray flowers 8-12, the ligules bright yellow to orange-yellow, about 1 cm.
long; disc corollas about 6 mm. long, puberulent, yellow; achenes black or mottled,
appressed-pubescent, 3-5 mm. long; pappus of 2 awns 2-3 mm. long.
Simsia foetida (Cav.) Blake, Proc. Amer. Acad. 49: 385. 1913.
Coreopsis foetida Cav., Icon. 1: 55, t. 77. 1791.
Clearings in mixed forest or on rocky, brushy slopes, 900-1,100
m.; Guatemala; Huehuetenango. Mexico.
Erect, branching annuals, 1-2.5 m. high, the stems usually densely glandular and
more or less hispid; uppermost leaves alternate, sessile or short-petiolate, the blades
lanceolate, the lower ones opposite, petiolate, the blades triangular-ovate, mostly 3-10
(-12) cm. long, sometimes trilobate or hastate-lobate, acute, truncate or subcordate at
314 FIELDIANA: BOTANY, VOLUME 24
the base, the margins crenate-dentate, both surfaces densely or sparsely glandular
and more or less setose or hirsute, at least along the costae, the petioles of lower
leaves naked, neither dilated nor clasping at the base; inflorescences cymose-
paniculate, the heads 10-14 mm. high, the disc 7-12 mm. broad; phyllaries 3-seriate,
subequal, lanceolate to linear-lanceolate, striate, densely glandular-hispid, the
herbaceous tips spreading; pales about 9 mm. long, appressed-pubescent on the keel
and just below the mucronate tip; ray flowers commonly 8-12, the ligules yellow, 7-8
mm. long; disc corollas about 6 mm. long, puberulent, yellow, becoming purplish near
the apex; achenes blackish, 4-5 mm. long, appressed-pubescent; pappus of 2 awns 3-4
mm. long.
Simsia ghiesbreghtii (A. Gray) Blake, Proc. Amer. Acad. 49:
392. 1913. Encelia ghiesbreghtii A. Gray, op. cit. 8: 658. 1873.
Damp, open forest, often in oak-pine forest, 1,700-2,200 m.; Baja
Verapaz; El Progreso; El Quiche. Southern Mexico.
Erect, coarse, branching herbs about 1.5 m. high, the stems glandular-puberulent
and long-pilose; leaves mostly opposite (the uppermost ones sometimes alternate),
short -petiolate, the blades lanceolate, mostly 4-10 cm. long, long-acuminate, truncate
or rounded at the base or sometimes abscurely hastate-lobate, the margins crentate-
dentate or serrate, the upper surfaces more or less glandular-strigillose, the lower
surfaces densely and softly silvery-sericeous; heads about 1 cm. high, few or
numerous, on short or elongated peduncles, the cymes sometimes disposed in large,
leafy panicles; phyllaries 3-seriate, graduate, glandular-puberulent, hispid-ciliate, and
sometimes pilose, especially at the base of the keels and near the margins; ray flowers
commonly 7-9, the ligules pale yellow, about 1 cm. long; disc corollas yellow, about
6.5 mm. long, at least the tubes puberulent; pales 7-8 mm. long, stiff, acuminate,
more or less glandular-puberulent on keel and apex, otherwise glabrous, the margins
lacerate-dentate; achenes about 3 mm. long, mottled or black, shining, glabrous;
pappus none.
Perhaps only a form of S. sericea (Hemsl.) Blake.
Simsia grandiflora Benth. ex Oersted, Vidensk. Medd. Kjoeb.
1852: 92. 1853. S. polycephala Benth. ex Oersted, torn. cit. 93.
Encelia polycephala Hemsl. Biol. Cent. Am. Bot. 2: 184. 1881. S.
megacephala Sch. Bip. ex Blake, Proc. Amer. Acad. 49: 391. 1913. S.
guatemalensis Robins. & Brettell, Phytologia 24: 372. 1972.
Wet to dry thickets, on plains and hillsides, sometimes on
gravel bars along streams, often a weed in cultivated fields, 500-
1,600 m.; Chiquimula; Escuintla; Guatemala; Jalapa; Jutiapa;
Santa Rosa. Southern Mexico; El Salvador; Honduras; Nicaragua.
Erect, coarse herbs, 1-2 m. high, much branched, the stems densely glandular and
pilose-hispid; uppermost leaves alternate, usually sessile and clasping, rarely very
short-petiolate, the middle and lower ones opposite, petiolate, the petioles more or
less winged and often with a small, foliar disc attached to the base, or scarcely if at
all winged and the base dilated and the disc apparently distinct, or sometimes the
base dilated and the disc absent, the blades ovate or broadly triangular-ovate,
NASH: FLORA OF GUATEMALA 315
sometimes obscurely hastate-lobate, mostly 4-12 cm. long, acuminate, subcordate to
broadly cuneate at the base, the margins crenate-dentate, triplinerved, more or less
glandular-hispid on both surfaces with spreading hairs; heads few or numerous, on
short or elongated peduncles, these densely glandular-pilose, the cymes often disposed
in panicles; heads mostly 1.3-1.5 cm. high, the disc 1.5-2.5 cm. broad; phyllaries 20-40,
green, 2-3-seriate, subequal, or the outer ones slightly shorter, oblong-lanceolate to
almost linear, subacute, 7-12 mm. long, glandular and long-hispid; pales 8-10.5 mm.
long, scarious but usually greenish toward the apex, pubescent or puberulent above,
abruptly acute, the margins laciniate; ray flowers 10-20, the ligules pale yellow, 7-8
mm. long; disc corollas yellow, glandular below, 6-7 mm. long; achenes 5-7 mm. long,
blackish, mottled with gray, more or less appressed-pubescent or almost glabrous;
pappus of 2 awns 3-4 mm. long.
A weedy species, especially variable in leaf and petiole,
indument, and even in heads. According to Robinson and Brettell,
S. guatemalensis has heads with "ca. 20-25 phyllaries and ca. 10-12
rays" while S. grandiflora has heads with "ca. 40 phyllaries and ca.
20 rays." It is often difficult to ascertain the number of ray flowers
on herbarium material as they are so soon deciduous, but counts
made from heads of several specimens (including some cited by
Robinson and Brettell as S. guatemalensis} show that the alleged
differences are not constant. While it is true that many Guatema-
lan specimens tend to have fewer phyllaries and often have fewer
ray flowers than some Nicaraguan specimens, there is considerable
intergradation. For instance, specimens from Guatemala (Molina &
Molina 25110, Standley 77667, and Steyermark 30968) have 22-30
phyllaries and 10-16 ray flowers per head. Specimens from
Honduras (Molina & Molina 24566) have 30-32 phyllaries and 14-20
ray flowers, and one Nicaraguan collection (Molina 23157) has 30-
34 phyllaries and 14-18 ray flowers. One very robust specimen from
Nicaragua (Williams & Molina 42449) has 34-40 phyllaries and 14-
20 ray flowers.
Simsia holwayi Blake, Contr. Gray Herb. 52: 46. 1917.
Known only from the type collection, about 2,000 m., Agua
Caliente, El Progreso, Holway 854.
Erect, branching herbs to about 1 m. high, the stems slender, hispidulous-
puberulent and shortly hispid -pi lose; uppermost leaves alternate, subsessile or short-
petiolate, mostly entire or subentire, sometimes with conspicuous basal lobes, middle
and lower leaves opposite, on naked petioles sometimes dilated at the base and/or
small, rounded, foliar disks about 8 mm. broad often forming there, the blades
triangular-ovate, acute, mostly 5-7 cm. long, broadly cordate at the base,
subhastately dilated at the base, the margins crenate-serrate, densely and rather
softly hispid-pilosulous above, densely and softly grayish pilose beneath;
inflorescences cymose-paniculate, the heads long-pedunculate, 7-11 mm. high, the disc
6-9 mm. broad; phyllaries 3-4-seriate, graduate, 8-9 mm. high, ovate-lanceolate to
316 FIELDIANA: BOTANY, VOLUME 24
lanceolate, acuminate, lineate, densely hispidulous- pilose, ciliate, purplish; pales
abruptly mucronate, about 8.5 mm. long, dorsally and apically pilose; ray flowers
about 10, the ligules yellow, bidentate, 3-3.5 mm. long; disc corollas yellow, stipitate-
glandular, 5-6 mm. long; achenes blackish, appressed-pilose, 4-5.5 mm. long; pappus
of 2 awns about 3.2 mm. long.
Simsia lagascaeformis DC. Prodr. 5: 577. 1836.
Damp thickets, hedges along roads, often a weed in coffee
plantations, 800-1,600 m.; Chimaltenango; Sacatepequez; Zacapa.
Mexico.
Erect, rather coarse annuals, 1-2.5 m. high, usually much branched, the stems
often red or purplish, glandular-pubescent and often pilose, especially near the nodes;
uppermost leaves alternate, often sessile or subsessile, the lower ones opposite,
petiolate, the blades thin, broadly triangular-ovate, very rarely hastate-lobate, acute,
the margins crenate to subentire, truncate or cordate at the base, strigillose and
pilose above, more or less puberulent beneath, often minutely glandular and usually
with some scattered long hairs, at least on the costae, the petioles naked or with very
narrow, inconspicuous margins; inflorescences cymose- paniculate; heads mostly 10-12
mm. high, the disc 5-9 mm. broad; peduncles pilose and short stipitate-glandular;
phyllaries 2-3-seriate, distinctly unequal, the outer ones ovate or lance-ovate, the
inner oblong-lanceolate, acute or acuminate, rather rigid, reddish to dark purple,
glandular-puberulent, ciliate; pales truncate or retuse and mucronate, ciliate near the
apex, glandular, purplish above, 6-8 mm. long; ray flowers 5-8, the ligules 5-8 mm.
long, yellow; disc flowers 10-15 (rarely 18-20), the corollas 5-6 mm. long, glandular-
puberulent, yellow becoming purplish; achenes black or mottled with gray, appressed-
pubescent, 4-5 mm. long; pappus of 2 awns about 3 mm. long (rarely the awns
wanting).
Simsia sanguinea Gray, PL Wright. 1: 107. 1852. Encelia
sanguined Hemsl. Biol. Cent. Am. Bot. 2: 185. 1881. S. sanguinea
f. albida Blake, Verh. Bot. Ver. Brandenb. 65: 120. 1923. S.
sanguinea subsp. albida Blake, Contr. U. S. Nat. Herb. 22: 630.
1924.
Open pine and oak forest, sometimes on rocky, brushy or grassy
slopes, 500-2,200 m.; Baja Verapaz; Chimaltenango; Escuintla;
Guatemala; Huehuetenango; Jalapa; Jutiapa; Sacatepequez. Mex-
ico; El Salvador.
Erect, perennial, branching herbs to about 1 m. high, from thick, somewhat
woody roots, the stems dark red or purplish, glandular-hirsute; principal leaves
opposite, mostly 4-12 cm. long, very variable in form, commonly hastately trilobate
with broad, petioliform bases, these more or less cordate-amplexicaul, the uppermost
leaves alternate, bractlike, linear or lanceolate, the margins of the principal ones
variously lacerate-lobate or sometimes merely crenate-dentate, more or less scabrous
and glandular-hirsute on both surfaces; inflorescences cymose, sometimes forming a
lax, open panicle; heads 10-15 mm. high, long-pedunculate; phyllaries about 3-seriate,
linear to linear-lanceolate, acuminate or subulate, densely glandular and setose,
usually purplish, sometimes reflexing, the outer ones shorter; ray flowers about 10,
NASH: FLORA OF GUATEMALA 317
the ligules pale purple, deep rose-purple, or sometimes pinkish (rarely white),
commonly 8-10 mm. long; disc corollas purple or yellow, 5-8 mm. long, the tubes
glandular-pubescent; pales 8-13 mm. long, glandular- pubescent toward the acute
apices; achenes 4.5-7 mm. long; pappus of 2 awns about 3 mm. long (rarely the awns
wanting).
Simsia sericea (Hemsl.) Blake, Proc. Amer. Acad. 49: 393.
1913. Encelia sericea Hemsl. Biol. Cent. Am. Bot. 2: 185. 1881.
Verbesina argentea Bertol. Fl. Guat. 435. 1840 (type from Volcan
de Agua, Sacatepequez, Velasquez s.n.), not V. argentea Gaud.
1826.
Damp or dry, brushy slopes or open forest, often in oak or oak-
pine forest, sometimes in cornfields, 1,500-3,000 m., the type Salvin
& Godman 133, from Guatemala without definite locality, said to
have come from the Motagua Valley; Sacatepequez; San Marcos;
Solola.
Erect, coarse herbs, 1-2 m. high, much branched, the stems glandular-puberulent
and long-pilose; leaves mostly opposite (the uppermost ones sometimes alternate),
short -petiolate, the blades lance-ovate to lanceolate, mostly 4-10 cm. long, long-
acuminate, truncate or rounded at the base, the margins more or less dentate or
inconspicuously serrate, the upper surface densely glandular-strigillose, the lower
surface densely and softly silvery-sericeous; heads about 1 cm. high, numerous, on
short or elongated peduncles, the cymes often disposed in large, leafy panicles;
phyllaries 3-seriate, graduate, the outer ones ovate-lanceolate, glandular-puberulent,
hispid-ciliate, and sometimes more or less pilose, the inner ones lance-acuminate; ray
flowers commonly 7-9, the ligules pale yellow, about 1 cm. long; disc corollas yellow,
about 6.5 mm. long, the tube and sometimes the limb puberulent; pales about 8 mm.
long, stiff, acuminate, more or less glandular-pubescent on the keel and apex, the
margins lacerate-dentate; achenes 3-3.2 mm. long, mottled, densely appressed-pilose;
pappus awns lacerate toward the base, 1.5-2 mm. long.
Simsia steyermarkii Robins. & Brettell, Phytologia 24: 375.
1972.
Known only from the type collection, Steyermark 42931, trail
between Santa Rosalia de Marmol and Vegas, Zacapa.
Erect, sparsely branched herbs to about 1 m. high, the stems terete, reddish,
more or less puberulent; leaves opposite, short-petiolate, the blades 1.5-3.2 cm. long,
triangular in outline, trilobate, acute, truncate to subcordate at the base, the margins
serrate, more or less scabrous on both surfaces and sparsely pilose; petioles narrowly
winged, the bases of the lower ones dilated and connate; heads 10-12 mm. high,
subsessile (the pedicels about 1 mm. long), 3-several disposed in dense, terminal and
axillary glomerules; phyllaries 3-4-seriate, very unequal, lanceolate to oblong-
lanceolate, acuminate, minutely puberulent and long-hirsute on the margins and
costae; pales with scarious margins, acute, more or less laciniate-serrate, densely to
moderately hirsute, usually more densely so near the apex; ray flowers about 9, the
ligules yellow, 8-10 mm. long; disc corollas 6.5 mm. long; achenes about 4 mm. long,
appressed-pubescent; pappus awns 2, about as long as the body of the achene.
318 FIELDIANA: BOTANY, VOLUME 24
SPILANTHES Linnaeus
Reference: A. H. Moore, Revision of the genus Spilanthes, Proc.
Amer. Acad. 42: 521-569. 1907.
Annual or perennial herbs, erect to prostrate, pubescent or essentially glabrous;
leaves opposite, sessile or petiolate; heads usually radiate, rarely discoid or sometimes
disciform with 2-4 inconspicuous ray flowers, usually long-pedunculate, globose to
conic; phyllaries 2-3-seriate (in ours), ovate to lanceolate; receptacle convex or
elongated; pales complicate; ray flowers pistillate, fertile, the ligules yellow or white;
disc flowers hermaphrodite, fertile, the corolla tubular, the limb ampliate or narrowly
campanulate, 4-5-cleft; style branches obtuse or acute; anthers truncate and entire at
the base or minutely sagittate; achenes often ciliate on the margins, the ray achenes
triquetrous or dorsally compressed, the disc achenes laterally compressed; pappus
bristles (in ours) commonly 2 or none, rarely 1 or 3.
Moore recognized 63 species, in both hemispheres, chiefly in
tropical regions. However, the genus is in serious need of revision,
for it appears that Moore did not have a clear idea of what
constitutes a species or of the characters upon which species of
Spilanthes could be based. His key to the American species is based
largely upon leaf shape and other variable characters. Some of the
species he recognized are evidently valid but not on the characters
by which he separated them. Six species are treated here, but until
the nomenclature of the entire genus is revised, one cannot be sure
that they are correctly named.
Heads discoid or with 2-4 inconspicuous ray flowers; ray and disc flowers white;
achenes with pappus awns (at least those of the inner flowers) often half as long
as the body of the achene S. ocymifolia.
Heads conspicuously radiate; ray and disc flowers yellow; achenes without pappus
awns or with awns about one- third as long as the body of the achene or shorter.
Disc at anthesis commonly 4-5 mm. high; pappus awns conspicuously unequal.
Leaves rounded or acute at the base, essentially glabrous S. filipes.
Leaves almost truncate or subcordate at the base, more or less pilose.
S. uliginosa.
Disc at anthesis commonly 6-12 mm. high; pappus awns (when present) subequal.
Achenes with 2 small, subequal awns S. papposa.
Achenes commonly lacking true awns (sometimes crowned with one or more
tufts of cilia); very rarely with a single short awn.
Leaves narrowly lanceolate or elliptic, the margins essentially entire; stems
usually reddish to very dark red S. phaneractis.
Leaves mostly ovate, sometimes lance-ovate, the margins dentate, crenate-
serrate, or subentire; stems usually not reddish S. americana.
Spilanthes americana (Mutis) Hieron. ex Sodiro, Bot. Jahrb.
29: 42. 1900. Anthemis americana Mutis ex L. f. Suppl. PI. 378.
1781. S. mutisii HBK. Nov. Gen. & Sp. 4: 209. 1820. S. beccabunga
DC. Prodr. 5: 622. 1836. S. lateraliflora Klatt, Bot. Jahrb. 8: 43.
NASH: FLORA OF GUATEMALA 319
1886 (type from Coban, Alta Verapaz, Lehmann 1319). S.
beccabunga var. parvula Robins. Proc. Amer. Acad. 27: 176. 1892. S.
americana var. parvula Moore, Proc. Amer. Acad. 42: 546. 1907. S.
americana var. parvula f. parvifolia Moore, loc. cit. S. americana
var. parvula f. lanitecta Moore, torn. cit. 547 (type from Sansiguan,
El Quiche, Heyde & Lux 3381). Grana de oro (Guatemala); rem
rem'Q en (Quecchi, Alta Verapaz); xux (Huehuetenango).
Damp or wet thickets, fields, or forest, often in marshes or wet
soil along streams, frequent in waste ground, sometimes on open
banks, 300-2,600 (-3,000) m.; Alta Verapaz; Baja Verapaz; Chimal-
tenango; Guatemala; Huehuetenango; Jalapa; Jutiapa; Peten; El
Progreso; Quezaltenango; El Quiche; Sacatepequez; San Marcos;
Santa Rosa; Zacapa. Southeastern United States; Mexico; British
Honduras to El Salvador and Panama; West Indies; northwestern
South America.
Perennials, the stems usually decumbent to prostrate, rarely erect, often rooting
at the lower nodes, pilose with rather long, spreading, whitish hairs, or almost
glabrous; leaves short-petiolate, the blades mostly ovate, sometimes lance-ovate,
mostly 1.5-4 cm. long, triplinerved, acute or obtuse, broadly cuneate to subcordate at
the base, the margins crenate or coarsely dentate, often conspicuously so, sometimes
merely undulate or subentire, villous, pilose, or glabrate; heads radiate, disposed on
long, naked peduncles, these few or numerous, the disc ovoid, 6-12 mm. long, more
elongated in fruit; phyllaries green, 4-5 mm. long, ovate or broadly ovate to lance-
ovate, mostly acute or subacute, sometimes obtuse, appressed, usually ciliate and
more or less pubescent dorsally; ray flowers commonly 8-14, the ligules bright yellow,
4-6 mm. long, spreading, rather conspicuous; disc corollas yellow; achenes dark brown
or blackish, about 1.5 mm. long, strongly compressed, usually ciliate, sometimes
naked, the faces glabrous; pappus awns wanting or rarely an occasional achene with
a single short awn, sometimes appearing to be crowned with one or more tufts of
cilia, but lacking true awns.
Highly variable plants, at least in Guatemalan material, and it
is possible that more than a single species is represented among
those that I have annotated S. americana sens lat. Moore described
several varieties and forms of this species and some of the forms are
probably better marked taxonomic units than some species he
proposed. The whole group is so confused that it does not seem wise
to separate these forms until a revision can be made of the entire
genus. Material referable to Moore's forma lanitecta, including most
of the specimens from British Honduras, is remarkable for the
abundant pubescence of long, soft hairs, and may well represent a
distinct species, however, it would be ill considered to separate this
material until a careful examination can be made of the status of
related species.
320 FIELDIANA: BOTANY, VOLUME 24
Spilanthes filipes Greenm. Proc. Amer. Acad. 35: 314. 1900.
Open, grassy, damp places, often in pine forest, near sea level to
2,200 m., Alta Verapaz; Huehuetenango; Quezaltenango. Yucatan
Peninsula of Mexico; British Honduras; Honduras.
Perennial or perhaps also annual plants, usually erect, sometimes procumbent
and rooting at the lower nodes, often much branched, the stems very sparsely villous-
pilose or almost glabrous; leaves petiolate, the blades broadly ovate to lance-oblong,
mostly 2.5-6 cm. long, acute or obtuse, rounded to acute at the base and sometimes
abruptly short-decurrent on the petiole, the margins crenate-dentate or merely
undulate, almost glabrous on both surfaces or with a few long hairs on the veins
beneath; heads radiate, long-pedunculate, the disc commonly 4-5 mm. high at
anthesis, sometimes to 8 mm. in fruit; phyllaries oblong-ovate, 2-2.5 mm. long, acute
or subacute, ciliate; ray flowers about 5, the ligules yellow, about 4 mm. long; disc
flowers yellow; disc achenes 1.5 mm. long, glabrous, ciliate; pappus of 2 minute,
unequal, awns.
Spilanthes ocymifolia (Lam.) A. H. Moore, Proc. Amer. Acad.
42: 531. 1907. Bidens ocymifolia Lam. Encycl. Meth. Bot. 1: 416.
1783. S. albus L'Her. Stirp. Nov. 1: 7, t. 4. 1784. S. exasperata Jacq.
Icon. PI. Rar. 3: 15, t. 584. 1793. S. ocymifolia f. radiifera A. H.
Moore, Proc. Amer. Acad. 42: 533. 1907. S. ocymifolia var.
acutiserrata A. H. Moore, loa. cit. Duerme-lengua (Jalapa); hierba
de sapo (Jutiapa). Figure 103.
Damp or wet thickets or open fields, often along stream banks
or irrigation ditches, sometimes on sandbars along riverbeds, near
sea level to 1,900 m.; Alta Verapaz; Chimaltenango; Chiquimula;
Escuintla; Guatemala; Huehuetenango; Jalapa; Jutiapa; Quezal-
tenango; Retalhuleu; San Marcos; Santa Rosa; Zacapa. Mexico;
Honduras and El Salvador to Panama; South America.
Erect or ascending annuals, often much branched, sometimes as much as 1 m.
tall but probably more commonly 10-60 cm. tall, the stems often reddish, sparsely
hirsute or glabrate; leaves petiolate, the blades broadly ovate to oblong-ovate, mostly
3-9 cm. long, obtuse or acute, rounded to subacute at the base and often more or less
decurrent on the petiole, the margins crenate-dentate or subentire, essentially
glabrous; heads few or numerous, on peduncles mostly 1-5 cm. long, the disc globose-
ovoid to ovoid, 8-12 mm. long; phyllaries few, 4-5 mm. long, oblong-ovate, subacute,
costulate, usually ciliate, glabrous or nearly so; pales subacute; ray flowers 2-4 or
none, when present the ligules inconspicuous, 2-3 mm. long, white, bilobate; disc
flowers numerous, greenish white to cream-colored; achenes about 2.5 mm. long,
almost black, the margins long-ciliate; pappus of the inner flowers of filamentous
awns often half as long as the body of the achene, those of the outer flowers often
much shorter.
The roots of these plants are often employed as a toothache
remedy; when a piece is chewed for a few minutes, there is a
NASH: FLORA OF GUATEMALA 321
gradual loss of sensation in the mouth, similar to that produced in
dental practice by local anesthesia.
Spilanthes papposa Hemsl. Biol. Cent. Am. Bot. 2: 193. 1881.
Sajon (Quezaltenango).
Damp or wet thickets or rather open forest, occasionally in pine
forest or a weed in cultivated ground, rarely in rather dry, rocky
places, 400-2,500 m.; Alta Verapaz; Baja Verapaz; Peten; Quezal-
tenango; El Quiche; Retalhuleu; San Marcos; Suchitepequez.
Honduras; Nicaragua (type from Chontales); Panama.
Plants usually erect, sometimes decumbent, commonly 30-75 cm. high, the stems
rather stout, sometimes rooting at the lower nodes, simple or sparsely branched,
usually more or less villous; leaves petiolate, the blades oblong-ovate to rounded-
ovate, mostly 3.5-10 cm. long, triplinerved, the margins coarsely crenate to subentire,
obtuse or acute, broadly rounded to cuneate at the base, thinly villous or long-pilose,
principally on the veins beneath, or essentially glabrous; heads radiate, few, long-
pedunculate; phyllaries narrowly oblong or lance-oblong, acute or obtuse, ciliate,
green and glabrous or nearly so; pales glabrous, complicate, more or less rounded at
the apex; ray flowers 10 or more, the ligules yellow, 5-10 mm. long; disc 8-12 mm.
long at anthesis, the disc flowers yellow, puberulent; achenes blackish, about 2 mm.
long, densely ciliate; pappus of 2 short, subequal awns or sometimes only one awn
present.
In general appearance similar to some forms of S. americana
(Mutis) Hieron. but usually a more robust plant, and the achenes of
S. americana normally lack pappus awns.
Spilanthes phaneractis (Greenm.) A. H. Moore, Proc. Amer.
Acad. 42: 543. 1907. S. disciformis Robins, var. phaneractis Greenm.
op. cit. 39: 108. 1903.
In shallow water on shores of lakes or ponds, sometimes in wet
meadows in oak forest, 1,300-1,800 m.; Jalapa; Huehuetenango.
Central and western Mexico; Honduras.
Plants perennial, decumbent to almost prostrate, rooting at the lower nodes,
conspicuously succulent when fresh, glabrous throughout or nearly so, the stems
usually red or dark red, mostly 30-60 cm. long; leaves short-petiolate, the blades very
narrowly lanceolate or linear-lanceolate, sometimes elliptic, mostly 3-6.5 cm. long and
0.3-1 cm. wide, very obtuse or subacute, attenuate to the base, the margins entire or
nearly so, the lateral nerves inconspicuous; heads radiate, on long, slender, almost
glabrous peduncles, the disc ovoid, 6-10 mm. long; phyllaries ovate, glabrous or nearly
so, obtuse or subacute; ray flowers about 10, the ligules yellow, conspicuous; disc
flowers yellow; achenes ciliate; pappus awns wanting.
Guatemalan material has consistently smaller heads than the
Mexican type specimen but otherwise appears to agree. It is possible
that the southern plant might be considered a distinct species, but
322 FIELDIANA: BOTANY, VOLUME 24
in this genus of poorly marked species it seems inadvisable at this
time to add another possible synonym to the great number already
published.
Spilanthes uliginosa Sw. Prodr. Descr. Veg. Ind. Occ. 110.
1788. Acmella uliginosa Cass. Diet. Sci. Nat. 24: 331. 1822. Jaegeria
uliginosa Spreng. Syst. Veg. 3: 590. 1826. Calea savannarum
Standl. & Steyerm. Field Mus. Bot. 23: 257. 1947.
In savannas near sea level, Izabal. Southern Mexico; Nica-
ragua; Costa Rica; Panama; West Indies.
Erect, perennial herbs, simple or sparsely branched above, 40-90 cm. high, the
stems densely villous-pilose with lax, spreading white hairs, the internodes mostly
longer than the leaves; leaves sessile or the lower ones short-petiolate, the blades
triangular-ovate or oblong-triangular, 1.5-3.5 cm. long, 0.7-1.6 cm. wide, obtuse or
subacute, shallowly cordate or truncate at the base, the margins shallowly crenate or
undulate-crenate, villous-pilose on both surfaces with white, multicellular hairs;
heads conic, radiate, solitary or 2-3 at the end of a branch, the peduncles very
slender, 4-7 cm. long; involucres about 3 mm. high; phyllaries few, obscurely biseriate,
ovate, obtuse, about 3 mm. long, sparsely strigillose or almost glabrous; pales thin,
about 3 mm. long, oblanceolate, obtuse; ray flowers few, the ligules yellow, about 2.5
mm. long; disc corollas almost 2 mm. long, the tube short, the throat rather long,
funnelform; achenes minutely pubescent or glabrate, ciliate on the margins; pappus
awns unequal in length.
SYNEDRELLA Gaertner
Erect or procumbent herbs, annual or perhaps sometimes more enduring, strigose
or hirsute; leaves opposite, short-petiolate, the blades 3-nerved, the margins more or
less crenate-serrate; heads heterogamous, radiate, axillary or terminal, sessile, usually
glomerate, sometimes solitary; involucres rather narrowly campanulate; phyllaries
few, the outer 2 or 3 foliaceous, the inner paleaceous; receptacle small; pales narrow,
scarious, subtending the disc flowers at least the outer ones rounded or obtuse at the
apex, somewhat erose and ciliate; ray flowers pistillate, fertile, the corolla with a
filiform tube and a short, 2-3-dentate ligule; disc flowers hermaphrodite, fertile, the
corolla tubular, the limb 4-cleft; stamens 4, the anthers entire or obtuse at the base,
the apical appendage obtuse or almost truncate; achenes of two kinds, those of the
ray flowers ellipsoidal, compressed, smooth, the margins winged, the wings deeply
lobate and appearing almost setaceous, with 2 (rarely more) short, spinelike pappus
awns at the apex, the achenes of the disc flowers narrower, clavate, when mature
more or less striate on one side, tuberculate on the other, not winged, bearing 2-3 (-4)
stiff, scabrous pappus awns at the apex, these becoming more divergent in maturity.
The genus consists of one species.
Synedrella nodiflora (L.) Gaertn. Fruct. & Sem. 2: 456. 1791.
Verbesina nodiflora L. Cent. PL 1: 28. 1755. Ucacou nodiflora
Hitchc. Rep. Mo. Bot. Gard. 4: 100. 1893. Figure 104.
NASH: FLORA OF GUATEMALA 323
Damp thickets, or often a weed in waste or cultivated ground,
near sea level to about 800 m.; Escuintla; Guatemala; Izabal;
Jutiapa; Retalhuleu; Santa Rosa. Southern Mexico; British
Honduras to El Salvador and Panama; West Indies; South
America; naturalized in some parts of the Old World tropics.
Plants usually erect and 20-75 cm. high, often much branched, strigose or
hispidulous; leaves short-petiolate, the blades broadly ovate to elliptic, mostly 2-7 cm.
long, acute or obtuse, abruptly contracted at the broad base and short-decurrent on
the petiole, the margins crenate-serrate or rarely subentire, strigose, more densely so
beneath; heads sessile, densely glomerate (rarely solitary), mostly terminal,
sometimes axillary, the clusters subtended by leaves; involucres 8-10 mm. long, 3-4
mm. broad; outer phyllaries lance-oblong, obtuse or acute, greenish, strigose, the
inner ones narrower, glabrous, thin and chafflike; pales rounded or obtuse at the apex
and often somewhat erose and ciliate; ray flowers commonly 3-6, the ligules bright
yellow, about 2 mm. long; disc flowers commonly 7-12, the corollas about 2 mm.
long; achenes 4-5 mm. long, those of the ray flowers winged, the wings with
ascending, sometimes almost setaceous lobes and with 2 (rarely 3) short, stout pappus
awns, the achenes of the disk flowers not winged, with 2-3 stout, spinose, scabrous
pappus awns about 3 mm. long.
Common weeds of waste ground in many parts of Central
America.
TITHONIA Desfontaines
Reference'. S. F. Blake, Revision of the genus Tithonia, Contr.
U. S. Nat. Herb. 20: 423-436, 1921.
Erect, coarse, often annual herbs or shrubs, with sparse or abundant, often rough
pubescence; leaves alternate or the lower ones opposite, petiolate or sessile, the blades
triplinerved, sometimes lobate, the margins usually dentate; peduncles often dilated
below the heads; heads heterogamous, radiate; involucres hemispheric or broadly
campanulate; phyllaries 2-5-seriate, graduate or subequal, lanceolate or ovate to
oblong or oval, the tips rounded to acute or acuminate; receptacles convex; pales
persistent, rigid, striate, acuminate to aristate, concave, embracing the achenes; ray
flowers 8-20, uniseriate, neutral, the ligules yellow or orange-yellow, emarginate,
minutely bifid, or tridenticulate; disc flowers numerous, fertile, the corolla tube
slender, the limb 5-dentate; anthers obtuse or cordate-sagittate at the base, ovate-
appendaged; style branches slender, recurved, the appendages lanceolate or linear-
lanceolate, acuminate, hispidulous; achenes oblong, thickened or somewhat qua-
drangular, pubescent or glabrous; pappus of 1-2 awns and 4-12 free or united
squamellae, or sometimes the awns wanting, or both awns and squamellae wanting.
Usually showy, often handsome plants, in general appearance
closely resembling various species of Helianthus found in the
United States. Ten species, all Mexican or Central American, with
one, T. diversifolia (Hemsl.) Gray, introduced in the West Indies
and now fairly widespread widespread there, also introduced into
324 FIELDIANA: BOTANY, VOLUME 24
Ceylon, where it has been collected in waste places. Five species are
known in Guatemala.
Phyllaries 4-seriate, their apices rounded or obtuse or sometimes the outermost ones
subacute or rounded and subapiculate.
Leaves (except the uppermost) conspicuously lobate; achenes appressed-pilose, with
pappus commonly of 2 awns and squamellae (rarely the awns wanting).
T. diversifolia.
Leaves not lobate; achenes glabrous, the pappus wanting.
Stems more or less finely hispidulous and sparsely hispid-pilose with short hairs;
involucres 1-2 cm. broad T. pittieri.
Stems usually densely pilose to hispid-pilose with spreading hairs; involucres 2-
3.5 cm. broad T. longiradiata.
Phyllaries 2-3-seriate, their apices acute to acuminate.
Peduncles and outer phyllaries densely pilose with long, spreading, white hairs;
pales long-aristate, the elongated apices 3-4 mm. long; achenes 4-5.5 mm. long,
the 2 pappus awns 1-3.5 mm. long T. tubaeformis.
Peduncles and outer phyllaries merely pilosulous or short-pilose or in age glabrate;
pales acuminate to somewhat cuspidate; achenes 6-7 mm. long, the 2 pappus
awns 3-6 mm. long (the awns early deciduous, especially those of the outer disc
flowers) T. rotundifolia.
Tithonia diversifolia (Hemsl.) Gray, Proc. Amer. Acad. 19: 5.
1883. Mirasolia diversifolia Hemsl. Biol. Cent. Am. Bot. 2: 168, t.
47. 1881. T. diversifolia var. glabriuscula Blake, Contr. U. S. Nat.
Herb. 20: 435. 1921. Mirasol (Alta Verapaz, Chiquimula, Santa
Rosa); k'onon, q'il, sun (Quecchi, Alta Verapaz); quil (Suchitepe-
quez); quil amargo (Guatemala); sajan grande (Jutiapa).
Damp thickets or dry, brushy slopes, 200-2,300 m.; Alta
Verapaz; Chimaltenango; Chiquimula; Escuintla; Guatemala; Ju-
tiapa; Quezaltenango; San Marcos; Sacatepequez; Santa Rosa;
Solola; Suchitepequez. Southern Mexico; British Honduras and
Honduras to El Salvador, Costa Rica, and Panama.
Plants herbaceous or shrubby, 1.5-4 m. tall, the branches stout, at first hispid-
pilose or subtomentose, in age often glabrous or nearly so; leaves alternate, petiolate,
the blades mostly 7-20 cm. long, 4-20 cm. wide, usually deeply 3-5-lobate, cuneate to
subtruncate at the base and then decurrent almost to the base of the petiole, there
minutely auriculate; the margins crenate-serrate, tuberculate-hispidulous above,
usually densely pilose beneath with rather soft, spreading hairs and glandular-
punctate, sometimes glabrate; peduncles stout, mostly 5-20 cm. long, sparsely hispid-
pilose or glabrate; involucres 1.5-4 cm. broad; phyllaries 4-seriate, broadly oval and
usually broadly rounded at the apex, or the outer ones ovate and subacute or
rounded and subapiculate, sparsely appressed-hispid or glabrate; ray flowers 12-14,
the ligules bright yellow to orange, 3-6 cm. long; disc corollas about 8 mm. long,
yellow; pales abruptly pointed, sparsely hispidulous above, 9-11 mm. long; achenes 4-
6 mm. long; pappus awns 2, unequal, 3-4 mm. long, the squamellae 6-10, connate
below, lacerate at the apex, 1.5-2.5 mm. long.
NASH: FLORA OF GUATEMALA 325
A decoction of the leaves, which contain a bitter oil, is
sometimes used as a "remedy" for malaria (especially in the Coban
area of Alta Verapaz) and also employed in the treatment of
eczema and sores on the skins of domestic animals.
Tithonia longiradiata (Bertol.) Blake, Bull. Torr. Bot. Club
53: 217. 1926. Helianthus longiradiatus Bertol. Fl. Guat. 436. 1840.
T. scaberrima Benth. ex Oerst. Vid. Medd. Kjoeb. 1852: 91. 1852. T.
platylepis Sch. Bip. ex Benth. & Hook. Gen. PI. 2: 368. 1873, nom.
nud. Mirasolia scaberrima Benth. & Hook, ex Hemsl. Biol. Cent.
Am. Bot. 2: 168. 1881. T. glaberrima Kuntze, Rev. Gen. 1: 371. 1891.
Gymnolomia scaberrima Greenm. Field Mus. Bot. 2: 268. 1907. Flor
amarilla (Sacatepequez); mirasol (Chimaltenango and Huehue-
tenango); sun (Alta Verapaz); yumo (Jalapa). Figure 105.
Damp or dry thickets or open forest, often in pine-oak forest,
1,100-3,000 meters; Alta Verapaz; Baja Verapaz; Chimaltenango;
Huehuetenango; Jalapa; Jutiapa; Quezaltenango; El Quiche; Saca-
tepequez; San Marcos; Solola; Zacapa. Mexico; El Salvador;
Honduras; Nicaragua; Costa Rica.
Erect, coarse, branched herbs or suffrutescent plants, 1-4 m. tall, the stems
densely pilose with long, spreading hairs; lower leaves opposite, petiolate, the upper
ones alternate, subsessile or short-petiolate, the blades lanceolate to broadly ovate,
not lobate, mostly 9-17 cm. long, 2.5-10 (-14) cm. wide, acuminate, cuneately or
abruptly contracted at the base and decurrent on the petiole, the margins crenate-
serrate, scabrous above, densely and softly pilose beneath with spreading hairs; heads
on elongated peduncles, these sparsely or densely hispid-pilose and hispidulous;
involucres mostly 2-3.5 cm. broad; phyllaries 4-seriate, strongly graduate, 12-18 mm.
long, oval or obovate, rounded at the apex, the outer ones hispid-pilose, usually
ciliate, the inner ones glabrate; ray flowers 15-26, the ligules bright yellow to orange-
yellow, 2.5-4 cm. long; disc corollas yellow, puberulent, 5-7 mm. long; pales acute to
acuminate, not cuspidate, about 8 mm. long, hispidulous near the apex; achenes
glabrous, 3-4.5 mm. long; pappus none.
Tithonia pittieri (Greenm.) Blake, Contr. Gray Herb. 54: 9.
1918. Gymnolomia pittieri Greenm. Proc. Amer. Acad. 39: 101. 1903.
Damp or dry thickets, rocky or grassy slopes, 150-1,700 m.; Alta
Verapaz; Chiquimula; Escuintla; Huehuetenango; Peten; El
Progreso. Mexico; British Honduras; Honduras; El Salvador; Costa
Rica.
Herbs or shrubs, mostly 2-4 m. tall, the branches finely hispidulous and sparsely
hispid-pilose with short hairs; lower leaves opposite, the upper ones alternate, the
uppermost sessile or short-petiolate, the lower ones petiolate, the blades linear-
lanceolate to ovate-lanceolate, not lobate, mostly 7-16 cm. long, acuminate, cuneate
at the base and decurrent on the petiole, the margins crenate-serrate, rather densely
326 FIELDIANA: BOTANY, VOLUME 24
hispidulous above, hispidulous and glandular-punctate beneath or densely soft-pilose;
heads pedunculate; involucres 1-2 cm. broad; phyllaries 4 -seriate, strongly graduate,
7-14 mm. long, the outermost ones obovate-oval to oblong, commonly 3-5 mm. long,
obtuse or subacute, appressed-hispidulous, often ciliate, the inner ones obovate,
glabrate, broadly rounded at the apex; ray flowers about 12, the ligules bright yellow,
1.5-3 cm. long; disc flowers yellow, the corollas hispidulous below, about 5 mm. long;
pales abruptly acute, often sparsely tuberculate above, 7-8 mm. long; achenes
glabrous, about 3.5 mm. long; pappus none.
Tithonia rotundifolia (Mill.) Blake, Contr. Gray Herb. 52: 41.
1917. Tagetes rotundifolia Mill. Card. Diet. ed. 8: 4. 1768. Tithonia
tagetiflora Desf. Ann. Mus. Paris 1: 49, t. 4. 1802. T. aristata Oerst.
Vid. Medd. Kjoeb. 1852: 114. 1852. T. heterophylla Griseb.
Bonplandia 6: 9. 1858. T. speciosa Hook, ex Griseb. Cat. PI. Cub.
155. 1866. T. macrophylla S. Wats. Proc. Amer. Acad. 26: 140. 1891.
T. vilmoriniana Pampanini, Bull. Soc. Bot. Ital. 1908: 133. 1908.
Mirasol.
Damp to dry, open or brushy fields, rocky or grassy slopes, 120-
600 m.; Chiquimula; Escuintla; El Progreso; Retalhuleu; Santa
Rosa; Zacapa. Mexico; British Honduras to El Salvador and
Panama; West Indies (perhaps introduced).
Erect, coarse annuals, 1-3 m. tall, the stems at first densely pilosulous with short
hairs, in age glabrate; leaves alternate, petiolate, the blades rather thin, ovate to
triangular-ovate, mostly 7-20 cm. long, acuminate, cuneate (or sometimes almost
truncate) at the base and then contracted and decurrent on the petiole, simple or
sometimes trilobate, the margins serrate, hispid-pilose on both surfaces, especially on
the veins, scabrous, glandular-punctate beneath; heads long-pedunculate; involucres
2-3 cm. broad; phyllaries biseriate, 1.5-2.5 cm. long, subequal or graduate, the outer
ones lance-oblong to ovate-oblong, acute or acuminate, finely pilosulous, the
herbaceous apex often lax or reflexed, the inner phyllaries similar but usually shorter;
ray flowers 9-13, the ligules golden yellow or orange-yellow, 2-3 cm. long; disc flowers
yellow, the corollas puberulent, about 9 mm. long; pales acuminate to cuspidate,
hispidulous above, 12-18 mm. long; achenes more or less appressed- pilose or glabrous,
6-7 mm. long; pappus awns 2, early deciduous, or those of the outermost flowers
sometimes wanting, 3-6 mm. long, the squamellae united nearly to the apex,
irregularly dentate, about 2 mm. long.
The flowers of this species are reported to yield a good grade of
honey.
Tithonia tubaeformis (Jacq.) Cass. Diet. Sci. Nat. 35: 278.
1825. Helianthus tubaeformis Jacq. PI. Hort. Schoenbr. 3: 65, t. 375.
1798. T. tubaeformis var. bourgaeana Pampanini, Bull. Soc. Bot.
Ital. 1908: 134. 1908. Flor amarilla (Guatemala, Sacatepequez); flor
de sol (Escuintla); mirasol (Escuintla, Jalapa, Jutiapa, Santa
Rosa); mirasol cimarron (Guatemala); sum (Chimaltenango).
NASH: FLORA OF GUATEMALA 327
Damp or dry, open, often rocky slopes or in thickets, common
in cultivated fields, pastures, sometimes in open oak, pine, or mixed
forest, 500-2,500 m.; Baja Verapaz; Chimaltenango; Chiquimula;
Escuintla; Guatemala; Huehuetenango; Jalapa; Jutiapa; Peten;
Sacatepequez; Santa Rosa; Solola. Mexico; Honduras; El Salvador.
Erect, coarse annuals, 0.5-4 m. tall, the stems densely hispid-pilose with
spreading hairs or in age glabrate; leaves alternate, petiolate, the blades ovate to
triangular-ovate, mostly 7-18 cm. long, not lobate, acuminate, cuneately narrowed
from a subtruncate base, the margins crenate-serrate, hispid-pilose above, densely and
softly pilose beneath; heads on long peduncles, these densely hispid-pilose with long,
spreading, white hairs; involucres 1.5-3.5 cm. broad; phyllaries 2-3-seriate, 1.5-3 cm.
long, lanceolate or oblong-lanceolate, acute or acuminate, subequal, the outer ones
densely pilose with long, spreading, white hairs; ray flowers 12-14, the ligules bright
yellow, 1.5-3 cm. long; disc corollas hispidulous, 6-7 mm. long; pales long-aristate, the
elongated apices 3-4 mm. long; achenes pilose, 4-5.5 mm. long; pappus awns 2,
unequal, 1-3.5 mm. long, the squamellae lacerate-fimbriate, 0.3-1.2 mm. long.
TRAGOCERAS HBK.
Reference: Andrew M. Torres, Revision of Tragoceras
(Compositae), Brittonia 15: 290-302. 1963.
Erect or ascending, small, branched annuals; stems terete, pubescent; leaves
opposite, sessile or short-petiolate, the margins entire; heads heterogamous, radiate,
pedunculate or not, solitary at the ends of or in the forks of branches; involucres
cylindrical to campanulate; phyllaries few-seriate, imbricate, graduate, obtuse;
receptacle flat or low-conical, paleaceous; pales hyaline, enclosing the disc flowers,
rounded or almost truncate and often incised at the apex; ray flowers 5-7, pistillate,
fertile, lacking a distinct tube, persistent on the achenes, the ligules white or yellow,
entire or bifid; disc flowers few to numerous, sterile; styles undivided or
inconspicuously bifid; anthers sagittate at the base; ray achenes oblong, dorsally
compressed, becoming tuberculate, especially on the inner face; pappus none.
Five species, all except the following confined to Mexico.
Tragoceras schiedeanum Less. Linnaea 9: 269. 1834. Melam-
podium anomalum M. E. Jones, Contr. West. Bot. 18: 72. 1933.
Figure 106.
Damp or dry, often rocky, open or brushy plains and hillsides,
200-1,400 m.; Baja Verapaz; Chiquimula; Huehuetenango; Jalapa;
Jutiapa; El Quiche; Zacapa. Mexico; Honduras; Nicaragua.
Erect or spreading annuals, often much branched, commonly 10-20 cm. high, the
stems puberulent or strigose; leaves sessile or subsessile, the blades thin, linear to
lance-oblong, 1-3 cm. long, obtuse or acute, rounded or subtruncate at the base,
thinly pilose, the margins entire; heads sessile or on peduncles to 1 cm. long;
receptacle conical; involucres broadly campanulate, 4-6 mm. high; phyllaries very
unequal, obovate, glabrous, green or purplish; ray flowers about 5, the ligules
328 FIELDIANA: BOTANY, VOLUME 24
triangular, white or greenish white, about 3 mm. long, bifid or emarginate at the
apex, becoming more conspicuously cleft in age; achenes about 4 mm. long, tricostate,
becoming tuberculate, especially on the inner face.
TRICHOSPIRA HBK.
Reference: H. Robinson and R. D. Brettell, The relationship of
Trichospira, tribal revisions in the Asteraceae, II, Phytologia 25:
259-261. 1973.
Annual or perennial, chiefly prostrate herbs, the stems often numerous,
branched, densely leafy; leaves alternate or the upper ones subopposite, sessile, the
basal ones petiolate, the blades glabrate or sparsely tomentose above, white-
tomentose beneath, the margins of the cauline leaves crenate or dentate, those of the
basal ones deeply crenate or shallowly pinnatifid; heads homogamous, discoid, many-
flowered, sessile in the leaf axils, the flowers all hermaphrodite and fertile; involucres
depressed; phyllaries few, membranaceous-scarious, usually more or less viscid at
least near the apex, the outer ones shorter, the inner ones subtending the outer
flowers; receptacle flat; pales membranaceous, narrow, flat; corollas regular, bluish
or purplish, the tube short, the limb deeply 5-cleft, the lobes narrow, usually more or
less viscid, the exudation yellowish; anthers minutely sagittate at the base, their
appendages flat; style branches slender, hirtellous, the stigmatic surfaces on the inner
side; achenes dorsally compressed or rare'y triquetrous, oblong-cuneate, the sides 3-4-
costate, truncate, more or less pubescent; pappus bristles usually 2, opposite the
angles of the achene, sometimes with 3-5 small, intermediate squamellae.
The genus consists of a single species.
Trichospira verticillata (L.) Blake, Torreya 16: 105. 1915.
Bidens verticillata L. Sp. PL 833. 1753. T. menthoides HBK. Nov.
Gen. & Sp. 4: 28. t. 312. 1820. Figure 107.
Wet soil on plains or on dried mud, sea level to 900 m.; Peten;
Santa Rosa. Mexico (Chiapas); British Honduras; El Salvador;
Nicaragua; Cuba; tropical South America.
Plants prostrate and often forming mats, the stems stout, sparsely branched,
often rooting at the lower nodes, laxly tomentose, to 30 cm. long; leaves glabrate or
sparsely tomentose on the upper surface, densely white-tomentose beneath, the basal
ones long-petiolate, the blades obovate-oblong, obtuse or rounded at the apex, the
margins deeply crenate or shallowly pinnatifid, the cauline leaves sessile, mostly 2-3
cm. long, obovate or oblong-obovate, rounded at the apex, narrowed to the clasping
base; heads closely sessile in the upper leaf axils, 4-5 mm. high; phyllaries and pales
usually more or less viscid- glandular, at least near the apex, the pales oblanceolate;
corollas 1-2 mm. long, bluish or purplish, at least the limb usually somewhat viscid-
glandular; achenes 3-4 mm. long; pappus awns spinelike and spreading, about 2 mm.
long.
NASH: FLORA OF GUATEMALA 329
TRIDAX Linnaeus
References: B. L. Robinson & J. M. Greenman, Revision of the
genus Tridax, Proc. Amer. Acad. 32: 4-10. 1896; A. M. Powell,
Taxonomy of Tridax (Compositae), Brittonia 17: 47-96. 1965.
Annual or perennial, herbaceous or somewhat suffrutescent plants, erect or
procumbent, more or less pubescent, often somewhat glandular; leaves opposite
(rarely alternate above), petiolate or sessile, simple, trilobate, pinnately cleft, or
pinnatifid, the margins usually serrate or dentate, sometimes entire; heads solitary or
disposed in a cymose panicle, radiate or discoid, long-pedunculate; involucres
campanulate to subcylindrical; phyllaries (1-) 2-5-seriate, subequal or unequal,
imbricate, at least the inner ones scarious, often purple-marginate and longitudinally
striate; receptacle commonly conic, sometimes convex or almost flat; pales scarious;
ray flowers (when present) fertile, the ligules yellow, white, pink, or purplish, obovate
or oblong, obscurely or conspicuously bilabiate, 2-4-dentate or deeply trilobate,
spreading; disc flowers yellow, usually regular, 5-dentate; anthers sagittate at the
base, appendaged at the apex; style branches terminating in long or short, subulate
appendages; achenes turbinate or narrowly obconical, terete or ridged, pubescent,
villous, or rarely glabrous; pappus of the disc flowers of 10-45 plumose or fimbriate
squamellae and/or setae (rarely absent); pappus of the ray flowers similar or
sometimes reduced or absent.
About 26 species, all American, the great majority in Mexico.
Only two are known in Central America, but a third, from Chiapas,
Mexico, is also treated here.
Peduncles densely and conspicuously glandular-pilose; phyllaries 4-5-seriate; rays
pinkish purple, about 15 mm. long; disc achenes crowned with pappus of 10-14
fimbriate squamellae 1-1.5 mm. long T.purpurea.
Peduncles hirsute, if glandular, sparsely or inconspicuously so; phyllaries 2-3-seriate;
rays pale yellow to white, 3-7 mm. long; disk achenes crowned with pappus of
about 20 plumose bristles.
Plants erect; involucres 9-10 mm. high T. platyphylla.
Plants procumbent; involucres 5-8 mm. high T. procumbens.
Tridax platyphylla Robins. Proc. Amer. Acad. 43: 41. 1907. T.
scabrida Brandg. Univ. Calif. Publ. Bot. 6: 73. 1914.
Not reported from Guatemala, but may be expected, as it
occurs in nearby Chiapas, Mexico.
Erect herbs, probably annual, the stems 30-75 cm. high, sparsely to densely
hispid; leaves short-petiolate, the blades broadly rhombic-ovate to ovate-lanceolate,
5-15 cm. long, reduced upward, acuminate, the base rounded or subtruncate and then
cuneate to petiole, the margins coarsely and unevenly serrate- dentate or sometimes
obscurely serrate, scabrous-strigose above, strigose beneath; heads radiate, solitary on
rather stout peduncles, the peduncles usually 2 or 3 from a leaf axil, 2-11 cm. long,
hirsute, sometimes also more or less glandular but usually not conspicuously so;
involucres 9-10 mm. long; phyllaries biseriate, lanceolate to ovate, acute or
acuminate, hispid, striate, the inner ones purplish tinged; pales persistent, glabrous,
330 FIELDIANA: BOTANY, VOLUME 24
oblong, acuminate to cuspidate; ray flowers about 5, the ligules white or pale yellow,
4-7 mm. long; disc corollas pale yellow, 6-8 mm. long; achenes blackish, narrowly
turbinate, 2-3 mm. long, densely pilose; disc pappus of about 20 plumose bristles 2-6
mm. long; pappus of the ray flowers reduced, 1-3 mm. long.
Tridax procumbens L. Sp. PL 900. 1753. Balbisia elongata
Willd. Sp. PI. 3: 2214. 1803. T. procumbens var. ovatifolia Robins. &
Greenm. Proc. Amer. Acad. 32: 7. 1896. Hierba del toro. Figure 108.
Damp or dry, open or brushy fields and hillsides, frequent in
sandy soil, along stream beds, often a weed in waste or cultivated
ground, sea level to 2,300 m. (most common at low elevations); Alta
Verapaz; Chimaltenango; Escuintla; Guatemala; Izabal; Jutiapa;
Peten; El Quiche; Retalhuleu; Sacatepequez; San Marcos; Santa
Rosa; Suchitepequez; Zacapa. Mexico; Honduras and El Salvador
to Panama; West Indies; tropical South America; naturalized in the
Old World tropics.
Plants perennial from a woody base, the stems branching, procumbent, 15-50 cm.
long, sometimes rooting at the nodes, sparsely or densely hirsute; leaves short-
petiolate, the blades broadly rhombic-ovate to ovate-lanceolate, mostly 2-7 cm. long,
acute or acuminate, acute or broadly cuneate at the base, often obscurely or
conspicuously trilobate or subhastate, the margins coarsely dentate or serrate, hirsute
on both surfaces and often scabrous above; heads radiate, solitary, on naked
peduncles mostly 10-20 cm. long, the peduncles hirsute with spreading or retrorse
hairs; involucres broadly campanulate, 5-8 mm. long; phyllaries biseriate, the outer
ones ovate to oblong, hispid-hirsute, acute or acuminate, the inner ones with scarious
margins and dark green costae, acute to minutely cuspidate; pales persistent,
scarious, about 8 mm. long, often purplish striate near the apex; ray flowers 3-6, the
ligules pale yellow to creamy white, suborbicular or oblong, 3-5 mm. long, 2-3-lobate;
disc corollas yellow, 5-7 mm. long; achenes blackish, about 2.5 mm. long, pilose, the
apex truncate; pappus a crown of about 20 plumose bristles, those of the disk achenes
4-7 mm. long, those of the ray achenes reduced, 2-3 mm. long.
One of the most widely distributed and abundant lowland
weeds of all Central America. Poultices of the leaves, usually fried
with lard, are often used as an application to reduce inflammation.
Tridax purpurea Blake, Brittonia 2: 351. 1937.
Damp or dry woods, chiefly in pine-oak forest, 1,800-2,000 m.;
Jalapa; Huehuetenango (type collected near Huehuetenango,
Skutch 1632).
Perennials from rather hard, more or less woody rootstocks, the stems
procumbent, mostly 40-90 cm. long, simple or very sparsely branched, hispid-hirsute,
glandular-pilose above; leaves on short petioles, the blades lanceolate to rhombic-
lanceolate, or elliptic, mostly 3-7 cm. long, acute or obtuse, cuneate at the base and
narrowly decurrent on the petioles, the margins shallowly repand-serrate or subentire,
hispid-hirsute; heads solitary on slender, naked peduncles 1-17 cm. long, the
NASH: FLORA OF GUATEMALA 331
peduncles densely glandular-pilose; involucres 9-12 mm. high, imbricate; phyllaries 4-
5-seriate, oblong or oblong-oval, ciliolate, rounded at the apex, the inner, longest ones
with scarious, purple apices; pales perisitent, obovate, glabrous; ray flowers 3, the
achenes appressed-pilose, about 3.5 mm. long; pappus a crown of 10-14 fimbriate
bristles or squamellae 1-1.5 mm. long.
TRIGONOSPERMUM Lessing
Reference: R. McVaugh and C. W. Laskowski, The genus
Trigonospermum Less. (Compositae, Heliantheae), Contr. Univ.
Mich. Herb. 9: 495-506. 1972.
Suffrutescent annuals or perennials or shrubs, more or less glandular-puberulent;
leaves opposite, on winged petioles or the upper ones subsessile, the blades rhombic-
ovate to elliptic, acute or acuminate, cuneate at the base and /or abruptly contracted
and decurrent on the petiole; inflorescences basically cymose-corymbiform, forming a
large, open panicle; heads small, heterogamous, radiate; involucres biseriate; outer
phyllaries linear to elliptic, the inner ones carinate, obovate, each inner one partially
enclosing an achene; receptacles convex; pales scarious, obovate to linear; ray flowers
1-10, pistillate, fertile, the ligules yellow or white, cuneate to flabelliform, trilobate;
disc flowers hermaphrodite, sterile, the tubular to funnelform corollas often darker
than the ray flowers, the limb 5- (4)-lobate; anthers minutely sagittate, the apical
appendages ovate; styles of the hermaphrodite flowers very minutely bifid; achenes
black, shining, ellipsoid to obovoid, longitudinally striate; pappus none.
Four species, all Mexican, with only one extending into
Guatemala.
Trigonospermum annuum McVaugh & Laskowski, Contr.
Univ. Mich. Herb. 9: 500. 1972. So/an (Chimaltenango). Figure 109.
Damp thickets and fields, rarely in forest, often a weed in
cultivated ground, 1,650-2,100 m.; Chimaltenango; Sacatepequez.
Mexico.
Erect annuals to 1.5 m. tall, branching only in the area of inflorescence, the
stems often purplish, glandular-puberulent; uppermost leaves sessile or subsessile, the
principal leaves on narrowly winged petioles, the blades ovate to elliptic, mostly 4-12
(-20) cm. long, 2-5 (-9) cm. wide, acuminate, obtuse or cuneate at the base,
hispidulous and usually scabrous above, sparsely strigose beneath, especially on
costae and veins and more or less glandular-punctate, the margins serrulate or rarely
subentire; heads pedicellate, the pedicels stipitate-glandular, disposed in pedunculate
cymes arising from the leaf axils; phyllaries more or less puberulent to strigose and
usually glandular, the (4) 5 outer ones mostly 2.5-3.5 mm. long, the inner ones
partially enclosing the achenes 3.5-5.5 mm. long, apiculate; pales scarious, the apices
erose and sparsely ciliate; ray flowers commonly 3, the ligules yellow, broad, more or
less flabelliform 2.5-6 mm. long, 3-9 mm. broad, deeply trilobate; disc flowers 10-20,
funnelform, acutely 5-lobate; achenes broadly elliptic to obovoid, 3-4 mm. long,
tipped by the persistent style-base.
332 FIELDIANA: BOTANY, VOLUME 24
Previously reported as T. melampodioides DC.
VERBESINA Linnaeus
References: B. L. Robinson & J. M. Greenman, Synopsis of the
genus Verbesina, with an analytical key to the species, Proc. Amer.
Acad. 34: 534-566. 1899; S. F. Blake in Standl. Contr. U. S. Nat.
Herb. 23: 1572-1586. 1926.
Herbs, shrubs, or small trees; leaves opposite or alternate, entire, lobate, or
pinnatifid, the margins dentate or subentire, often scabrous, frequently decurrent as
wings on the petioles and on the stems; heads small or large, radiate or discoid,
usually numerous and disposed in more or less corymbose panicles, but sometimes
solitary on long, terminal peduncles; involucres hemispheric or campanulate,
sometimes much shorter than the disc flowers; phyllaries 2-6-seriate, ovate to linear;
receptacle generally conic; pales concave, enfolding the laterally compressed disc
achenes, sometimes squarrose in age; ray flowers (when present) pistillate and usually
fertile, sometimes sterile, (rarely neutral), the ligules white, yellow, or orange, short
and inconspicuous or large and showy; ray pappus of 2-3 awns; disc flowers fertile,
the corollas regular, the tube short, the limb cylindric, 5-dentate; anthers not
appendaged at the base; style tips acute or attenuate; achenes glabrous or upwardly
pubescent, sometimes tuberculate, strongly compressed laterally, usually oblong or
obovate, usually winged on each margin, rarely on only one margin; disc pappus of 2
(rarely 1) deciduous or persistent, usually straight awns, rarely obsolete.
One of the largest composite genera of tropical America, with
about 250 species. All are American, and mostly tropical, a few
extending into temperate areas. A few species besides the 30 listed
here are known from other parts of Central America.
Heads yellow or orange, the rays (when present) also yellow or orange.
Achenes, at least the outer ones, commonly with only one awn.
Leaf blades mostly rhombic-ovate, 3-11 cm. wide V. sousae.
Leaf blades narrowly elliptic to linear-lanceolate, less than 2 cm. wide.
V. eperetma.
Achenes commonly with two awns (rarely the sterile ray achenes with 3 awns).
Stems winged.
Principal leaves ovate to broadly ovate in outline, lobate or pinnatifid.
Heads discoid.
Outer phyllaries obovate to spathulate, rounded or obtuse V. fraseri.
Outer phyllaries lanceolate to linear-lanceolate, acute or subacute.
V. crocata.
Heads radiate V. greenmanii.
Principal leaves narrowly lance-elliptic or lance-oblong, never lobate nor
pinnatifid; heads radiate V. neriifolia.
Stems not winged.
Lower leaf surfaces white, with dense, usually strigillose indument, or white
and silvery with appressed, microscopic hairs.
Leaves opposite; ligules of ray flowers 10-15 mm. long V. hypoglauca.
Leaves alternate; legules of ray flowers less than 5 mm. long or wanting.
NASH: FLORA OF GUATEMALA 333
Leaf blades elliptic, lanceolate, or lance-oblong, the hairs microscopic.
V. hypargyrea.
Leaf blades ovate to rhombic-ovate, the hairs not microscopic.
V. abscondita.
Lower leaf surfaces green, but may be more or less strigillose to pilose or
tomentose.
Ray flowers commonly only 5 or 6 V. pleistocephala.
Ray flowers commonly 8-14.
Heads small, in anthesis 2-4 mm. broad; ligules of ray flowers commonly
2-3 mm. long.
Leaves narrowly lanceolate to narrowly oblanceolate, usually about 4
times longer than broad, attenuate to the base V. steyermarkii.
Leaves ovate, rhombic-ovate, or oblanceolate, usually 2-3 times longer
than broad, the base cuneate, then decurrent on the petiole.
Phyllaries more or less cuspidate V. agricolarum.
Phyllaries rounded or obtuse to subacute, not cuspidate.
Leaf blades ovate to rhombic-ovate; heads few, disposed in small
corymbs often surpassed by the leaves V. abscondita.
Leaf blades lanceolate or oblong to elliptic; heads numerous in
broad corymbs V. perymenioides.
Heads larger, in anthesis mostly 6-10 mm. broad; ligules of ray flowers 4-
15 mm. long.
Ligules 12-15 mm. long V. calciphila.
Ligules 4-10 mm. long.
Leaves 2-3 times longer than broad, more or less scabrous above,
commonly soft-pilose beneath, often densely so; phyllaries 6-9
mm. long V. apleura.
Leaves 4-5 times longer than broad, smooth above or nearly so,
glabrous beneath or inconspicuously appressed-puberulent; phyl-
laries 3-4 mm. long.
Lower leaf surfaces finely and inconspicuously appressed-puberu-
lent; ligules about 8 mm. long V. chiapensis.
Lower leaf surfaces essentially glabrous from the first; ligules 4-5
mm. long V. persicifolia.
Heads white, greenish white, or if the disc flowers (rarely) greenish yellow, the ray
flowers (when present) always white.
Principal leaves undivided, the margins merely dentate or serrate.
Stems winged by the decurrent petioles or leaf bases.
Leaves essentially glabrous beneath; achenes more or less puberulent.
V. punctata.
Leaves densely short -pilose beneath; achenes sparsely and minutely tubercu-
late V. petzalensis.
Stems not winged.
Petioles broadly winged to the base, there dilated and clasping.
V. guatemalensis .
Petioles naked, or if winged, not dilated nor clasping at the base.
Leaves glabrous or glabrate.
Leaf blades mostly 4-10 cm. long; rays 1.5-2 mm. long V. minarum.
Leaf blades mostly 10-20 cm. long; rays 15-20 mm. long V. standleyi.
Leaves more or less pilose or hispidulous, at least on the lower surfaces.
334 FIELDIANA: BOTANY, VOLUME 24
Upper leaf surfaces scabrous, the margins coarsely dentate; phyllaries
acute to acuminate, mucronulate V. scabriuscula.
Upper leaf surfaces smooth, the margins inconspicuously dentate or
subentire; phyllaries obtuse V. lanata.
Principal leaves usually deeply lobate or pinnatifid, rarely some of them merely
undulate-lobate as in V. turbacensis.
Stems conspicuously winged.
Branches and stems densely pubescent V. turbacensis.
Branches and stems glabrous or nearly so V. hypsela.
Stems not winged.
Branches and stems densely tomentose or puberulent V. sublobata.
Branches and stems glabrous or only sparsely puberulent.
Leaves with only 1-3 rounded lobes on each side, the margins serrate.
V. holwayi.
Leaves with usually more than 3 (commonly 4-5) acute or acuminate lobes
on each side, the margins mostly entire or subentire.
Petioles broadly winged, auriculate-amplexicaul at the base V. gigantea.
Petioles not winged V. giganteoides.
Verbesina abscondita Klatt, Leopoldina 20: 93. 1884. V.
smithii Robins. & Greenm. Proc. Amer. Acad. 32: 46. 1896.
Not reported from Guatemala, but may be expected, as it
occurs in nearby Chiapas, Mexico.
Branching shrubs to about 3 m. tall, the stems not winged, at least the young
branches tomentose; leaves alternate, short-petiolate, the blades ovate to rhombic-
ovate, the principal ones mostly 3-10 (-15) cm. long, acute or acuminate, cuneate at
the base and more or less decurrent on the short petiole, the margins serrate- dent ate,
scabrous above, tomentose beneath; heads disposed in small corymbs often surpassed
by the leaves, the short pedicels tomentose; involucres campanulate, about 5 mm.
high; phyllaries about 3-seriate, elliptic-oblong, rounded at the apex or subacute,
more or less pubescent, ciliolate; pales puberulent, more densely so toward the
subacute apex; ray flowers about 8, the ligules yellow, 2-3 mm. long; disc flowers 30-
35; achenes 3-4 mm. long, winged; pappus of 2 subequal awns.
Verbesina agricolarum Standl. & Steyerm. Field Mus. Bot.
22: 319. 1940. Vara de sum (Escuintla).
Damp thickets or oak forest, 100-2,300 m.; Chimaltenango (type
from Finca La Alameda, J. R. Johnston 1026); Escuintla;
Sacatepequez. Honduras; Nicaragua.
Erect, branching shrubs, 1.5-3 m. high, the branches viscid-puberulent and more
or less pilose with short, spreading, white hairs; leaves alternate, on petioles 0.5-2.5
cm. long, or the uppermost sessile, the blades broadly rhombic-ovate to oblong-ovate,
mostly 6-12 cm. long and 2.5-8 cm. wide, acute or short-acuminate, rather abruptly
narrowed to a cuneate base and decurrent on the petiole, scabrous above, pilose
beneath with short but somewhat matted hairs, the margins closely dentate or
undulate-dentate; inflorescences corymbiform, mostly 3-8 cm. broad; heads few or
numerous, about 5 mm. long and 2-3 mm. broad in anthesis, in age about 5 mm.
NASH: FLORA OF GUATEMALA 335
broad, on slender pedicels to 8 mm. long; phyllaries about 3-seriate, mostly linear-
oblong, sparsely short-pilosulous, more or less cuspidate, or the outermost ones
sometimes obtuse or subacute, commonly somewhat reflexed; ray flowers 8-10, the
ligules yellow, 2-3 mm. long; pales acute to subacute; disc flowers commonly 16-24;
achenes 2-2.5 mm. long, blackish and glabrous or sometimes white-papillose, usually
rather broadly winged; pappus awns often about as long as the body of the achene.
Verbesina apleura Blake, Contr. Gray Herb. 52: 53. 1917. V.
apleura var. foliolata Standl. & Steyerm. Field Mus. Bot. 23: 109.
1944.
Damp or wet forest or thickets, 1,400-3,600 m.; Baja Verapaz;
Chimaltenango; Guatemala; El Progreso; Quezaltenango; El
Quiche; Sacatepequez; San Marcos; Solola; Totonicapan. Mexico
(Chiapas).
Shrubs or small trees to about 6 m. tall (rarely weak and subscandent), the
branches sordid-pilose with lax, subappressed hairs; leaves alternate, petiolate, the
blades broadly ovate to lanceolate, mostly 10-15 (-20) cm. long, 4-7 (-10) cm. wide,
acuminate to long-acuminate, the base rather abruptly or gradually cuneate and
more or less decurrent on the petiole, the margins irregularly serrate, usually scabrous
to thinly scaberulous above but sometimes short-pilose, usually rather densely soft-
pilose beneath; heads numerous, on pedicels mostly 1-3 cm. long, disposed in large or
small rounded or flat-topped panicles; involucres commonly about 1 cm. broad;
phyllaries 3-seriate, 6-9 mm. long, the outermost obovate to spathulate, obtuse,
herbaceous, the inner ones lanceolate, acute or acuminate, sparsely pilose; ray
flowers about 12, the ligules yellow, 5-10 mm. long; disc corollas numerous, yellow, 5-
6 mm. long, pilose on the tube; pales acuminate, usually cuspidate, appressed-
pilosulous, about 7.5 mm. long; achenes blackish, about 6 mm. long, sometimes
pubescent near apex, broadly winged when mature; pappus awns 2, subequal, 3-4
mm. long.
Immature Mexican specimens have sometimes been confused in
herbaria with V. perymenioides Sch. Bip. ex Klatt, which has
smaller heads (in fruit only 5-7 mm. thick).
Verbesina calciphila Standl. & Steyerm. Field Mus. Bot. 23:
262. 1947. Dalia de monte.
Damp forest, sometimes in oak-pine forest, sometimes on
limestone cliffs, 2,400-3,700 m.; Huehuetenango (Sierra de los
Cuchumatanes; type collected between Tajquia and Caxin bluff,
Steyermark 50132); Totonicapan.
Shrubs or small trees to 6 m. high, the branches slender, when young densely
hispidulous and scabrous; leaves alternate, sessile or on petioles to 3 mm. long, the
blades oblanceolate or oblong-oblanceolate, mostly 4-14 cm. long, 1.5-5 cm. wide,
acuminate, narrowing to a cuneate base, the margins serrulate, the upper surfaces
very scabrous, the lower surfaces glabrate or sparsely hispidulous except on costae
and veins, there the indument often dense; inflorescences laxly corymbose-
336 FIELDIANA: BOTANY, VOLUME 24
paniculate; heads few or numerous, on pedicels 1-4 cm. long; involucres 8-10 mm.
high and broad; phyllaries about 3-seriate, laxly imbricate, sometimes spreading, the
outer ones oblanceolate to oblong or subspathulate, obtuse or acute, scabrous or
appressed-hispidulous, ciliolate, at least on the lower half; pales 6-8 mm. long, acute;
ray flowers 8-12, the ligules bright yellow, 12-15 (-20) mm. long; achenes cuneate-
oblong, about 4 mm. long, glabrous, ciliate, when immature appearing very narrowly
winged or merely ciliate, but in maturity the wings about 0.5 mm. wide; pappus awns
about 4 mm. long.
Verbesina chiapensis Robins. & Greenm. Proc. Amer. Acad.
34: 554. 1899.
Not reported from Guatemala, but may be expected as the type
collection is from Chiapas, Mexico, 1,200-1,700 m.
Plants probably shrubby, the stems striate, glabrous, the young branches
appressed-puberulent; leaves alternate, short-petiolate or subsessile, the blades lance-
oblong, attenuate to each end, mostly 15-22 cm. long, the margins inconspicuously
serrulate, the upper surface essentially glabrous, the lower one finely appressed-
puberulent; inflorescences corymbose, about 15 cm. broad; heads 20-30, pedicellate,
about 1.2 cm. in diameter (excluding the rays); pedicels pubescent; phyllaries 2-3-
seriate, 3-4 mm. long, ovate-oblong, subacute; ray flowers about 12, the ligules
yellow, about 8 mm. long; disc flowers probably yellow; immature achenes obovate,
about 2 mm. long, puberulent, winged; pappus awns 2, about as long as the achene.
Verbesina crocata (Cav.) Less. Syn. Gen. Comp.: 232. 1832;
DC. Prodr. 5: 617. 1836. Bidens crocata Cav. Icon. 1: 66, t. 99. 1791.
Spilanthes crocata Sims, Curt. Bot. Mag. 39, t. 1627. 1814.
Platypteris crocata HBK. Nov. Gen. & Sp. 4: 201. 1820.
Damp thickets, 1,000-1,400 m., Huehuetenango. Mexico; Costa
Rica; Nicaragua.
Shrubs, 2-3 m. high or the branches weak and more or less scandent, the stems
conspicuously winged, hispidulous or glabrate; leaves opposite, petiolate, the blades
thin, the uppermost, smaller ones ovate, the margins entire or dentate, the principal,
larger ones ovate to broadly ovate in outline, at least the lower ones lobate to
pinnatifid and coarsely dentate, acuminate, rather abruptly cuneate and decurrent on
the petiole, scabrous on the upper surface, less scabrous beneath; heads pedunculate,
solitary or several at the ends of the branches, discoid, 1-3 cm. broad; involucres 6-11
mm. high; phyllaries 4-5-seriate, lanceolate to linear-lanceolate, acute to acuminate;
corollas orange-yellow or reddish orange, about 7 mm. long; achenes 6-7 mm. long,
broadly winged; pappus awns 2, equal, 2-3 mm. long.
Verbesina eperetma Blake, Proc. Biol. Soc. Wash. 60: 43.
1947.
Known only from the type collection, Steyermark 42933, Sierra
de Las Minas, trail between Santa Rosalia de Marmol and Vegas,
Zacapa.
NASH: FLORA OF GUATEMALA 337
Large, coarse herbs or shrubs to 3 m. tall, the stems glabrous or nearly so, not
winged but marked by lines decurrent from the leaf bases; leaves alternate, subsessile
or short- petiolate, the blades narrowly elliptic to linear-lanceolate, mostly 13-16 cm.
long, 1.5-1.8 mm. wide (except those subtending the inflorescence much reduced),
acuminate, somewhat falcate, attenuate to the base and decurrent on the short
petiole, the margins revolute, obscurely serrulate, glabrous or nearly so, feather-
veined with about 40 pairs of lateral veins; heads 1-5 at the ends of the branches, the
peduncles and petioles minutely appressed-pubescent; involucres about 1 cm. high,
broadly campanulate; phyllaries 3-4-seriate, the outermost ones narrowly lance-linear
with wide-spreading herbaceous tips, the others ovate or oblong with herbaceous,
glabrous, acute appendages; pales acute or acuminate, about 12 mm. long; disc
corollas yellow, about 7.5 mm. long; achenes 5-7 mm. long, hispidulous, narrowly
winged, those of the outer flowers with only one pappus awn, those of the inner
flowers with either one pappus awn or two unequal ones, 4-6 mm. long.
Because the heads of the only specimen are very mature, it is
not known whether they are discoid or radiate.
Verbesina fraseri Hemsl. Biol. Cent. Am. Bot. 2: 187. 1881. V.
fraseri var. nelsonii Donn.-Sm. Bot. Gaz. 23: 9. 1897 (type from
Huehuetenango, Nelson 3551). Arnica (Guatemala); capitaneja
(Sacatepequez); mirasol de bejuco (Chiquimula). Figure 110.
Damp or wet thickets, damp to dry forest, sometimes in
roadside hedges, occasionally a weed in fields, 500-2,000 m.; Alta
Verapaz; Chimaltenango; Chiquimula; Escuintla; Guatemala;
Huehuetenango; Quezaltenango; Sacatepequez (type from Duenas,
Fraser, Salvin, & Godman s.n.); Santa Rosa; Solola. Honduras and
El Salvador to Costa Rica.
Large, coarse, branching herbs, the stems usually weak, arching or subscandent*
sometimes as much as 5 m. long, conspicuously winged, hispidulous or glabrate;
leaves opposite, on broadly winged petioles, the blades thin, the uppermost, smaller
ones ovate, the margins entire or dentate, the principal, larger ones mostly 10-20 cm.
long, ovate to broadly ovate in outline, usually deeply pinnatifid with 3-7 broad,
acute, coarsely dentate or with somewhat lobate segments, acute or acuminate,
nearly truncate at the base and decurrent on the petiole, forming the broad wings,
very scabrous on the upper surface, scabrous beneath, with prominent, reticulate
venation; heads few, long-pedunculate, often 1-3 at the ends of the branches, discoid,
2-3 cm. broad; involucres about 1 cm. high; outer phyllaries obovate to spathulate,
rounded or obtuse at the apex, viscid-scabrous, the inner ones narrower, obtuse or
acute; corollas orange, about 8 mm. long; achenes glabrous, 8-9 mm. long, with broad,
thin wings; pappus awns 2-3 mm. long.
Verbesina gigantea Jacq. Icon. PI. Rar. 1: t. 175. 1781; DC.
Prodr. 5: 615. 1836. V. myriocephala Sch. Bip. ex Klatt, Leopoldina
23: 144. 1887.
Wet to dry thickets, fields, hedgerows, on plains, hillsides, or in
ravines, mostly 120-1,100 m. (rarely 3,000 m.); Chiquimula;
338 FIELDIANA: BOTANY, VOLUME 24
Escuintla; Guatemala; Huehuetenango; Retalhuleu; Solola; Suchi-
tepequez. Mexico; British Honduras to El Salvador and Panama;
West Indies.
Erect, coarse herbs, mostly 2-4 m. high, rarely branching below the inflorescence,
the stems thick, sometimes reddish, not winged, glabrous or nearly so; leaves
alternate, on broadly winged petioles, these auriculate-amplexicaul at the base, the
blades thin, more or less ovate in outline, deeply pinnatifid, mostly 15-50 cm. long,
the upper surface sparsely or densely scabrous, the lower surface densely sericeous or
strigose or occasionally glabrate in age, the lateral lobes 3-5 pairs, acute or
acuminate, the margins essentially entire or subentire (sometimes the larger, lower
leaves irregularly dentate); inflorescences mostly 10-30 cm. broad, the panicles
flattopped or rounded; heads numerous, pedicellate; involucres turbinate-camp-
anulate, 4-6 mm. high; phyllaries pale, linear to lanceolate or oblanceolate, ciliate,
more or less pilosulous, acute or subacute; ray flowers 5-6, inconspicuous, the ligules
white, 2-3 mm. long; achenes 4-5 mm. long, sparsely strigillose, broadly winged;
pappus awns 2, 2-3 mm. long.
Verbesina giganteoides Robins., Proc. Amer. Acad. 47: 213.
1911. Mano de lagarto (Peten).
Damp, brushy slopes or open, rocky places, 300-2,000 m.;
Chiquimula; Peten. Mexico (Chiapas); Honduras; Nicaragua; Costa
Rica.
Erect, coarse, herbaceous or shrubby plants, 1-5 m. high, the stems thick, often
red or purple, wingless, essentially glabrous; leaves alternate, on wingless petioles 5-16
cm. long, the blades more or less ovate in outline, deeply pinnatifid, mostly 15-40 cm.
long, 10-25 cm. wide, the lateral lobes (3-) 5-6 pairs, acute or acuminate, entire or
undulate, the upper surface smooth or scaberulous, somewhat pubescent or glabrate,
the lower surface softly pubescent; inflorescences large, often 30 cm. broad, the
panicles rounded or flat-topped; heads numerous, pedicellate, radiate; involucres
turbinate-campanulate, 4-6 mm. high; phyllaries pale but the outermost ones dark-
tipped near the apex, linear to oblanceolate-oblong, ciliate, more or less pilosulous,
acute; ray flowers 5-6, the ligules white, about 4 mm. long; disc flowers about 15,
white; achenes 3-3.5 mm. long, sparsely pubescent, winged; pappus awns 2, subequal,
about 2 mm. long.
Verbesina greenmanii Urban, Symb. Ant. 5: 265. 1907. V.
pinnatifida Cav. Icon. 1: 67, t. 100. 1791, not V. pinnatifida Sw.
(1788).
Not reported from Guatemala but may be expected there as it
has been collected in nearby Chiapas, Mexico.
Shrubs or trees to 6 m. high or more, the stems broadly winged; leaves opposite,
on winged petioles, the uppermost blades entire or lobate, the principal blades more
or less broadly ovate in outline but sinuately 3-5-lobate, to 20 cm. or more long,
contracted at the base and then cuneately decurrent on the petiole, scabrous on both
surfaces; heads radiate, on pubescent pedicels 1-3 cm. long; involucres campanulate,
5-6 mm. high; phyllaries 3-seriate, the outermost 2 very small, ovate, acute, the inner
NASH: FLORA OF GUATEMALA 339
third series narrowly lanceolate, acuminate to somewhat cuspidate, puberulent; ray
flowers 8-12, the ligules yellow, 3-4 mm. long; disc flowers numerous; achenes about 4
mm. long, essentially glabrous, broadly winged; pappus awns 2, unequal, 2-3 mm.
long.
Verbesina guatemalensis Robins. & Greenm. Proc. Amer.
Acad. 34: 550. 1899. V. medullosa Robins, op. cit. 45: 411. 1910 (type
from Fiscal, Guatemala, Deam 6250). V. guatemalensis var.
glabrata Standl. & Steyerm. Field Mus. Bot. 23: 109. 1944.
Juronero bianco (Chiquimula).
Damp or dry thickets on hillsides and in ravines, often on open,
rocky slopes, rarely in coniferous forest, usually 250-1,350 m., rarely
to 2,800 m.; Chiquimula; Guatemala (type from Palin, J. D. Smith
2860); Jalapa; Jutiapa; Quezaltenango; El Quiche; San Marcos;
Santa Rosa; Zacapa. Honduras; El Salvador.
Shrubs or small trees, 1.5-5 m. high, the branches stout, wingless, tomentose to
glabrate; leaves alternate, subsessile or petiolate, the blades thin, lanceolate or lance-
oblong, mostly 7-25 cm. long, acuminate, broadly cuneate at the base and then rather
abruptly attenuate and decurrent to the base of the petiole, there dilated and usually
more or less auriculate, the upper surface short-pilose with very slender, commonly
deciduous hairs, in age usually glabrate and smooth to the touch, rarely scabrous, the
lower surface glabrous or glabrate or rarely somewhat pilose, the margins serrate,
undulate-dentate, or subentire; inflorescences broadly corymbiform; heads usually
very numerous, short-pedicellate or subsessile, in flower 7-8 mm. high and 6 mm.
broad, in fruit about 10 mm. in diameter; ray flowers small and inconspicuous, the
ligules white, bi- or tridentate; involucres 6-7 mm. high; phyllaries linear or lance-
linear, stramineous, densely ciliate, acute or obtuse, pilosulous or glabrate; achenes
obovate, about 5 mm. long, broadly winged; pappus awns 2, equal.
Verbesina holwayi Robins. Proc. Amer. Acad. 51: 539. 1916.
Damp thickets or forest, often in oak forest, sometimes in deep
sand, 1,800-3,000 m.; Huehuetenango; Quezaltenango (type from
Quezaltenango, Holway 96).
Coarse, herbaceous or perhaps sometimes suffrutescent plants, 1-3 m. high, the
stems stout, purplish brown, not winged, glabrous; leaves alternate, on broadly
winged petioles, these auriculate-amplexicaul at the base, the blades ovate or
triangular-ovate in outline, shallowly lobate, mostly 12-20 cm. long and 6-14 cm.
wide, the lobes few, commonly 1-3 on each side near the base, unequal, somewhat
falcate, rounded at the apex, the margins irregularly serrate, the upper surface very
scabrous, the lower surface softly tomentulose or short-pilose; panicles 10-30 cm.
broad, flat-topped; heads long-pedicellate, radiate; involucres turbinate-campanulate,
4-5 mm. high; outer phyllaries linear, unequal, sordid-pubescent, subacute, the inner
ones spathulate-oblong, mucronulate; ray flowers inconspicuous, the ligules white,
about 2 mm. long; achenes black, 3-4 mm. long, sparsely pubescent or more or less
tuberculate, broadly winged; pappus awns 2, 1.5-3 mm. long.
340 FIELDIANA: BOTANY, VOLUME 24
Verbesina hypargyrea Robins. & Greenm. Proc. Amer. Acad.
34: 556. 1899.
Rocky cliffs and slopes, mixed or broadleaf forest, 800-1,800 m.,
Huehuetenango. Mexico (Chiapas).
Shrubs or trees to about 5 m. tall, the stems not winged, the branches cinerous-
puberulent with appressed hairs or glabrate; leaves alternate, short-petiolate or
subsessile, the blades elliptic, lanceolate, or lance-oblong, mostly 6-20 cm. long, 1.5-4
cm. wide, acuminate, attenuate to the base, the margins serrate or subentire,
conspicuously bicolored, the upper surface dark green, essentially glabrous, the lower
surface densley white-strigillose, pinnately veined; inflorescences corymbiform; heads
numerous, subglobose, about 8 mm. high, on cinerous-puberulent pedicels; phyllaries
2-3-seriate, the outer ones longest; pales mucronulate; ray flowers 7-10, with pale
yellow ligules less than 5 mm. long; disc flowers yellow, numerous; achenes about 2
mm. long, glabrous, narrowly winged, one wing slightly broader than the other;
pappus awns 2, subequal, 1-1.5 mm. long.
Verbesina hypoglauca Sch. Bip. ex Klatt, Leopoldina 23: 144.
1887.
Damp or wet thickets, dense, mixed forest, and coniferous
forest, 1,200-3,600 m.; Chimaltenango; Huehuetenango; Quezal-
tenango; San Marcos; Solola; Totonicapan. Southern Mexico.
Shrubs or small trees to 6 m. tall, usually densely branched, the young branches
white- to mentu lose; leaves opposite, short-petiolate, the blades lanceolate to oblong-
lanceolate, mostly 5-12 cm. long, acuminate, the base cuneate and then attenuate
and decurrent on the petiole but not to its base, the margins denticulate or serrulate,
the upper surfaces thinly strigillose or glabrate, the lower surfaces white or gray,
densely sericeous-strigose; inflorescences cymose-paniculate and sometimes appearing
corymbiform, broad, open; heads numerous, on pedicels mostly 1-3 cm. long;
involucres 5-8 mm. high; outermost phyllaries longest, linear to spathulate-obovate,
usually foliaceous, obtuse or broadly subtriangular or subacute, grayish strigose;
pales acute; ray flowers about 10, the ligules bright yellow, 10-15 mm. long; achenes
3-5 mm. long, black, almost glabrous or sparsely appressed-pubescent, very narrowly
winged (the margins sometimes appearing merely ciliolate); pappus awns 2-3 mm.
long, deciduous.
Verbesina hypsela Robins. Proc. Boston Soc. Nat. Hist. 31:
269. 1904. Capitana (Guatemala); lengua de vaca (Chimaltenango);
vara de carrizo (Chiquimula).
Wet to dry thickets or forest, 200-2,800 m.; Chimaltenango;
Chiquimula; Escuintla; Guatemala; Jutiapa; Retalhuleu; Santa
Rosa; Suchitepequez. Mexico (Chiapas).
Erect, coarse herbs to 3 m. high, simple below the inflorescence, the stems thick,
often reddish or purplish, glabrous, the internodes broadly winged by the decurrent
leaf bases; leaves alternate, on broadly winged petioles, the blades mostly 10-30 cm.
long, deeply and irregularly pinnatifid, the lateral lobes 3-6 pairs, acute or acuminate,
the margins essentially entire or inconspicuously denticulate, the upper surface thinly
NASH: FLORA OF GUATEMALA 341
pilose or almost glabrous, the lower one densely sericeous- pilose or in age glabrate,
the blades usually contracted at the base, sometimes broadly cuneate and then rather
abruptly decurrent to form the winged petiole, this broadly auriculate-clasping at the
base; panicles flat-topped or rounded, often as much as 30 cm. broad; heads
numerous, pedicellate; involucres rather narrowly campanulate, 4-5 mm. high;
outermost phyllaries short, lanceolate or oblong-ovate, the others linear-oblong to
linear-oblanceolate, acute or subacute, appressed-pilosulous; ray flowers about 5, the
ligules white, inconspicuous, about 2 mm. long; disc flowers about 20; achenes 4-5
mm. long, attenuate to the base, strigillose and somewhat tuberculate near the apex,
winged, pappus awns 2, about 2 mm. long.
Verbesina lanata Robins. & Greenm. Proc. Amer. Acad. 34:
558. 1899. Hoj; taxanx; xou (Quecchi, Alta Verapaz).
Wet forest, sometimes along streams in ravines, occasionally in
Liquidambar forest, 1,250-2,500 m.; Alta Verapaz (type from Coban,
Tuerckheim 1344); Huehuetenango; El Quiche; Zacapa. British
Honduras; Nicaragua; Costa Rica.
Large shrubs or trees to 10 m. high, the trunk sometimes as much as 20 cm. in
diameter, the branches stout, not winged, densely pilose with long, brownish, more or
less matted hairs; leaves very large, on short, naked petioles, the blades thin, elliptic
to oblong-ovate, mostly 20-60 cm. long and 10-30 cm. wide, acute to short-acuminate,
cuneate or attenuate to the base and more or less decurrent on the petiole, the
margins obsoletely denticulate or almost entire, sparsely pilose above or almost
glabrous, usually thinly short-pilose beneath or in age sometimes glabrate;
inflorescences large, the panicles sometimes as much as 50 cm. broad; heads very
numerous, pedicellate; involucres 6-7 mm. high, turbinate-campanulate, sordid-
tomentulose; phyllaries about 4-seriate, oblong, obtuse; ray flowers about 10, the
ligules white, about 6 mm. long; disc flowers yellowish; achenes 5 mm. long, narrowly
attenuate to the base, sparsely appressed-pilosulous; pappus awns white, about 3 mm.
long.
Handsome plants, the large inflorescences much used in the
Coban area of Alta Verapaz for decorating altars during the
Christmas season.
Verbesina minarum Standl. & Steyerm. Field Mus. Bot. 23:
263. 1947.
Damp mountain forest, 2,000-3,000 m.; El Progreso; Zacapa
(type from Rio Repollal, upper slopes to summit of mountain,
Sierra de las Minas, Steyermark 42489). Mexico (Chiapas).
Shrubs or small trees to 4.5 m. high, apparently densely branched, the branches
stout, subterete, the older ones glabrate and brownish, the young ones densely
strigillose with pale hairs, the internodes usually very short; leaves alternate, on
petioles 0.4-13 mm. long, the blades oblanceolate or oblong-oblanceolate, 4.5-11 cm.
long, 1-4 cm. wide, acute or subacute, gradually long-attenuate to the base, the
margins entire or with a few serrations near the apex, glabrous, pale beneath, when
young sometimes minutely and inconspicuously strigillose, the lateral nerves 7-8
342 FIELDIANA: BOTANY, VOLUME 24
pairs; inflorescences terminal, corymbose, mostly 3-7 cm. broad, shorter than the
leaves; heads numerous, radiate, pedicellate or subsessile; involucres campanulate, 4-
4.5 mm. high; phyllaries greenish, imbricate, appressed, the outer ones oblong-ovate,
the inner ones oblong, all subacute, sparsely and minutely puberulent; pales linear-
oblong, about 4 mm. long, pale, membranaceous, obtuse or subacute; ray flowers 6-8,
the ligules white, about 2 mm. long; disc corollas white, about 3.5 mm. long,
puberulent near the base; immature achenes compressed, glabrous, obscurely ciliate;
pappus awns 1-2 mm. long.
Although the heads were described by Standley and Steyer-
mark as discoid, the ray flowers on the immature type specimen are
easily seen.
Verbesina neriifolia Hemsl. Biol. Cent. Am. Bot. 2: 188. 1882.
Not reported from Guatemala but may be expected, as several
collections have been made in Chiapas, Mexico, 1,300-1,400 m.
Shrubs, 1-2 m. high, the branches narrowly or broadly winged; leaves alternate,
subsessile or short-petiolate, the blades narrowly la nee- elliptic to linear-lanceolate or
lance-oblong, mostly 6-12 (-15) cm. long, 1-2 cm. wide, acuminate, attenuate to the
base and decurrent on the petiole, glabrous above or minutely and inconspicuously
puberulent, silvery-tomentulose beneath with minute, appressed hairs, the margins
subentire or serrate; inflorescences cymose, corymbiform; heads few to several,
radiate, the peduncles and pedicels often winged; involucres 5-8 mm. high; phyllaries
3-4-seriate, the outermost ones longest, linear-lanceolate, sericeous-tomentose; pales
acute; ray flowers 18-20, the ligules yellow, 7-10 (-15) mm. long; immature achenes
puberulent, about 3 mm. long, pappus awns 2, about 2 mm. long.
Verbesina persicifolia DC. Prodr. 5: 614. 1836.
Not reported from Guatemala but to be expected, as it has
been collected in nearby Chiapas, Mexico. Mexico.
Shrubs to about 4 m. tall, the stems striate, minutely puberulent or glabrate;
leaves alternate, short-petiolate, rather narrowly lanceolate, mostly 15-25 cm. long, 3-
5.5 cm. wide, long-acuminate, attenuate to the base, the margins serrate, essentially
glabrous on both surfaces; inflorescences corymbose; heads radiate, usually
numerous, the pedicels appressed-puberulent; involucres soon saucer-shaped, in
anthesis 6-10 mm. broad; phyllaries 3-seriate, oblong, rounded to subacute, 3-4 (-5)
mm. long, ciliolate; pales linear, glabrous or nearly so, triangular to subacute at the
apex; ray flowers 12-21, the ligules yellow, 4-5 mm. long; disc flowers numerous; ray
achenes glabrous, winged; ray pappus of 1 or 2 awns; disc pappus of 2 short awns.
Verbesina perymenioides Sch. Bip. ex Klatt, Leopoldina 23:
143. 1887. Otopappus perymenioides Klatt, Ann. Naturh. Hofmus.
Wien 9: 362. 1894.
Damp forest or thickets, 1,100-1,800 m.; Guatemala. Mexico.
Shrubs or small trees to 6 m. tall, the branches not winged, puberulent to
somewhat tomentose or glabrate; leaves alternate, short-petiolate, the blades
NASH: FLORA OF GUATEMALA 343
lanceolate to lance-oblong or elliptic, mostly 8-16 cm. long, acuminate, cuneately
decurrent on the petiole, the margins serrate, often scabrous above, the lower surface
soft-pilose or tomentose or glabrous; heads radiate, numerous, small, in fruit only 5-7
mm. broad, disposed in rather broad corymbs, the pedicels usually densely short-
tomentose; phyllaries about 3-serieate, rounded or obtuse, at least the outer ones
pubescent and ciliolate; pales obtuse; ray flowers about 8, the ligules yellow, 2-4 mm.
long, often bifid at the apex; achenes 2-3 mm. long, broadly winged or merely ciliate,
or the wings very narrow, the surface papillate or not; pappus awns 2, subequal, 1-2
mm. long.
Verbesina petzalensis Standl. & Steyerm. Field Mus. Bot. 23:
147. 1944. Damp hillside thickets, sometimes in cornfields, 900-1,800
m.; Huehuetenango (type from near Rio San Juan Ixtan, east of
San Rafael Petzal, Standley 82921). Mexico (Chiapas).
Shrubs, 2-3.5 m. high, the branches thick, striate, densely glandular and sordid-
pilosulous or tomentulose, winged by the decurrent petiole bases; leaves alternate,
sessile or on broadly winged petioles, the blades membranaceous, oblong-ovate or
lance-oblong, the upper ones as much as 40 cm. long and 11 cm. wide, the lower ones
often 50 cm. long and 30 cm. wide, acute or acuminate, cuneately narrowed or
attenuate to the base of the petiole, obsoletely denticulate or almost entire, very
densely scabrous above, densely pilose beneath with short, sordid, spreading or
subappressed hairs; inflorescences large, corymbiform, the pedicels 0.5-1 cm. long,
rigid, densely villosulous-hispidulous; heads numerous, radiate; involucres camp-
anulate, 4-4.5 mm. high; phyllaries about 3-seriate, pale, linear-lanceolate to linear-
spathulate or oblong-obovate, obtuse or rounded and subcuspidate at the apex,
pilosulous; ray flowers about 8, the ligules white, broadly oblong, about 3 mm. long;
achenes about 3 mm. long, black, glabrous, sparsely and minutely tuberculate, with
rather narrow, white wings; pappus awns 2, about 2 mm. lone.
Verbesina pleistocephala (Donn.-Sm.) Robins. Proc. Amer.
Acad. 43: 41. 1907. Encelia pleistocephala Donn.-Sm. Bot. Gaz. 13:
189. 1888. V. donnell-smithii Coulter, Bot. Gaz. 20: 50. 1895 (type
from El Quiche, Heyde & Lux 3385).
Damp or wet forest and thickets, 800-2,000 m.; Alta Verapaz
(type from Coban, Tuerckheim 1121); Baja Verapaz; Huehue-
tenango; El Quiche. Mexico.
Shrubs to 2.5 m. high, the branches angulate or stnate, often dark red or purple,
hispidulous when young; leaves alternate, short-petiolate, the blades oblong-
lanceolate, mostly 10-17 cm. long, acute or acuminate, gradually or abruptly cuneate
at the base and decurrent on the petiole but not usually to its base, the margins
serrate, rarely entire, scabrous on both surfaces but more conspicuously so on the
upper surface; heads numerous, pedicellate, radiate, disposed in large or small,
usually dense, corymbiform panicles; involucres 5-6 mm. long, turbinate-campanulate
in anthesis, then campanulate; phyllaries 2-3-seriate, puberulent, purplish green, the
outer ones shorter, obtuse, the inner ones oblong, acute; ray flowers commonly 5-6,
the ligules yellow, spreading, conspicuous, 3-5 mm. long; achenes black, glabrous, 4-6
mm. long, broadly winged when mature; pappus awns commonly 3-4 mm. long.
344 FIELDIANA: BOTANY, VOLUME 24
Verbesina punctata Robins. & Greenm. Proc. Amer. Acad. 34:
561. 1899. Chimaliote (Jutiapa); lengua de vaca (Chimaltenango);
vara blanca (Jalapa); yucar (Santa Rosa).
Damp or wet thickets or mixed forest, sometimes in oak forest,
600-1,800 m.; Chimaltenango; Huehuetenango; Jalapa; Jutiapa;
Santa Rosa (type from Casillas, Heyde & Lux 4241}. Mexico; El
Salvador.
Coarse herbs, 2-4 m. high, simple or branched, the stout stems pale, puberulent
or almost glabrous, conspicuously and broadly winged by the decurrent leaf bases;
leaves alternate, the blades thin, ovate to lance-ovate or oblanceolate, mostly 18-40
cm. long, 7-20 cm. wide, acute or acuminate, rounded or broadly cuneate at the base
and narrowing rather abruptly into the winged petiole, the margins serrate-dentate,
the upper surface slightly scabrous and white-punctate, glabrous or glabrate beneath;
inflorescences large, more or less corymbose, the peduncles winged; heads very
numerous, on pubescent pedicels, radiate; involucres campanulate, 4-6 mm. high;
phyllaries 3-seriate, lanceolate or oblong-lanceolate, acute or acuminate, more or less
cuspidate, puberulent, ciliate; ray flowers about 8, the ligules white, oval-oblong, 3-4
mm. long; disc flowers greenish white; achenes more or less puberulent, 3-4 mm. long,
narrowly winged; pappus awns 2, about 2 mm. long.
This species has been reported from Guatemala as V. leprosa
Klatt.
Verbesina scabriuscula Blake, Contr. Gray Herb. 52: 54.
1917. Suquinai bianco (Suchitepequez); toquillo (Sacatepequez);
s-suq sa'an, saqi mank (Quecchi, Alta Verapaz).
Damp thickets and in oak, pine, or mixed forest, on cliffs and
rocky banks, in ravines, sometimes along roadsides, 1,300-2,600 m.;
Chimaltenango; Guatemala; Jalapa; Quezaltenango; Sacatepequez;
Solola; Suchitepequez.
Large, coarse, herbaceous or shrubby plants, 2-5 m. high, simple or sparsely
branched, the branches thick, not winged, densely sordid-pilosulous or tomentulose;
leaves alternate, petiolate, the blades ovate to broadly ovate, mostly 25-50 cm. long,
acuminate, cuneate and abruptly contracted at the base and decurrent on the petiole
(often but not always to the base of the petiole), the margins irregularly repand-
dentate to sinuate-dentate, densely scabrous above, densely hispidulous or scabe-
rulous beneath; inflorescences large, the corymbiform panicles often more than 30
cm. broad, the pedicels mostly 5-12 mm. long; heads radiate; involucres 3-4 mm. high;
phyllaries 2-3-seriate, linear to somewhat spathulate, acute or acuminate, mucronu-
late, pale, pubescent or pilosulous; ray flowers about 8, the ligules white, 3-5 mm.
long; disc flowers greenish white; achenes about 3 mm. long, tuberculate-hispidulous,
winged; pappus awns 2 mm. long, about equal.
Handsome, highly ornamental plants, the large panicles often
sold in markets to be used for decoration.
NASH: FLORA OF GUATEMALA 345
Verbesina sousae Fay, Brittonia 25: 195. 1973.
Not reported from Guatemala but may be expected, as it occurs
in the mountains of Chiapas, Mexico.
Shrubs, about 3 m. high, the branches thick, densely puberulent; leaves
alternate, petiolate, the blades mostly rhombic-ovate, the uppermost ones more
narrow, 10-20 cm. long, 3-11 cm. wide, acuminate, the base cuneate, the margins
sinuate-serrate, almost glabrous above, the indument mostly of scattered, tuberculate
hairs, but puberulent on costae and veins, densely and softly hirtellous beneath;
inflorescences corymbose, the stems and peduncles ascending; heads radiate;
involucres 0.5-1 cm. high, 2-2.5 cm. wide; phyllaries about 3-seriate, oblong, indurate,
puberulent, especially near the base, the outermost ones often with herbaceous
apices, rounded, sometimes apiculate, the inner ones acute; ray flowers 30-50, fertile,
the ligules yellow, about 1 cm. long; disc flowers commonly 200-300; pales rigid, 6-7
mm. long, narrowly oblanceolate, acute, carinate; disc achenes black, 3-4 mm. long,
compressed, very narrowly winged on one side; pappus of only one awn; ray achenes
similar but more or less trigonous.
Verbesina standleyi (Steyerm.) D. Nash, Phytologia 31: 362.
1975. Calea standleyi Steyermark, Field Mus. Hot. 22: 299. 1940.
Damp or wet, mixed forest, 2,000-3,000 m.; San Marcos (type
from slopes of Volcan de Tajumulco, between Todos Santos and
Finca El Porvenir, Steyermark 37004); also on Volcan de Tacana.
Shrubs or small trees, 3-5 m. high, the branches glabrous; leaves on petioles 1.5-5
cm. long, the blades oblong-elliptic, mostly 7-18 cm. long, 3-7.5 cm. wide, long-
acuminate, acute or attenuate to the base, the margins serrulate, triplinerved far
above the base, glabrous above, minutely appressed-pilose beneath along the veins or
glabrous; inflorescences corymbose-paniculate; heads few or numerous, radiate, the
pedicels 2-6 cm. long, pilose with mostly spreading hairs; involucres 4-5 mm. high;
phyllaries 2-3-seriate, the outer ones herbaceous, puberulent, oblong-lanceolate or
liguliform, obtuse, 3-5 mm. long, the inner ones oblong or ovate, obtuse or rounded at
the apex, shorter; ray flowers 8-10, pistillate, the ligules white, 1.5-2 cm. long; disc
flowers about 100, the corollas yellow, about 3 mm. long, glandular-puberulen.t;
achenes 2-3 mm. long, black, hirtellous at the apex, obovate or turbinate, 2-3-
angulate, ciliolate on the angles; ray pappus of 1-3 awns or none; disc pappus of 2
fimbriate awns, about one third as long as the body of the achene.
Verbesina steyermarkii Standl. Field Mus. Bot. 22: 320. 1940.
Julgunero (Guatemala, fide Morales); malacate (Huehuetenango);
tatascame (Solola).
Damp or dry thickets and deciduous forest, frequently in oak
forest, 1,000-2,100 m.; Chiquimula (type collected between Chiquim-
ula and La Laguna, Steyermark 30698); Guatemala; Huehue-
tenango; Jalapa; Solola. Mexico (Chiapas).
Shrubs or small trees to 6 m. high, the branches yellowish to pale brown,
sericeous-pilose at first but soon glabrate, not winged; leaves alternate, short-
346 FIELDIANA: BOTANY, VOLUME 24
petiolate, the blades thin, narrowly lanceolate to oblanceolate, 6-15 (-25) cm. long, 2-4
(-9) cm. wide, acuminate to narrowly long-acuminate, narrowly attenuate to the base
and decurrent on the petiole, the margins usually crenate-serrate, sometimes coarsely
serrate, the upper surfaces at first sparsely pilose and scabrous, but usually glabrous
in age, pilose beneath or sparsely and laxly pilose when young and in age essentially
glabrous; inflorescences corymbiform-paniculate, 6-16 cm. broad; heads numerous,
crowded, radiate, on slender pedicels to about 16 mm. long, the heads 6-8 mm. high
and about 4 mm. broad in anthesis; phyllaries about 3-seriate, pale, ovate-oblong to
oblong, ciliate, glabrous or glabrate, usually obtuse or rounded at the apex but
sometimes broadly triangular and subacute; pales acute; ray flowers about 8, the
ligules yellow, 2-3 mm. long; achenes black, glabrous, about 3.5 mm. long, the outer
ones broadly winged; pappus awns about 2 mm. long.
Verbesina sublobata Benth. PL Hartweg. 76. 1841. Toquillo
(Sacatepequez).
Damp or dry thickets or forest, often in pine-oak forest,
sometimes in open fields, 100-2,600 (rarely to 3,700) m.; Alta
Verapaz; Chimaltenango; Chiquimula; Guatemala; Huehue-
tenango; Quezaltenango; Sacatepequez; San Marcos; Santa Rosa;
Solola. Mexico (Chiapas); Honduras; Costa Rica.
Erect, coarse herbs or shrubs, 1.5-3 m. high, simple or branched, the stems thick
not winged, densely sordid-tomentose; leaves alternate, on broadly winged petioles,
the blades rather thick, ovate in outline, sinuate-lobate to deeply lobate-pinnatifid,
mostly 10-25 cm. long, acute to acuminate, usually almost truncate at the base and
then abruptly decurrent on the petiole, the margins denticulate, dentate, or undulate,
the upper surface very scabrous, the lower one densely pilose-tomentose or
hispidulous, the venation usually conspicuous; inflorescences large, often as much as
30 cm. broad, the panicles rounded or flat-topped; heads numerous, radiate, on
slender pedicels; involucres about 3 mm. high; phyllaries linear to oblanceolate, acute
or acuminate, ciliate, pubescent or glabrate; ray flowers 8-10, the ligules white, about
2 mm. long; achenes about 3 mm. long, usually sparsely tuberculate, sometimes
glabrous, broadly winged; pappus awns 2, 1.5-2.5 mm. long.
Variable, coarse, unsightly, weedy plants.
Verbesina turbacensis HBK. Nov. Gen. & Sp. 4: 203. 1820. V.
nicaraguensis Benth. in Oerst. Vid. Medd. Kjoeb. 1852: 97. 1852.
Mango, pitaneja (Alta Verapaz); obes-te, varilla blanca (Huehue-
tenango).
Damp or wet thickets or in thin forest, especially of pine and
oak, sometimes in Liquidambar forest, often on steep banks or
hillsides, near sea level to 1,800 m.; Alta Verapaz; Chiquimula;
Guatemala; Huehuetenango; Izabal; Jalapa; Jutiapa; Quezal-
tenango; Retalhuleu. Southern Mexico; British Honduras,
Honduras and El Salvador to Costa Rica; Colombia and Venezuela.
NASH: FLORA OF GUATEMALA 347
Herbaceous or shrubby plants or sometimes weak trees, commonly 1-5 m. high,
the branches stout, densely puberulent or tomentulose, winged by the decurrent
petiole bases; leaves alternate, on broadly winged petioles, broadly angulate-ovate to
to almost rhombic in outline, shallowly or deeply pinnate-lobate or rarely, merely
undulate or dentate, mostly 12-35 cm. long, the lobes obtuse or acute, very scabrous
above, usually densely grayish puberulent to tomentulose or velutinous beneath;
panicles rounded or flat-topped, mostly 15-30 cm. broad; heads numerous, radiate,
pedicellate; involucres 2-3 mm. high, campanulate; phyllaries pale, linear to obovate-
oblong, puberulent or pilosulous, ciliate, the outer ones obtuse or subacute, the inner
ones acute, the innermost ones acuminate; ray flowers 8-10, the ligules white, 2-4 mm.
long; achenes 2-2.5 mm. long, usually more or less strigillose, sometimes tuberculate,
winged, pappus awns 2, 1-1.5 mm. long.
This species has been reported from Guatemala as V.
diversifolia DC.
VIGUIERA HBK.
Reference'. S. F. Blake, A revision of the genus Viguiera, Contr.
Gray Herb. 54: 1-205. 1918.
Annual or perennial herbs or sometimes shrubs, usually pubescent, the stems
terete; leaves opposite or alternate, the lowest ones usually opposite, petiolate or
sessile, the blades ovate or rhombic-ovate to narrowly lanceolate or linear, rarely
lobate, the margins entire, dentate, or crenate; inflorescences cymose or of solitary,
pedicellate heads; heads heterogamous, radiate; involucres campanulate to subcylin-
dric; phyllaries graduate or subequal, 2-5-seriate, mostly linear-lanceolate to lance-
ovate, the base usually indurate but the apex herbaceous, sometimes entirely
herbaceous; receptacles flat to low-conic; pales firm, scarious or sometimes
subherbaceous at the apex, carinate, embracing the achenes and persistent after their
fall; ray flowers 5-30, neutral, the ligules (in ours) yellow, 2-3-denticulate or
subentire; disc flowers fertile, the corollas commonly yellow, the tube short, usually
pubescent, the limb 5-dentate; anthers sagittate at the base, the apical appendages
ovate; style branches rather long, more or less recurved, hispid above, with subacute
to acuminate appendages; achenes somewhat thickened, often subquadrangular,
usually appressed-pubescent, sometimes glabrous, not marginate, truncate or rounded
at the apex; pappus of the sterile ray achenes of 2-3 awns or squamellae, or none,
that of the disc achenes of 2 lacerate or fimbriate awns and several shorter, free or
united squamellae, or sometimes wanting.
Perhaps 150 species, all American, in tropical or warm-
temperate regions, with three in Guatemala; a fourth, V. longi folia
(Robins. & Greenm.) Blake, is also treated here as it has been
collected in nearby Chiapas, Mexico.
Leaves linear, linear-lanceolate, or linear-elliptic, pinnately nerved V. longifolia.
Leaves ovate to rhombic-ovate or lanceolate, triplinerved.
Leaf blades mostly 3-12 cm. long; ligules of ray flowers 7-25 mm. long.
Heads 7-10 mm. high V. dentata.
Heads 13-18 mm. high V. mima.
Leaf blades mostly 1.5-3.5 cm. long; ligules of ray flowers 2-3 mm. long.
V. graciUima.
348 FIELDIANA: BOTANY, VOLUME 24
Viguiera dentata (Cav.) Spreng. Syst. 3: 615. 1826. Helianthus
dentatus Cav. Icon. PL 3: 10, t. 220. 1795. V. helianthoides HBK.
Nov. Gen. & Sp. 4: 226, t. 379. 1820. V. dentata var. helianthoides
Blake, Contr. Gray Herb. 54: 86. 1918. Figure 111.
Open forest, brushy hillsides, or marshy, open ground, 300-1,800
m.; Baja Verapaz; Huehuetenango; Peten; El Progreso; Zacapa.
Southwestern United States; Mexico; British Honduras; Honduras.
Erect herbs to about 2 m. high, the stems slender, sparsely strigillose or glabrate;
lower leaves opposite, the upper alternate, petiolate, the blades ovate to rhombic-
ovate or lanceolate, mostly 3-12 cm. long, acute to long-acuminate, cuneate or
attenuate to the petiole, the margins serrate, serrulate, or subentire, strigillose and
usually scabrous above, beneath strigillose to densely soft-pilose; inflorescences
pedunculate; heads on pedicels mostly 2-8 cm. long, disposed in lax, open, cymose
panicles; disc of the head 7-10 mm. long, 10-14 mm. broad; involucres broadly
campanulate, 5-10 mm. high; phyllaries 3-seriate, lance-ovate or ovate-oblong to
linear or linear-oblong, acute or acuminate, herbaceous, spreading or appressed,
hispidulous or densely appressed-pilose; ray flowers 10-12, the ligules yellow, 7-15
mm. long; disc corollas yellow, more or less hirtellous, 3-4 mm. long; achenes
obovate-oblong, black or mottled, appressed-pubescent, 3-4 mm. long; pappus a
crown of fimbriate squamellae less than 1 mm. long and 2 slender, unequal awns, 2.2-
2.8 mm. long.
These weedy plants are reported to supply forage for horses;
the flowers are said to yield a superior grade of honey.
Viguiera gracillima Brandg. Univ. Calif. Publ. Bot. 6: 74.
1914.
A weed of cornfields and roadsides, about 300 m., Zacapa.
Mexico (Oaxaca).
Erect, much-branched annuals, the stems slender, more or less strigose; leaves
short -petiolate, the upper ones alternate, the lower ones opposite, the blades rhombic-
ovate to lance-ovate, 1.5-3.5 cm. long, acuminate or acute, broadly cuneate at the
base, the margins serrate, more or less hispid-strigose above, usually densely strigose
beneath; heads numerous, on strigose pedicels mostly 2-4 cm. long, the disc 7-8 mm.
high, 5-8 mm. wide; involucres broadly campanulate; phyllaries biseriate, lanceolate,
acuminate, canescent-strigose, herbaceous, appressed; ray flowers about 5, the ligules
yellow, 2-3 mm. long; disc corollas yellow, 3-4 mm. long; achenes about 3 mm. long,
black or mottled, appressed-pilose; pappus a crown of lacerate squamellae about 1.5
mm. long and 2 awns 3-4 mm. long.
Viguiera longi folia (Robins. & Greenm.) Blake, Contr. Gray
Herb. 54: 111. 1918. Gymnolomia longifolia Robins. & Greenm.,
Proc. Boston Soc. Nat. Hist. 29: 92. 1899.
Not reported from Guatemala, but may be expected there as it
has been collected in nearby Chiapas, Mexico. Arizona; New
Mexico; Texas; Mexico.
NASH: FLORA OF GUATEMALA 349
Erect herbs to about 1 m. high, sparsely branched, the stems strigillose or almost
glabrate in age; leaves opposite below, often alternate above, subsessile or short-
petiolate, the blades linear to narrowly lanceolate or linear -elliptic, mostly 4-12 cm.
long, 0.3-1.4 cm. wide, acuminate, attenuate to the base, pinnately nerved, the
margins entire or obscurely dentate, somewhat revolute, the upper surface strigillose
and scabrous, the lower surface more or less strigillose and glandular-punctate;
involucres broadly campanulate, 4-7 mm. high; phyllaries biseriate, herbaceous,
lanceolate or linear-lanceolate, hispid below, strigillose above; pales about 4 mm.
long, more or less hispid above, short- cuspidate; ray flowers about 12, the ligules
yellow, 6-12 mm. long; disc corollas about 3 mm. long; achenes black, shining,
glabrous, about 2 mm. long; pappus none.
Viguiera mima Blake, Contr. Gray Herb. 54: 69. 1918.
Known only from the type, Retalhuleu, Bernoulli & Carlo
1520.
Branching herbs, the stems purplish, pilose; leaves alternate, on petioles 6-13
mm. long, the blades ovate-lanceolate, acuminate, subtruncate to rounded or cuneate
at the base, 4-9.5 cm. long, 1.5-3 cm. wide, the margins crenate-serrate, 3-nerved,
softly a ppressed- pilose and glandular-punctate on the upper surface, paler beneath,
densely pilose with short, spreading hairs and glandular -punctate; heads few, long-
pedunculate; disc 13 mm. high or in fruit 18 mm. high, 20-28 mm. broad; involucre
biseriate; phyllaries subequal, narrowly oblong, about 1 cm. long, acute or obtuse,
pilosulous, appressed or lax at the apex; ray flowers about 16, the ligules oblong, 1.5-
2.5 cm. long; disc corollas 6.5 mm. long; achenes black, rather sparsely pubescent,
about 4 mm. long; pappus awns 2, ciliolate, 3 mm. long, the squamellae fimbriate,
about 1 mm. long.
I have seen only a fragment of the type specimen and have
followed Blake's description.
WEDELIA Jacquin
Herbs or shrubs, scabrous-pubescent or hirsute; leaves opposite, petiolate or
sessile, the margins commonly dentate or serrate, sometimes lobate, rarely entire;
heads heterogamous, radiate, pedunculate at the ends of branches or in the upper leaf
axils; involucres campanulate to hemispheric; phyllaries 2-4-seriate, the outer ones
usually herbaceous or foliaceous, the inner ones dry or membranaceous; receptacle
flat or convex; pales complicate or concave, enfolding the disc flowers; ray flowers
pistillate, fertile or sometimes sterile, the ligules spreading, entire or 2-3-dentate,
yellow; disc flowers hermaphrodite, fertile or the innermost sterile, the corollas
regular, tubular, the limb cylindric or narrowly campanulate, 5-dentate; anthers
entire at the base or sagittate; style branches short or elongated, the appendages
subacute; achenes glabrous or pubescent, cuneate-oblong or obovate, more or less
compressed laterally or those of the ray flowers triquetrous, the angles obtuse or with
cartilaginous wings, rounded at the apex and constricted into a short neck bearing
the cyathiform pappus, with or without one or two setiform, erect, often caducous
awns.
Perhaps 50 species, many of them weedy, in tropical and
subtropical regions of both hemispheres, with eight in Guatemala.
350 FIELDIANA: BOTANY, VOLUME 24
The genus has not been monographed recently and the taxonomy is
in a rather chaotic state. Treatment of the genus here is
unsatisfactory, but perhaps not much better can be prepared until
the group is monographed.
Plants usually prostrate, procumbent, or decumbent, the stems often rooting at the
lower nodes; leaves (at least the larger ones) trilobate or more or less hastate;
achenes not winged, commonly lacking pappus awns (rarely one very short awn
present).
Stems thinly strigose or glabrate; leaves essentially sessile, the blades cuneate at
the base or tapering into a broad, petioliform base, the margins rather
remotely dentate, denticulate, or crenate; achenes about 5 mm. long,
conspicuously tuberculate W. trilobata.
Stems hirsute with spreading, white hairs; leaves petiolate, the blades usually
abruptly contracted at the broad base and then cuneate-decurrent on the
petiole, the margins closely serrate; achenes about 3 mm. long, not tuberculate.
W. filipes.
Plants usually erect or ascending; leaves neither trilobate nor hastate (in ours);
achenes winged or not, commonly with 1 or 2 (3) awns, but these sometimes
caducous.
Ray achenes (when present) winged or not, with 2 awns, sometimes 3, one or more
of these sometimes caducous; disc achenes with 2 awns, these sometimes
caducous but always evident on immature material.
Leaves oval to oval-ovate, disc flowers about 12, pappus awns 1-1.5 mm. long,
unequal W. adherent.
Leaves ovate to narrowly lanceolate; disc flowers more than 12; pappus awns 2-4
mm. long, subequal or unequal.
Phyllaries mostly 8-16 mm. long.
Leaves mostly ovate; ray flowers sterile W. acapulcensis.
Leaves mostly lanceolate; ray flowers usually fertile.
W. hispida var. ramosissima.
Phyllaries mostly 5-8 mm. long W. pinetorum.
Ray achenes usually broadly winged, often without awns; disc achenes with 1 short
or long awn (rarely 2), sometimes none.
Disc flowers commonly 8-12 W. parviceps.
Disc flowers commonly 20-36 W. fertilis.
Wedelia acapulcensis HBK. Nov. Gen. & Sp. 4: 215. 1820;
McVaugh, Contr. Univ. Mich. Herb. 9: 459. 1972. W. strigosa Hook.
& Arn. Bot. Beechey Voy. 435. 1841. Aspilia strigosa Hemsl. Biol.
Cent. Am. Bot. 2: 171. 1881. Chinchiguaste (Zacapa); tatascamite
(Guatemala). Figure 112.
Damp thickets, brushy hillsides, scrub forest or cutover forest,
200-1,800 m.; Chiquimula; Guatemala; Zacapa. Mexico; Honduras;
El Salvador; Nicaragua; Costa Rica. Peru.
Erect, branching shrubs or suffrutescent perennials, 0.5-2 m. high, the stems
hispid or hispid-pilose; leaves short-petiolate, the blades ovate to broadly ovate, 2-6
NASH: FLORA OF GUATEMALA 351
cm. long, acuminate, rounded at the base and then more or less abruptly cuneate and
decurrent on the petiole, the margins dentate or serrate, more or less hispid-pilose on
both surfaces, often scabrous above; heads long-pedunculate, the involucres broadly
campanula te, 8-16 mm. high; phyllaries about 3-seriate, oblong, obtuse or subacute,
more or less hispid or hispidulous; ray flowers sterile, the ligules -yellow or orange;
disc flowers numerous, fertile, the corollas 5-7 mm. long; anthers 2-2.5 mm. long;
achenes 4-5 mm. long, narrowly to broadly winged; pappus a crown of 2 subequal or
unequal awns 2-4 mm. long, these sometimes caducous when mature.
Wedelia adhaerens Blake, Journ. Wash. Acad. Sci. 22: 383.
1932.
Known only from the type, in logwood (Haematoxylon] swamp,
Dos Arroyos, Peten, Bartlett 12111.
Herbs to about 50 cm. high, the stems densely hispidulous with short, mostly
spreading hairs, sparsely hispid, especially below; leaves opposite, subsessile to short -
petiolate, the blades oval or oval-ovate, the larger ones 3-3.5 cm. long and 1.5-2.5 cm.
wide, acute, broadly rounded at the base, appearing almost entire but remotely
callous-serrulate, spreading- hispidulous above, densely so beneath, triplinerved; heads
usually in clusters of 3 at the ends of the stems and branches, on slender peduncles to
1 cm. long, or sometimes these elongating in age; involucres campanulate, 7.5-9.5 mm.
high; phyllaries about 3-seriate, the outer ones lanceolate or lance-ovate, densely
hispidulous and tuberculate-hispid, acuminate or acute; disc about 4 mm. broad; ray
flowers about 6, the ligules yellow, mostly 5-9 mm. long; disc flowers about 12, the
corollas 5 mm. long, hispidulous on the teeth, elsewhere essentially glabrous; ray
achenes thick, obovate, with winglike margins, mottled, glabrous except at the apex, 4
mm. long; pappus a minute, lacerate cup and 2 awns 1.3 mm. long or the awns
caducous; pappus of the immature disc achenes a lacerate cup and 2 minute awns.
Wedelia fertilis McVaugh, Contr. Univ. Mich. Herb. 9(4): 462.
1972. Lipochaeta strigosa DC. Prodr. 5: 610. 1836, not W. strigosa
Hook. & Arn. (1841). Zexmenia strigosa Sch. Bip. in Seem. Bot.
Voy. Herald 306. 1856.
Wet thickets, pine-oak or Cupressus forest, 200-2,100 m.;
Chimaltenango; Huehuetenango; Jalapa; El Quiche; Sacatepequez;
Santa Rosa; Zacapa. Mexico; Honduras.
Herbaceous or suffrutescent perennials, 30-75 cm. high, the stems hispid; leaves
subsessile to short -petiolate, the blades mostly ovate, usually acuminate, sometimes
acute, mostly 3-6 cm. long, rounded at the base and usually rather abruptly
decurrent on the petiole, sometimes almost to the base, the margins serrate-dentate,
more or less hispid and scabrous above, hispid or hispid-pilose beneath; heads on
peduncles mostly 2-12 cm. long; involucres 5-7 mm. high; outer phyllaries hispid or
hispidulous; pales hyaline, the apices triangular, puberulent; ray flowers commonly
12 or more, fertile, the ligules yellow, 5-8 mm. long; disc flowers 20-36, the corollas
pale yellow, 4-5 mm. long; achenes 2.5-4 mm. long, more or less pubescent, the disc
achenes not winged, the ray achenes usually broadly winged; pappus a lacerate cup
of squamellae, with or without one or two short awns.
352 FIELDIANA: BOTANY, VOLUME 24
Wedelia filipes Hemsl. Biol. Cent. Am. Bot. 2: 170. 1881.
Damp or wet thickets or fields, sometimes in marshes or along
stream margins, 1,350-1,850 m.; Alta Verapaz; Baja Verapaz;
Chimaltenango; Guatemala; Sacatepequez (type from Duenas,
Salvin s.n.}. Costa Rica.
Decumbent or procumbent perennials, simple or sparsely branched, the stems
rather thick, hirsute with spreading, white hairs, often very leafy; leaves petiolate, the
blades rhombic-ovate to ovate-oblong, acuminate, at least the larger ones shallowly
trilobate or more or less hastate, mostly 4-8 cm. long, the margins closely and
unevenly serrate, triplinerved, densely hispid-pilose on both surfaces and scabrous
above, the hairs appressed or subappressed; heads few, solitary or 2 or 3 in a leaf axil,
the disc 8-9 mm. broad; phyllaries 3-seriate, oblong or ovate-oblong, obtuse or acute,
8-11 mm. long, strigose or hispidulous, the outer ones essentially herbaceous; pales
obtuse or acute; ray flowers 10-12, the ligules yellow, oblong-elliptic, about 1 cm.
long; disc flowers numerous, the corollas essentially glabrous but the lobes more or
less pubescent within; achenes about 3 mm. long, brown, cuneate-oblong, turgid,
shortly pubescent or glabrate; pappus short, cyathiform, fimbriate, without awns, or
rarely one short, caducous awn present.
Wedelia hispida HBK. Nov. Gen. & Sp. 4: 216. t. 371. 1820.
Zexmenia texana A. Gray, PL Wright. 1: 112. 1860. Z. hispida A.
Gray, Proc. Amer. Acad. 19: 10. 1883.
Erect to procumbent, branching herbs, the stems terete, striate, hispid; leaves
opposite, short-petiolate, the blades rather narrowly lanceolate to subovate-
lanceolate, acuminate, narrowing at the base and more or less decurrent on the
petiole, scabrous above, more or less hispid-pilose beneath, the margins irregularly
serrate-dentate; heads usually solitary on peduncles mostly 10-22 cm. long; phyllaries
lanceolate to ovate-lanceolate; pales linear- lanceolate, acute, carinate, glabrous,
scarious; ray flowers about 8, pistillate, usually fertile, the ligules yellow; disc flowers
numerous, the corollas yellow; ray achenes usually winged when mature; ray pappus
a lacerate cup, without awns; disc pappus a lacerate cup with usually 1 short awn.
Only the following variety is found in Guatemala.
Wedelia hispida var. ramosissima (Greenm.) K. Becker,
Phytologia 31: 25. 1975. Zexmenia hispida var. ramosissima
Greenm. Field Mus. Bot. 3: 127. 1903.
Forests and savannas, sea level to 200 m.; Alta Verapaz; Peten.
Mexico; British Honduras.
Differs from the typical variety in its usually subentire leaf
margins (sometimes remotely dentate), its inflorescences (some-
times of solitary heads but more often 2 or 3 in a loose cyme) on
peduncles mostly 3-9 cm. long, and in having usually 10-15 ray
flowers.
NASH: FLORA OF GUATEMALA 353
Wedelia parviceps Blake, Contr. U. S. Nat. Herb. 24: 28, t. 9.
1922.
Damp or wet thickets, often in pine forest, sometimes in
savannas, near sea level to 1,200 m.; Alta Verapaz; Baja Verapaz;
Guatemala; Huehuetenango; Izabal; Peten; Zacapa (type from
Gualan, Blake 7681). Mexico (Yucatan); British Honduras;
Honduras; Nicaragua; Costa Rica.
Erect, branching herbs to about 1 m. high, the stems slender, usually hispidulous,
glandular; leaves opposite, short-petiolate, the blades lance-ovate to narrowly ovate
o" lanceolate, mostly 2-6 cm. long, acute or acuminate, more or less cuneate at the
base, the margins serrulate, more or less hispid above and often scabrous, hispidulous
beneath, but hispid on the veins, triplinerved; heads pedunculate, solitary or in
groups of 3 at the ends of branches; involucres campanulate, 3.5-7 mm. high;
phyllaries about 3-seriate, hispid-pilose, ciliate, subacute, the tips herbaceous,
somewhat spreading; ray flowers 5-8, the ligules bright yellow, 4-8 mm. long; disc
flowers about 9, the corollas about 4 mm. long; achenes gray or blackish, more or less
appressed-pubescent, or glabrate, narrowly or broadly winged; pappus a minute,
lacerate cup, with or without a caducous awn on the inner side.
Wedelia pinetorum (Standl. & Steyerm.) Becker, Phytologia
31: 25. 1975. Zexmenia pinetorum Standl. & Steyerm. Field Mus.
Bot. 23: 149. 1944. Chinchiguaste (Huehuetenango); tatascamito
(Guatemala).
Wet, mixed forest and thickets, swampy areas, 200-2,100 m.;
Baja Verapaz; Chiquimula; Guatemala; Huehuetenango; Jalapa;
Jutiapa (type from hills between Jutiapa and Plan de Urrutia,
Standley 75590); El Quiche; Santa Rosa. Honduras.
Erect or ascending, branching herbs, commonly 50-100 cm. tall, the stems densely
or sparsely hispid; leaves short-petiolate or subsessile, the blades ovate to rhombic-
ovate or lanceolate, mostly 3-7 cm. long, acuminate or acute, broadly cuneate at the
base or almost truncate and then rather abruptly cuneate and decurrent on the
petiole, triplinerved, the margins serrate or subentire, rather densely hispid above and
scabrous, densely or sparsely hispid-pilose beneath; heads usually long-pedunculate,
1-3 at the ends of branches and in the upper leaf axils; involucres broadly
campanulate, 5-8 mm. high; phyllaries 3-seriate, triangular-oblong to linear-oblong,
obtuse to subacute, the outer ones hispid or hispidulous, the inner sparsely
hispidulous or glabrate; pales linear, acuminate, about 9 mm. long, ciliolate; ray
flowers 5-8, the ligules yellow, 4-8 mm. long; disc flowers 20 or more; achenes 4-5
mm. long, subcompressed, more or less appressed-pilose, winged; pappus a short,
lacerate cup and 2 unequal or subequal awns 2-4 mm. long.
Wedelia trilobata (L.) Hitchc. Rept. Mo. Bot. Gard. 4: 99.
1893. Silphium trilobatum L. Syst. ed. 10: 1233. 1759. W. carnosa L.
Rich, ex Pers. Syn. PI. 2: 490. 1807. Verbesina tridentata Spreng.
Syst. Veg. 3: 577. 1826. Figure 113.
354 FIELDIANA: BOTANY, VOLUME 24
Damp or wet soil, sometimes in marshes, often along sea
beaches, sea level to about 600 m.; Alta Verapaz; Huehuetenango;
Izabal; Peten. Florida; southern Mexico; British Honduras, along
the Atlantic coast to Panama; West Indies; northern South
America.
Prostrate or procumbent, perennial herbs, somewhat fleshy, sparsely branched,
the stems thinly strigose or glabrate, usually rooting at the lower nodes, to about 1
m. long; leaves essentially sessile, elliptic to obovate or obovate- oblong, mostly 3-12
cm. long, obtuse or acute, cuneate below, or narrowing into a broad, petioliform base,
this sometimes slightly dilated near the stem, all or at least the lower ones rather
shallowly trilobate or subhastate, the margins rather obscurely or coarsely dentate,
denticulate, or crenate, thinly strigose, especially on the upper surface, triplinerved;
peduncles solitary in the upper leaf axils, 2-14 cm. long; involucres 8-12 mm. high;
phyllaries oblong-obovate, herbaceous throughout or nearly so, obtuse, sparsely or
moderately pilose; pales acuminate or acute, more or less serrate to lacerate; ray
flowers about 10, the ligules bright yellow, 0.8-1.5 cm. long, usually tridentate at the
apex; disc flowers yellow, the corolla lobes densely pubescent within, glabrous
outside; achenes about 5 mm. long, usually conspicuously tuberculate in maturity,
glabrous or somewhat pubescent, at least on the margins; pappus short, cyathiform,
fimbriate, without awns.
ZEXMENIA La Llave
By KENNETH M. BECKER
Herbs, subshrubs, or weak shrubs, sometimes clambering; leaves all opposite or
becoming subopposite or alternate above, generally ovate or lance-ovate, sometimes
narrower, the margins serrate, occasionally revolute, triplinerved; heads radiate,
solitary and axillary and/or terminal or in umbelliform, cymose clusters; involucres
campanulate to hemispherical, 2-4-seriate, weakly to strongly graduate, phyllaries
generally ovate or oblong, the outer indurate below and foliaceous or at least
herbaceous above, the inner generally membranaceous below and often somewhat
herbaceous near apex; pales membranaceous, conduplicate, sometimes with a dark
line along keel; rays fertile, yellow; disc corollas campanulate, 5-lobate; anther
thecae generally auriculate, brown or black; style branches filiform, hispidulous
dorsally; disc achenes cuneate or broad-cuneate, body usually thickened, generally
constricted above and bearing at the apex a pappus of often connate squamellae and
one or two generally fragile awns, or awnless, margins often with true wings, wings
sometimes ciliate, faces often tuberculate and/or hispid; ray achenes 3-angled, 3-
awned or toothed, in most species otherwise similar to disc achenes.
About 20 species, ranging from the southwestern United States
to South America. Closely related to Wedelia Jacq. Generic limits
in the Zexmenia-Wedelia-Aspilia complex are in need of revision.
Outer phyllaries similar to leaves in size, shape and pubescence Z. phyllocephala.
Outer phyllaries not similar to leaves.
Leaf margins decurrent onto petiole and clasping stem Z. guatemalensis.
Leaf margins not decurrent onto petiole.
Outer phyllaries longer than inner.
BECKER: FLORA OF GUATEMALA 355
Outer phyllaries lance-oblong or linear, 1-2 mm. wide above Z. virgulta.
Outer phyllaries ovate-oblong, often spathulate, 5-8 mm. wide above.
Z. salvinii.
Outer phyllaries shorter than inner or subequal Z. serrata.
Zexmenia guatemalensis Donn.-Sm. Bot. Gaz. 13: 188. 1888
(type from Coban, Alta Verapaz, J. D. Smith 853).
Damp thickets, savannas, moist pine forest or oak forest, often
in clearings, 1,200-1,500 m.; Alta Verapaz; Peten. Mexico (Chiapas).
Coarse-hairy herbs to 2 m. high, often swollen at nodes, the stems several-angled,
coarsely hispid to pilose, especially above; leaves opposite below, alternate in
inflorescence, petioles to 1.5 cm. long, hispid, blades thin, often drying black or matte-
brown above, decurrent onto petiole and clasping stem, usually lance-ovate,
occasionally lanceolate or ovate, 4-16 cm. long, 1.5-6 cm. wide, acute or acuminate at
the apex, cuneate to rounded at the base, scabrous and hispid above, with longer
hairs along veins, white-hirsute to pilose, especially along the veins below, the
margins shallowly serrate and somewhat revolute to serrate, triplinerved; heads
solitary, terminal and from the usually condensed upper leaf axils, on peduncles to
5.5 cm. long, slender, white-hirsute to pilose; involucres campanulate; fruiting heads
hemispherical, about 1 cm. high, 1.5 cm. wide; phyllaries in 2 series, outer phyllaries 5,
longer than inner series, foliaceous and often ampliate above, somewhat indurate
below, 10-13 mm. long, 5-6 mm. wide, rounded and mucronate to acute at apex,
white-hirsute and puberulent dorsally, the inner phyllaries membranous, ovate-
oblong, acute to obtuse at apex; ray flowers typically 8, yellow, ligules 1-1.5 cm. long;
disc corollas yellow, the lobes hireutulous; anther thecae black; pales stramineous,
exposed in ripe heads and enveloping disc achenes; disc achenes 4 -angled, exaristate
or 1-awned, 3-4 mm. high, about 1 mm. wide, often tuberculate when mature, not
winged; disc pappus a squamulate cup to 0.5 mm. high; ray achenes triquetrous, 3-
awned or toothed, membranous wing-margined, including wings to 3 mm. wide; ray
pappus a squamulate cup to 2 mm. high.
Known from the area around Coban, Alta Verapaz, and
scattered localities in Peten and Chiapas, this species is rendered
quite distinctive vegetatively by its decurrent-clasping leaf margins.
Zexmenia phyllocephala (Hemsl.) Standl. & Steyerm. Field
Mus. Bot. 23 (3): 150. 1944. Wedelia phyllocephala Hemsl. Biol.
Cent. Am. Bot. 2: 170. 1881 (type from between San Cristobal and
Rio Chisoy, Alta Verapaz, Bernoulli 1077). Z. phyllostegia
Klatt, Leopoldina 25: 105. 1889. Z. subsericea Blake, Contr. U. S.
Nat. Herb. 22: 635. 1924 (type from Agua Blanca, Tejada 57).
Figure 114.
Dry, open-wooded, often rocky slopes, often on serpentine,
1,100-2,800 m.; Baja Verapaz; El Quiche; Huehuetenango. Endemic.
Densely branched, slender-stemmed subshrubs, 0.5-1 m. high, stems viscid and
white-hirsute to pilose; leaves opposite below, becoming subopposite to alternate
356 FIELDIANA: BOTANY, VOLUME 24
above, on petioles to 0.5 cm. long, the blades green above, grayish green or whitish
below, narrowly lanceolate to lance-elliptic, 2.5-7 cm. long, 0.5-1.5 cm. wide, acute at
apex, cuneate at base and decurrent onto petiole, white-hispid to hispid above,
canescent or subcanescent-pilose with long, slender hairs below, the margins
denticulate-mucronulate to subentire, usually revolute, triplinerved; heads solitary,
terminal, to 4 cm. wide across extended rays, on slender, hispid peduncles to 20 cm.
long; involucres hemispherical, 3-4-seriate, about 1 cm. high, 1.5-2 cm. wide; outer
phyllaries 4-5, foliaceous and similar to cauline leaves in shape and pubescence,
generally much surpassing inner phyllaries in length, 2-5 cm. long, 0.5-2 cm. wide,
somewhat indurate below, the inner phyllaries membranous below, sometimes with
herbaceous tips, ovate-oblong, margins ciliate; ray flowers to about 20, orange-yellow
or yellow, ligules to 1.2 cm. long; disc corollas yellow; anther thecae dark brown;
pales stramineous, enveloping and surpassing the achenes; disc achenes 2-awned,
thickened, the body 3.5-4 mm. high, 2-3 mm. wide, the awns 3-4 mm. long, margins
usually winged, the wings ciliate, the faces usually tuberculate and hispid, glabrate
toward margins; disc pappus squamellae connate into a cuplike structure to about 1
mm. long; ray achenes triquetrous, 3-toothed or awned, otherwise similar to disc
achenes.
This montane endemic is rendered quite distinctive vegetatively
by its foliaceous outer phyllaries and canescent or subcanescent leaf
undersurfaces.
Zexmenia salvinii Hemsl. Biol. Cent. Am. Bot. 2: 173. 1881
(type from Volcan de Fuego, Salvin).
Common in dry or moist oak, pine, or mixed forest and thickets,
1,000-2,600 m.; Chimaltenango; El Quiche; Guatemala; Huehue-
tenango; Jalapa; Sacatepequez; San Marcos; Santa Rosa; Solola.
Mexico (Chiapas); British Honduras; Honduras.
Herbs or weak shrubs, 1-4 m. high, the young stems densely hirsute to finely and
densely sordid, whitish, or yellowish short -pilose, several angled; leaves all opposite,
on petioles 0.5-3 cm. long, the blades lance-ovate to ovate, occasionally lanceolate, 6-
15 (-21) cm. long, 2-10 cm. wide, acute or acuminate at the apex, cuneate at base,
scabrous and hispid above, hirsute to densely short-pilose and occasionally brownish
below, veins often brownish below, the margins serrate, triplinerved; heads 1-3 in
terminal and axillary clusters, the peduncles often stout and fistulose above, 0.5-10
cm. long, the heads to 3.5 cm. wide across extended rays; disc to about 1.5 cm. high,
1.5 cm. wide, or 2.5 cm. wide in fruit; involucres campanulate, becoming
hemispherical in fruit, biseriate; outer phyllaries longer than the inner, foliaceous,
generally spathulate, obtuse at apex, to 2 cm. long, the apex 5-8 mm. wide, white-
hirsute, the inner phyllaries membranous, oblong, obtuse at apex, occasionally
purple-tinged; rays 8-12, yellow or pale yellow (rarely purple?), to 1.5 cm. long; aisc
corollas yellow; anther thecae black; pales stramineous, occasionally purple-tinged,
surpassing achenes; disc achenes 2-awned, thickened, the body 4-5 mm. long, 2-2.5
mm. wide, margins and often 1 awn narrow-winged, faces often strongly tuberculate,
hispidulous; disc pappus squamellae free or united into a cuplike structure about 0.5
mm. long; ray achenes triquetrous, 3-toothed or short-awned, otherwise similar to the
disc achenes.
BECKER: FLORA OF GUATEMALA 357
This relatively large-headed species with its characteristic
spathulate outer phyllaries is common in central Guatemala.
Zexmenia serrata La Llave, Nov. Veg. Descr. 1: 16. 1824 (type
from San Jose del Corral, Mexico, La Llave}. Z. scandens Hemsl.
Biol. Cent. Am. Bot. 2: 174. 1881 (type from Mexico, Bourgeau
1930). Z. trachylepis Hemsl. torn, cit.: 175 (type from Mexico,
Johnson 24). Z. dulcis Coulter, Bot. Gaz. 16: 99. 1891 (type from
Rio Dulce, Izabal, J. D. Smith 1607).
Wet or dry thickets or forest, often in secondary growth and
scrambling over other vegetation, sea level to 700 (1,350) m.; Alta
Verapaz; Izabal; Peten. Southern Mexico; British Honduras; Costa
Rica.
Erect or clambering shrubs to 7 m. high, young stems densely brownish or
yellowish viscid-pubescent, older stems glabrate; leaves all opposite, the petioles
flattened, 0.5-2.5 cm. long, viscid-pubescent, the blades usually firm, rough, often
drying matte-brown above, lance-ovate, lance-oblong, or occasionally ovate, (3-) 6-15
cm. long, (1.5-) 2.5-6 cm. wide, acute or acuminate at apex, cuneate to rounded at
base, scabrous and hispid above, viscid brown-hirsutulous (sometimes densely so)
below, the margins occasionally revolute, generally serrate, varying to coarsely
serrate or denticulate-mucronulate, triplinerved; inflorescence of (1-) 3-5-headed,
terminal and axillary, umbelliform clusters, the peduncles slender, to 4 cm. long,
viscid-pubescent; heads to 2.5 cm. wide across extended rays; involucres broad-
campanulate to hemispherical, 3-4-seriate; phyllaries usually strongly graduate, to 6
mm. high and 10 mm. wide, the outer phyllaries shortest, generally with herbaceous,
often recurved, acute apex, ovate in outline, viscid-puberulent and hispid, margins
ciliate, the inner phyllaries membranous, oblong, obtuse or rounded at apex; ray
flowers 8-12, yellow or deep yellow, ligules to 1 cm. long; disc corollas yellow; anther
thecae blackish; pales shorter than achenial awns; disc achenes 2-awned, when
mature generally with 2 marginal wings, the body thickened, 3.5-4.5 mm. long, about
1 mm. wide (with wings included 3-4 mm. wide), when wingless the body with ciliate
margins, body constricted at apex, faces hispid, body and wings often with brownish
markings; pappus awns 2-4 mm. long, exserted from ripe heads, pappus squamellae
about 0.5 mm. long; ray achenes triquetrous, 3-awned or toothed, otherwise similar to
the disc achenes.
This species, the type of the genus Zexmenia La Llave, is quite
variable in leaf shape and serration. Tun Ortiz 1638, from Dolores,
Peten, is unusual in its ferrugineous pubescence and distally
ampliate inner phyllaries.
Zexmenia virgulta Klatt, Bull. Soc. Bot. Belg. 31: 203. 1892
(type from Costa Rica). Z. dulcis Coulter in Smith, J. D., Enum. PI.
Guat. 4: 86. 1895 (cites J. D. Smith 2854 from Rio Samala,
Retalhuleu), not Bot. Gaz. 16: 99. 1891. Z. chiapensis Brandg. Univ.
Calif. Publ. Bot. 6: 197. 1915 (type from Finca Irlanda, Chiapas,
358 FIELDIANA: BOTANY, VOLUME 24
Mexico, Brandegee 7192). Z. leucactis Blake, Journ. Bot. 53: 307.
1915 (type from Retalhuleu, Bernoulli & Carlo 1396).
Wet thickets, open places, forested flats, sea level-900 (1,500)
m.; Escuintla; Quezaltenango; Retalhuleu; San Marcos; Suchitepe-
quez. Mexico (Chiapas); Costa Rica
Much-branched herbs, sometimes clambering, 1.5-3 m. high, the stems densely
yellowish or brownish viscid-pubescent; leaves opposite, the petioles to 2 cm. long,
viscid-hirsute, the blades thin, lance-ovate or occasionally ovate, 6-14 cm. long, 2-7.5
cm. wide, long-acuminate or caudate-acuminate at apex, cuneate to rounded at base,
softly and densely viscid-hispid above, grayish and softly viscid-pilose to subsericeous
below, the veins generally brownish below, the margins crenate-serrate, triplinerved;
heads solitary or in terminal and axillary clusters of 3 or 5, the peduncles slender, to 3
cm. long, viscid-pubescent; the heads to 2.5 cm. wide across extended rays; involucres
hemispherical, 3-seriate, 5-7 mm. high (excluding tips of outer phyllaries), about 15
mm. wide; outer phyllaries longer than inner, with broad-ovate, membranous base
and lance-oblong or linear, generally recurved, hispid, acute, foliaceous apex 1-2 cm.
long, 1-2 mm. wide, the inner phyllaries membranous below, generally somewhat
herbaceous above, ovate-oblong, obtuse or rounded-mucronulate, the margins ciliate;
ray flowers 8-12, bright yellow; disc corollas yellow; anther thecae brown or dark
brown; pales shorter than achenial awns, sometimes purple-tipped; disc achenes 2-
awned, when mature usually with ciliate marginal wings, thickened, the body 3.5-4.5
mm. long, 2-2.5 mm. wide including wings, constricted at apex, faces hispid; pappus
awns 4-5 mm. long, exserted from ripe heads, pappus squamellae about 1 mm. long;
ray achenes triquetrous, 3-toothed, to 4 mm. wide, otherwise similar to disc achenes.
Near Z. serrata, but is readily distinguished vegetatively by its
herbaceous habit, crenate-serrate leaf margins, and the distinctive
narrow, spreading, foliaceous apices of its outer phyllaries.
ZINNIA Linnaeus
References: B. L. Robinson & J. M. Greenman, A revision of
the genus Zinnia, Proc. Amer. Acad. 32: 14-20. 1896; Andrew M.
Torres, Taxonomy of Zinnia, Brittonia 15: 1-25. 1963.
Annual or perennial herbs, rarely low shrubs; stems pubescent, usually striate;
leaves opposite, sessile or short-petiolate, the blades usually entire, mostly 3-nerved;
heads terminal, long-pedunculate to subsessile, radiate; involucres ovoid-cylindric,
campanulate, or hemispheric; phyllaries 3-many-seriate, broad, closely imbricate,
obtuse or rounded at the apex, often colored and slightly inflated or subsquarrose at
the apex; receptacles conic to slightly concave; pales carinate, becoming scarious,
enveloping the flowers, often erose at the usually obtuse apex; ray flowers pistillate,
fertile, persistent in fruit, the ligules often purple, sometimes red, pink, yellow,
orange, or white; disc flowers hermaphrodite, fertile, the corollas tubular, the limb
little ampliate, 5-dentate; anthers sagittate at the base, appendaged at the apex; style
branches acuminate or truncate, scarcely or not at all appendaged; ray achenes
triquetrous, tridentate, awnless or with 1-3 long or short pappus awns; disc achenes
laterally compressed, glabrous or pubescent, sometimes ciliate, bidentate at the apex
and often awned on the inner angle, rarely 2-awned.
NASH: FLORA OF GUATEMALA 359
Of the 17 species (all American and most of them Mexican, in
the mountains of the tropics or in warm-termperate regions), only
the following two are known in Central America.
Apices of pales conspicuously fimbriate and dentate-lobate; achenes 5-6 mm. long,
the disc achenes awnless; plants cultivated or naturalized Z. elegans.
Apices of pales erose, not dentate-lobate, more or less rounded; achenes 7-10 mm.
long, the disc achenes with a single awn; native plants Z. peruviana.
Zinnia elegans Jacq. Coll. Bot. 5: 152. 1789; Ic. PI. Rar. 3: t.
589. 1793. Z. violacea Cav. Ic. 1: 57. t. 81. 1797. Mulata
(Sacatepequez); berjima (British Honduras); ambolia (Honduras).
Native of the mountains of Mexico, but planted widely in
Guatemala, sometimes escaping from cultivation and more or less
naturalized in waste or cultivated ground. Cultivated for ornament
in almost all parts of the earth and often spontaneous after
cultivation.
Erect annuals, often branching, normally less than 1 m. tall (in cultivation
sometimes to 2.5 m.), the stems often striate, hirsute to pilose-strigose; leaves sessile
and somewhat clasping at the base, ovate to lance-oblong, acute or acuminate,
mostly 3-8 cm. long, the margins entire, more or less scabrous above, sparsely
scaberulous beneath and gland-dotted; peduncles stout, solitary at the ends of
branches; heads solitary; involucres campanulate to hemispheric; phyllaries broadly
obovate, essentially glabrous, broadly rounded at the apex, commonly black-
marginate, erose, squarrose, ciliate; receptacles conical; pales acute or acuminate,
dentate-lobate, the lobes conspicuously fimbriate, pink to reddish-purple; ray flowers
normally 8-20, the ligules 1-2 cm. long, purple or pinkish (considerably longer in
cultivars and variously colored); ray achenes oblanceolate to cuneate, dorsally
compressed, about 6 mm. long; pappus awns wanting; disc flowers numerous, about 7
mm. long, the tubes yellow or orange, the lobes velutinous, black outside; disc
achenes obovate, laterally compressed, about 5 mm. long, ciliolate, emarginate at the
apex; pappus awns wanting.
Zinnia peruviana (L.) L. Syst. ed. 10: 1221. 1759. Chrysogo-
num peruvianum L. Sp. PL 920. 1753. Z. multiflora L. Sp. PI. ed. 2:
1269. 1763. Z. pauciflora L. loc. cit. Z. tenuiflora Jacq. Ic. Rar. 3: t.
509. 1793. Z. revoluta Cav. Ic. 3: 26. 1797. Z. leptopoda DC. Prodr. 5:
535. 1836. Margarita (Huehuetenango); mulata, mulata silvestre
(Guatemala and Jutiapa). Figure 115.
Damp, open or brushy, often rocky slopes, sometimes in oak
forest, 800-2,000 m.; Chiquimula; Guatemala; Huehuetenango;
Jalapa; Jutiapa; El Quiche. Southern Arizona; Mexico; Honduras;
Nicaragua; West Indies; South America.
Erect annuals, simple or sparsely branched, commonly 20-50 cm. tall, the stems
slender or stout, strigose, often becoming purplish; leaves sessile, broadly ovate to
linear-lanceolate or elliptic, mostly 3-8 cm. long, the margins entire, acute or
acuminate, rounded at the base and more or less clasping, thinly scabrous on both
360 FIELDIANA: BOTANY, VOLUME 24
surfaces, glandular-punctate; peduncles short or elongated, slender or inflated
distally; heads solitary; involucres campanulate; phyllaries obovate, essentially
glabrous, rounded at the apex and usually dark-marginate, entire or erose, often
ciliate; receptacles convex to conical; pales glabrous, obtuse or rounded at the apex,
erose, often red, purple, or black near the apex, yellowish below; ray flowers 6-15, the
ligules 0.8-2.5 cm. long, entire, usually brick-red or purplish red, sometimes yellow or
orange; ray achenes linear-oblanceolate or oblong, stria te, sericeous, 9-10 mm. long;
pappus awns wanting, but achenes sometimes crowned by 2 low tufts of hairs; disc
flowers usually numerous, the tube yellow, the lobes yellow within but purplish to
black outside; disc achenes oblong, triquetrous to compressed, striate, ciliolate, 7-9
mm. long; pappus a single awn at the apex (or achenes rarely with 2 unequal awns).
The disc achenes of all specimens examined by me were
provided with only one awn, with the exception of Standley 74907
from Jutiapa. These plants matched the other collections in all
details but some of their achenes have two awns instead of one, the
second one varying from very short to only a little shorter than the
principal one.
EXCLUDED GENERA
OYEDAEA DeCandolle
Of the 13 species of Oyedaea in Mexico and Central and South
America, only one, O. steyermarkii Blake, has been reported from
Guatemala, and as it is certainly not Oyedaea, it has been omitted
here. In his original description, Blake stated, "This species is
ascribed to Oyedaea with some hesitation. The 3 sheets examined
are in very mature fruit, and I was able to find only a single, not
fully developed ray corolla and a few over-mature disc corollas . . .
In any case, further material in less mature condition is to be
desired." Unfortunately, to my knowledge no additional material
has been collected; I have been unable to place it to genus, but the
following description is given in the event it may again be collected.
The description of the ray flowers is based on Blake's description of
the single floret he found; color is not known. No other ray flowers
were found on the very mature heads, and the packet said to
contain the ray flower was empty when I examined the type
specimen.
OYEDAEA STEYERMARKII Blake, Proc. Biol. Soc. Wash. 60: 42.
1947.
Known only from the type collection, Alta Verapaz, Cerro
Chinaja, between Finca Yalpemech and Chinaja, above source of
Rio San Diego, alt. 150-700 m., Steyermark 45686.
WILLIAMS: FLORA OF GUATEMALA 361
Shrub to about 3 m. tall, the stems subterete, striate; leaves opposite, on short
petioles 3-5 mm. long, the blades ovate to lance-ovate or elliptic, 9-12 cm. long, 3-6
cm. broad, acuminate, broadly cuneate at the base, the margins inconspicuously
crenate-serrulate, the upper surface scabrous but shining, in age maculate with the
flattened white bases of scattered, short hairs, the lower surface more or less
pubescent with short, appressed hairs; inflorescences cymose, the heads clustered at
the ends of the branches or in the axils of upper leaves; peduncles 4-10 mm. long, the
pedicels 5-18 mm. long; involucre broadly campanulate, the phyllaries 2-3-seriate, the
outer ones about 6 mm. long, 2 mm. wide, the inner ones 6-7 mm. long, 2.5-3 mm.
wide; ray flowers neutral, the ligules (said to be) about 5 mm. long, tridentate; disc
flowers numerous, the corollas about 4.2 mm. long; pales abruptly acuminate, about
8 mm. long, minutely hispidulous on keel and near apex; abortive ray achenes
trigonous, with 3 awns; disc achenes obovate- oblong, about 4 mm. long, sparsely
strigillose, compressed, 3-angulate, with 2 unequal pappus awns 2-3.2 mm. long, these
smooth, rigid, triangular from a broad base, acuminate, decurrent on the apex of the
achene as narrow wings continuous with the margins of the achene, connate at the
base with the short, irregular squamellae.
TRIBE VI. HELENIEAE
By Louis O. WILLIAMS
Annual or perennial herbs, rarely suffrutescent and shrublike perennials; leaves
opposite with the uppermost alternate, or all alternate (Helenium, Gaillardia);
inflorescences cymose, corymbose, paniculate, or (in Dyssodia decipiens) a
pseudocephalium, often reduced to a single head; involucres cylindric to cam-
panulate, with often a fairly definite number of phyllaries within a genus; phyllaries
few (one in Flaveria) to rather numerous, usually in a single series and subequal in
length but sometimes in two or more series (Helenium, Espejoa, Gaillardia) and
unequal in length; receptacles plane or somewhat convex, naked or somewhat
foveolate, fimbrillate in Gaillardia; the florets in a head may be subdefinite in
number, as few as one (in Flaveria) or very numerous in several genera; heads
commonly radiate but the ray flowers may have very small ligules or these sometimes
completely reduced, PorophyUum has only disc flowers; anthers normally appendaged
at the apex, the base entire or rarely sagittate; achenes normally elongated, not
prominently if at all ridged but often 4- angled, glabrous to sericeous, usually black;
pappus squamellae entire or dissected, often into bristles or setae, or lacking.
The tribe contains a few attractive and decorative plants,
Tagetes, Dyssodia, Helenium, and Gaillardia, as well as a great
many inconspicuous and weedy ones.
The tribe Helenieae is not greatly different from the tribe
Heliantheae and by some botanists the two are joined. The
Guatemalan genera of the tribe Helenieae seem to be fairly distinct
from the helianthoid kinds, and as a matter of convenience the tribe
is maintained.
The principal and most consistent differences between the
362 FIELDIANA: BOTANY, VOLUME 24
Helenieae and the Heliantheae in our region, as elsewhere, is the
lack of pales on the receptacle of the Helenieae, while in the
Heliantheae the receptacle is paleaceous, i.e. with chaffy bracts
subtending the flowers in the heads.
There are 12 helenioid genera in Guatemala, to which these
comments apply. They are mostly found at low or middle
elevations, especially those that tend to be weedy. A few, Bahia
(one endemic species), Helenium, and Tagetes, are found at very
high elevations.
Heads l-(2-)flowered, the flowers either radiate and pistillate or perfect Flaveria.
Heads mostly 4-many-flowered, the flowers various.
Heads discoid, mostly more than 1.5 cm. long; plants glabrous Porophyllum.
Heads radiate, or if discoid the heads much less than 1.5 cm. long; plants glabrous
or pubescent.
Involucre of numerous, imbricated phyllaries.
Leaves opposite; plants glabrous or nearly so Espejoa.
Leaves alternate; plants copiously pubescent.
Style branches truncate; native plants Helenium.
Style branches with subulate appendages; plants cultivated or rarely
escaping and somewhat naturalized Gaittardia.
Involucre uniseriate or nearly so or, if imbricate, usually 5 or very few phyllaries.
Pappus none; ray achenes with dentate margins; pubescent annuals.
Galeana.
Pappus present; achenes with entire margins.
Foliage without conspicuous oil glands.
Plants very small, mostly 2 cm. high or less; leaves small, all 3-parted;
heads usually with about 8 flowers Bahia.
Plants larger, mostly 30-50 cm. high; the upper leaves linear to ovate.
Principal phyllaries of involucre 4-5, punctate Schkuhria.
Principal phyllaries 5-10, not punctate Florestina.
Foliage with conspicuous oil glands.
Pappus of setae, these sometimes reduced to squamellae; leaves simple.
Pectis.
Pappus of squamellae, these often dissected or awned; leaves simple or
pinnatisect.
Pappus squamellae not dissected Tagetes.
Pappus squamellae dissected into bristles Dyssodia.
BAHIA Lagasca
Reference: Ellison, W. L., Systematic studies in the genus
Bahia, Rhodora 66: 177-215, 281-311. 1964.
Diffuse, dwarf annuals; stems slender, hirsute and glandular-pilose with
multicellular hairs; leaves opposite, few, short petiolate, leaf blades tripartite, the
lobes sometimes redivided; inflorescence a single head or sometimes 2-3 terminal or
axillary heads, these mostly 6-8-flowered; involucre uniseriate; phyllaries about 5,
WILLIAMS: FLORA OF GUATEMALA 363
obovate, membranaceous or subherbaceous, with narrow, colorless, scarious margins
and yellowish, black-lineolate apices; ray flowers yellow, fertile, the tube glandular-
pilose, rays with 1-2 blunt teeth on the inner side, the lamina rounded and usually
emarginate at the apex; disc flowers perfect, broadly campanulate, 5-dentate;
receptacle small, flattened, glabrous, strongly scrobiculate; achenes all similar,
obovoid, trigonous, with conspicuous carpopodium; pappus very short, with 10
obovate or suborbicular, subtruncate or erose-dentate squamellae; style branches
with short, deltoid, dorsally papillose appendages.
A small genus of perhaps a dozen species, only the following in
Guatemala. The generic description is written from our species and
the closely related B. anthemoides (HBK.) Gray. These, if separated
from Bahia, might be placed in Achyropappus. Ellison, in his
account of Bahia, excluded both, placing B. anthemoides in
Achyropappus;-and B. depauperata into Vasquezia for no ex-
plained, nor obvious, reason.
Bahia depauperata Blake, Brittonia 2: 352. 1937. Vasquezia
depauperata Ellison, Rhodora 66: 308. 1964. Figure 116.
Alpine meadows or debris-covered rocks and cliffs, 3,000-3,700
m.; endemic; Huehuetenango (type, Skutch 1271); Solola; Quezal-
tenango.
Small, diffuse annuals, up to about 1.5 cm. high, the stems short-hirsute and
glandular-pilose; petioles 3-5 mm. long, hispid-ciliate; leaf blades 2.5-5 mm. long,
tripartite, the segments entire or again tripartite, obtuse, glandular-pilose and
hispidulous, especially beneath; peduncles slender, 3-4 mm. long; inflorescence
usually a single head, rarely 2-3, terminal or axillary; heads mostly 6-8-f lowered;
involucre about 3 mm. high; phyllaries about 5, obovate, with narrow scarious
margins, glandular-pilose and sparsely hirsute; rays yellow; disc corollas about 1.3
mm. long, glandular-pilose; achenes trigonous, obovoid, about 2 mm. long, black,
glabrous on the outer flat surface, sparsely to densely hispidulous on the inner
surface; pappus very short, with 10 obovate or suborbicular, subtruncate or erose-
denticulate squamellae.
Dr. Blake thought his specimens to be depauperate, but recent
collections and material in the field are much like those that he
had. Ellison, in his revision of Bahia, excluded this species by
placing it into Vasquezia. No explanation was given for the change,
and, in fact, the plant hardly belongs in Vasquezia. Dr. Cuatrecasas
(in lit.) believes that Blake was correct in his generic placement of
this species.
DYSSODIA Cavanilles
Reference: John L. Strother, Systematics of Dyssodia Cav-
anilles, Univ. Cal. Publ. Bot. 48: 1-88, illus. 1969.
364 FIELDIANA: BOTANY, VOLUME 24
Annual or perennial herbs, rarely low shrubs, glabrous or pubescent, usually
strong-scented and with numerous, conspicuous oil glands on the involucre; leaves
opposite or alternate, dentate or pinnately dissected; inflorescence single-pedunculate
to sessile heads or several heads glomerate into a pseudocephalium; heads small or
rather large, heterogamous, radiate, yellow; ray flowers pistillate, uniseriate, the disc
flowers fertile, the rays sometimes absent; involucre cylindric or campanulate;
phyllaries 1-2-seriate, subequal, rigid-membranaceous, free or united below, a
subtending calyculum present or absent; receptacle flat or nearly so, fimbrillate or
not; rays entire, spreading; hermaphrodite flowers regular, the limb scarcely
ampliate, shallowly 5-fid; anthers entire at the base or minutely bidentate; style
branches of the hermaphrodite flowers acute, hispiduluous at the apex, short or
elongate; achenes obconic or obpyramidal, pubescent, 3-5-angulate, striate; pappus of
10-20 squamellae, these setaceous, deeply divided into numerous setae.
Species about 30, all American, chiefly in tropical and warm-
temperate regions. Only the following are known in Central
America. The genus Syncephalantha, which is easily distinguished
superficially from Dyssodia, is included here, since the pseudocepha-
lium shows transition to some species of Dyssodia.
Heads several, borne in a pseudocephalium or glomerate into secondary heads.
D. decipiens.
Heads not borne in a pseudocephalium nor glomerate into secondary heads.
Heads small, involucre usually about 7 mm. high D. papposa.
Heads large, the involucre usually 12-15 mm. high
Leaves serrate; heads radiate D. montana.
Leaves mostly pinnatifid; heads either radiate or eradiate D. porophylla.
Dyssodia decipiens (Bartling) M. C. Johnston, Rhodora 64:
13. 1962; L. Wms. Phytologia 29. 273. 1974. Syncephalantha
decipiens Bartling, Ind. Sem. Hort. Goett. 1836: 6. 1836; Schlech-
tendahl, Linnaea 12: Lit. Ber. 80. 1838. S. sanguinea Klatt,
Leopoldina 25: 106. 1889 (type from Guatemala, Warszewicz 87). S.
macrophylla Klatt, torn. cit. 108. Dyssodia sanguinea Strother,
Univ. Cal. Publ. Bot. 48: 25, fig. 1969. Flor de muerto; boton de oro;
cambray de rosa.
Open, grassy, rocky slopes or meadows, moist or rather dry
brushy plains or hillsides, frequently on sandbars along streams,
often a weed in cultivated fields, 600-2,000 m.; Chiquimula; Jalapa;
Jutiapa; Santa Rosa; Guatemala; Sacatepequez; Chimaltenango; El
Quiche; San Marcos. Southern Mexico.
Erect or spreading, usually 30-60 cm. high, often much branched, the branches
angulate, sparsely pilose or glabrate; leaves 2-5 cm. long, pinnately dissected, the
divisions 4-9, oblanceolate or cuneate, dentate, sparsely pilose beneath on the veins,
with marginal oil glands; aggregate heads about 1.5 cm. broad; involucre 5-6 mm.
WILLIAMS: FLORA OF GUATEMALA 365
high; phyllaries oblong-obovate, glabrous, dark red or purplish, striate, with
membranaceous margins; rays bright yellow, 7-9 mm. long; disc corollas yellow, 3
mm. long; achenes black, 3 mm. long, pubescent; pappus yellowish, 4 mm. long.
The plant is a very common one in many places in Guatemala,
often a conspicuous and abundant weed in corn fields, in
appearance suggesting species of Tagetes.
Dyssodia montana (Benth.) Gray, Proc. Am. Acad. 19: 38.
1883. Clomenocoma montana Benth. PI. Hartw. 86. 1841 (type from
Mountains of Aceituno, Guatemala, Hartweg 592). Valeriana;
mirasol; flor amarilla; Paulina; flor de fuego (fide Aguilar). Figure
117.
Moist or rather dry thickets, sometimes on open slopes, 200-
1,900 m.; Baja Verapaz; Chiquimula; Jalapa; Jutiapa; Santa Rosa;
Guatemala; Chimaltenango; Quezaltenango. Southern Mexico; El
Salvador to Panama.
Probably perennial, herbaceous, erect, 1 m. high or less, or sometimes with more
elongate and somewhat scandent stems, the stems stout, sparsely branched, terete,
glabrous or sparsely hirsutulous, striate; leaves opposite, short-petiolate or sessile,
rather thick and firm, lanceolate or ovate, 4-8 cm. long, acute or acuminate at each
end, sharply and closely serrate, the very base of the blade usually dissected into a
few small and inconspicuous, almost filiform lobes, green above, paler beneath,
sometimes densely hirsutulous but usually almost glabrous; heads large, orange,
solitary at the ends of the long, stout, naked peduncles; involucre 12-14 mm. high,
hemispheric, as much as 2.5 cm. broad, the accessory bracts at the base of the
involucre few and small, subulate to ovate; phyllaries 12-15, oblong to elliptic, obtuse
or rounded at the apex, fimbriate-crenate at the apex, glabrous, bearing a few
conspicuous, linear or oblong glands; ray flowers 12-15, the rays oblong-obovate,
about 1 cm. long; disc corollas 8-9 mm. long; achenes striate, sericeous-villous, 3 mm.
long; pappus squamellae 10, 7-8 mm. long, dissected almost to the base into numerous
fulvous bristles.
This species has been most often determined as Dyssodia
grandiflora DC.
Dyssodia papposa (Vent.) Hitchc. Trans. Acad. Sci. St. Louis
5: 503. 1891. Tagetes papposa Vent. Descr. Cels. t. 36. 1802.
Dyssodia fastigiata DC. Prodr. 5: 640. 1836. Boeberia roseata Rydb.
N. Am. Fl. 34: 168. 1915.
Open slopes or banks, sometimes in oak forests and in old fields,
1,600-2,200 m.; El Quiche; Huehuetenango; San Marcos. A wide-
spread and often weedy species, from New England (where perhaps
introduced) to the Rocky Mountain states then south through
Mexico to Guatemala. Introduced in Argentina.
366 FIELDIANA: BOTANY, VOLUME 24
An erect, rather stout annual, 40 cm. high or less, sparsely or much branched, the
stems sparsely or densely villous or villous-hirsute; leaves opposite, 2-4 cm. long,
pinnatifid or bipinnatifid, the segments linear-oblanceolate, often dentate, spinulose-
mucronate, scabrous-hispidulous on the veins; heads small, sessile or short-
pedicellate, in small clusters at the ends of the branchlets; involucre turbinate, 5-7
mm. high and broad; principal phyllaries about 8, oval, obtuse, tinged with brown or
red, glabrous, with 1-few large glands, or sometimes eglandular; accessory bracts few,
linear, short; ray flowers few, the rays scarcely more than 1 mm. long, yellow; disc
corollas yellow, 3 mm. long; achenes 4 mm. long, compressed and somewhat angulate,
hirsute; pappus 4-5 mm. long, exceeding the disc flowers, the squamellae dissected
into slender bristles.
Dyssodia porophylla (Cav.) Cav. An. Cienc. Nat. 6: 334. 1803.
Pteronia porophyllum Cav. Ic. 3: 13, t. 225. 1794. Adenophyllum
porophyllum Hemsl. Biol. Cent. Am. Bot. 2: 218. 1881. Lebetina
porophylla A. Nels. Bot. Gaz. 47: 435. 1909. Dyssodia porophylla
var. radiata DC. Prodr. 5: 639. 1836. Dyssodia porophylla subsp.
cancellata (Cass.) Strother var. radiata (DC.) Strother, Univ. Cal.
Publ. Bot. 48: 39. 1969. Copa de rey; varilla de San Jose (fide
Aguilar).
Grassy, often stony, dry or moist plains or hillsides, sometimes
on sandbars, or a weed in cultivated ground, 400-1,900 m.:
Chiquimula; Jalapa; Jutiapa; El Progreso; Guatemala; Sacatepe-
quez; Chimaltenango; El Quiche; Huehuetenango. Southern Mex-
ico; El Salvador to Panama.
A stout, erect annual, 30-75 cm. high, simple or corymbosely branched above, the
branches suberect, glabrous, striate; leaves alternate, 5 cm. long or less, pinnately
parted, the segments oblong or obovate, short, dentate, the teeth bristle- tipped, the
lobes with conspicuous marginal glands; heads mostly solitary at the ends of the long
branches or somewhat corymbose, the peduncles bearing numerous, small, reduced
leaves; involucre turbinate, 12 mm. high and broad; phyllaries about 20, acuminate,
bearing a hornlike appendage 1 mm. long; accessory bracts 15-20, pectinately divided
into 3-6 pairs of rigid spinelike divisions; rays none or small; disc corollas bright
yellow, 5 mm. long; achenes 5 mm. long, striate; pappus squamellae 6-7 mm. long,
each dissected into 7-10 fulvous bristles.
This species bears heads with and without ray flowers. Those
plants without ray flowers (fairly rare in Guatemala) should be
called D. porophylla (Cav.) Cav.; those with small but usually
conspicuous ray flowers should be called D. porophylla var. radiata
DC.
ESPEJOA De Candolle
Reference: P. A. Rydberg, N. Am. Fl. 34: 3. 1914.
WILLIAMS: FLORA OF GUATEMALA 367
Erect annuals, almost glabrous, branched; leaves small, opposite, sessile, entire;
heads small, discoid, on long, naked peduncles, solitary in the forks of the branches;
involucre campanulate; phyllaries 8-12, herbaceous, 4-seriate, broad; receptacle
alveolate; flowers all fertile; corollas glabrous, with sparsely pubescent lobes, the
tube short, enlarged at the base, the throat funnelform, the teeth lanceolate; anthers
linear, subcordate at the base, the tips lanceolate; style branches obtuse, sparsely
hispidulous; achenes cuneate, somewhat compressed dorsiventrally, pubescent,
truncate at the apex; pappus scales about 16, ovate-oblong, 1-nerved, mucronate.
The genus consists of a single species.
Espejoa mexicana DC. Prodr. 5: 660. 1836. Jaumea mexicana
Benth. & Hook, ex Hemsl. Biol. Cent. Am. Bot. 2: 209. 1881. Figure
118.
Weedy field, 150 m.; Santa Rosa (along Avellana Road south of
Guazacapan, Standley 79432). Southern Mexico; El Salvador;
Honduras; Nicaragua; Costa Rica.
Plants erect, 30-75 cm. high, dichotomously branched, the stems pale, subterete,
glabrous or sparsely short-pilose above; leaves opposite, elliptic or oblong,
membranaceous, mostly 3-6 cm. long, obtuse, apiculate, rounded to acute at the base,
penninerved, glabrous, ciliate, somewhat paler beneath; peduncles slender but stiff, 3-
8 cm. long, phyllaries very unequal, the inner ones 7-9 mm. long, oval or obovate,
striate, broadly rounded and apiculate at the apex, ciliate, the outer ones very short,
orbicular; corollas yellow; achenes narrowly cuneate, 6-7 mm. long, very densely
white-sericeous; pappus scales white, with a brown costa, erose-denticulate or entire.
This species has often been referred to Jaumea but it does not
seem to the author of this work to be congeneric with the type
species of that genus. Rare in Central America, except common in
the Rio Yeguare valley in central Honduras. Usually in weedy
situations.
FLAVERIA Jussieu
Annual herbs, glabrous or minutely puberulent, the stems erect and much
branched; leaves opposite, entire or dentate; inflorescence of dense cymes or
glomerules (ours), rarely corymbose-paniculate, borne in the axils or at the ends of
branches; heads sessile or nearly so, heterogamous; involucre of 1-8 subequal
phyllaries (ours 1-2), the outer sometimes much reduced; ray flowers ligulate, small;
disc flowers tubular, the limb turbinate-campanulate, 5-dentate; anthers entire at the
base; style branches truncate; achenes oblong, 8-10-costate; pappus wanting (ours).
A genus of about a dozen species, only the following is known
from Central America, that from a single collection from British
Honduras.
Flaveria trinervia (Spreng.) C. Mohr, Contr. U. S. Nat. Herb.
6: 810. 1901. Oedera trinervia Spreng. Bot. Gart. Halle 63. 1800.
368 FIELDIANA: BOTANY, VOLUME 24
Probably a weed, Corozal District, British Honduras (Gentle
147}. Southern and southwestern United States; Mexico; West
Indies; South America.
Erect, divaricately branched annuals to perhaps 1 m. tall; stems subterete or 4-
angulate, glabrous or obscurely puberulent on new growths; leaves opposite, elliptic
to oblanceolate, 3-nerved, denticulate, the blades to 9 cm. long, attenuate to slender
petioles; inflorescence of sessile or short pedunculate, fasciculate heads, axillary or
terminal; heads small, usually 1-flowered; phyllaries 1-2, about 2 mm. long; corolla of
pistillate flowers about 3 mm. long, the ligule tridentate; corolla of perfect flowers
about 2 mm. long, the tube villous, the throat campanulate; achenes somewhat
clavate, about 2 mm. long; pappus none.
FLORESTINA Cassini
Reference: B. L. Turner, Taxonomy of Florestina (Helenieae,
Compositae), Brittonia 15: 27-46. 1963.
Plants annual, erect, pubescent and more or less glandular; leaves petiolate, the
lower ones opposite, the upper alternate, all except the uppermost pinnately,
palmately, or pedately 3-7-parted, or entire and 3-nerved; heads small, whitish,
numerous, discoid, in flat-topped panicles; involucre turbinate or campanulate;
phyllaries 5-10, obovate, scarious-marginate, flat or somewhat concave, not carinate;
flowers 10-20, hermaphrodite, fertile; corolla tube short, funnelform, glandular-
puberulent, the lobes oblong, longer than the tube; style branches terminated by
slender hispid appendages; achenes narrowly obpyramidal, truncate, 4-angulate,
pubescent; pappus squamellae scarious, somewhat thickened at the base, costate or
ecostate, sometimes awn- tipped.
Eight species, ranging from southwestern United States to
Guatemala.
Leaves simple, never deeply parted or divided, the blade triangular to cordate.
F. latifolia.
Leaves conspicuously lobate, at least above, usually 3-5 parted F. pedata.
Florestina latifolia (DC.) Rydb. N. Am. Fl. 34: 58. 1914.
Palafoxia latifolia DC. Prodr. 5: 125. 1836. Figure 119.
Thickets and savannas, 300-1,200 m. Southern Mexico;
Honduras; Nicaragua; to be expected in Guatemala.
Erect annuals to 60 cm. tall, sparsely branched below, short pubescent
interspersed with viscous-glandular hairs; leaves simple, the lower opposite, with
subscabrous pubescence, blade ovate to triangular, puberulent, serrate or lobulate, 2-8
cm. long and 1.5-6 cm. broad; inflorescence dichotomously branched, with few to
many turbinate heads; heads 0.5-1 cm. broad; phyllaries to 10, 5-9 mm. long and half
as broad, mostly obtuse; corollas mostly 6-7 mm. long with a short tube and a
cylindric throat, 4-5 mm. long, the lobes unequal, 0.5-2.5 mm. long; achenes
obpyramidal, 4-angled, puberulent to glabrous; pappus of about 8 scales, the longer
WILLIAMS: FLORA OF GUATEMALA 369
ones lanceolate, mucronate, the shorter ones truncate or obtuse, or all the scales
similar.
Florestina pedata (Cav.) Cassini, Diet. Sci. Nat. 17: 155. 1820.
Stevia pedata Cav. Ic. 4: 33. 1797. Schkuhria viscosissima Standl. &
Steyerm. Field Mus. Bot. 22: 318. 1940 (type from El Quiche,
Standley 62513). Florestina viscosissima Heiser, Ann. Mo. Bot.
Card. 32: 278. 1945.
Dry or wet rocky thickets or meadows, sometimes a weed in
cultivated fields, 800-2,000 m.; El Quiche; Huehuetenango. Mexico.
A slender or stout, erect annual, 25-50 cm. high, often much branched, the
branches ascending, striate, densely pilose, with spreading gland-tipped hairs; leaves
pinnately lobate, with few linear lobes, or simple and linear, densely scabrous or
hispidulous, the margins revolute; heads numerous, greenish white, slender-
pedicellate, 6 mm. high and broad; phyllaries 5-6, oblong-obovate, 4.5-5 mm. long,
very obtuse or rounded at the apex, appressed-hispidulous, pale-marginate; achenes
narrowly turbinate, 3 mm. long, blackish, densely hispidulous; pappus squamellae 8,
obovate-orbicular, almost 1 mm. long, rounded at the apex, whitish, slightly
thickened at the base.
The plant perhaps is common in El Quiche and Huehuetenango
during the wet months, but only a few scattered plants are seen
during the dry season. Apparently, they are eaten by the many
goats and sheep pastured in the region.
GAILLARDIA Fougeroux
Reference: P. A. Rydberg, N. Am. Fl. 34: 131-141. 1915.
Plants annual, biennial, or perennial, usually herbaceous, scapose or caulescent;
leaves alternate or basal, usually narrow, entire, dentate, or pinnatifid; heads radiate
or discoid, long-pedunculate; involucre rotate or saucer-shaped in anthesis; phyllaries
2-3-seriate, ovate to lanceolate, more or less reflexed in fruit; receptacle convex to
globose, alveolate, usually fimbrillate; ray flowers usually neutral, sometimes none,
the rays broad, 3-cleft, yellow or purple; disc flowers hermaphrodite, fertile; corolla
tube very short, glabrous, abruptly dilated into the throat, the lobes ovate to
subulate, pubescent with glandular, often moniliform hairs; anthers auriculate at the
base; style branches appendaged, the appendages short and glabrous or filiform and
hispidulous; achenes broadly obpyramidal, wholly or partly covered with long, stiff,
ascending hairs; pappus of 5-10 squamellae.
Species about 30, in temperate and warm-temperate regions of
America, mostly in North America. Numerous species are native in
Mexico but none in Central America.
Gaillardia pulchella Foug. Mem. Acad. Sci. Paris 1786: 5.
1788. G. picta Sweet, Brit. Fl. Gard. II, t. 267. 1834. G. pulchella var.
370 FIELDIANA: BOTANY, VOLUME 24
picta Gray, Syn. Fl. N. Am. 1, pt. 2: 352. 1884. Gallarda; gallardia;
gallardina.
Native in western and southern United States; often grown for
ornament in other regions; a common garden plant of Guatemala,
at almost all elevations; sparingly naturalized in cornfields about
Chichicastenango (El Quiche).
An erect or ascending annual, 25-50 cm. high, usually branched, short-hirsute or
puberulent, the stems pale, striate; lower leaves oblanceolate or linear-oblanceolate,
pinnate- lo bate to entire, pubescent; peduncles long and slender, naked; phyllaries
lanceolate, herbaceous, pubescent and ciliate, long-acuminate; receptacle fimbrillate,
the fimbrillae longer than the achenes; ray flowers neutral, ligules yellow or yellow
and purple, 1.5-2 cm. long, deeply 3-cleft, corollas yellow below, purple above, 6-7
mm. long, the lobes acuminate, ciliate with moniliform hairs; achenes 2-2.5 mm. long,
densely hirsute; pappus squamellae 5-6 mm. long, lanceolate, attenuate into a slender
awn.
Sometimes called "cambray" in El Salvador, "pompon" in
Honduras, and "jalacate" or "jalacate extranjero" in Nicaragua.
One of the common garden flowers of Guatemala, frequently
planted in beds in the city parks. The flowers often are on sale in
the markets. The usual cultivated plants are referable to var. picta
(Sweet) Gray.
GALEANA La Llave
Reference: P. A. Rydberg, N. Am. Fl. 34: 42-43. 1914.
Slender annuals, dichotomously branched, glandular-pubescent and glandular-
punctate; leaves chiefly opposite, petiolate, dentate; inflorescences small, leafy
cymes, in the forks of the branches, pedicellate; phyllaries 5, broad, concave, oval;
ray flowers 3, pistillate and fertile, the rays short, white, broadly cuneate, 3-dentate;
disc flowers 3-4, yellow, 1-2 of them hermaphrodite and fertile, the others staminate
and sterile; corolla tube of the hermaphrodite flowers short, glandular, the throat
short-campanulate; staminate corollas longer, glabrous; achenes of the hermaphro-
dite flowers pyriform, triquetrous, those of the ray flowers broader, their inner angles
with somewhat corky, concave wings, having curved crenate margins; pappus none.
Three species, the other Mexican.
Galeana pratensis (HBK.) Rydb. N. Am. Fl. 34: 42. 1914.
Unxia pratensis HBK. Nov. Gen. & Sp. 4: 279. 1820. Figure 120.
Moist or wet thickets or open fields, often in oak forest, 800-
2,100 m.; Chiquimula; Jalapa; Jutiapa; Santa Rosa; Guatemala;
Chimaltenango; El Quiche; Huehuetenango. Southern Mexico; El
Salvador; Honduras; Costa Rica.
An erect annual, commonly 25-50 cm. high, often much branched, viscid-
pubescent throughout, the stems angulate; leaves opposite, short-petiolate, deltoid-
WILLIAMS: FLORA OF GUATEMALA 371
ovate, mostly 2-3 cm. long, obtuse or acute, subtruncate or broadly cuneate at the
base, 3-nerved, coarsely and irregularly dentate, often subhastate, the upper leaves
narrower and often subentire; heads in small, pedunculate cymes, pedicellate;
involucre obovoid, 3 mm. long; phyllaries 5, green, densely glandular-pubescent and
punctate, obtuse, appressed; rays scarcely 1 mm. long; achenes black, glabrate, 3 mm.
long, obscurely tuberculate; wings of the marginal achenes narrow, pale at first, dark
in age, undulate, glabrous.
Sometimes known in El Salvador as "hierba sana," and
reported to be used there in domestic medicine as a febrifuge. The
Mexican G. hastata La Llave may not be distinct from this species.
HELENIUM Linnaeus
References: P. A. Rydberg, N. Am. Fl. 34: 119-131. 1915. Mark
W. Bierner, Taxonomy of Helenium sect. Tetrodus and a conspectus
of North American Helenium, Brittonia 24: 331-355. 1972.
Annual or perennial herbs, erect, puberulent or lanate; leaves alternate, often
decurrent, entire or coarsely dentate; heads heterogamous, radiate, pedunculate,
medium-sized and often corymbose, or large and solitary or few; ray flowers 1-seriate,
fertile or neutral, the rays and often the disc yellow, the disc flowers hermaphrodite,
fertile; involucre broad; phyllaries 1-2-seriate, the outer ones herbaceous, narrow,
subequal, spreading or in age reflexed, the inner ones usually broader, membrana-
ceous or hyaline; receptacle convex to subglobose or oblong, naked; rays spreading,
usually cuneate, 3-5-lobate at the apex; hermaphrodite flowers regular, tubular, the
limb 4-5-dentate, the teeth glandular-pilose; anthers sagittate at the base, the
auricles small, generally acute; style branches of the hermaphrodite flower truncate-
dilated at the apex; achenes turbinate, costate, sericeous or rarely almost glabrous;
pappus scales 5-8 or none or much reduced, when present hyaline, acuminate, aristate
or muticous, entire, dentate, or lacerate.
Some 30 species, most of which are to be found in Mexico and
the United States. Only two species are known in Central America,
although H. scorzoneraefolium (DC.) Gray is to be expected in
Guatemala.
Stems winged, puberulent above; disc of the head about 1 cm. broad.
H. mexicanum.
Stems not winged, often ridged, densely lanate above; disc mostly 2 cm. or more
broad.
Pappus present and conspicuous; pubescence of peduncle pale H. integrifolium.
Pappus absent or inconspicuous; pubescence of peduncle sordid to blackish purple.
H. scorzoneraefolium.
Helenium integrifolium (HBK.) Benth. & Hook, ex Hemsl.
Biol. Cent. Am. Bot. 2: 227. 1881. Actinea integrifolia HBK. Nov.
Gen. & Sp. 4: 297, t. 410. 1820. Dugaldia integrifolia Cass. Diet. Sci.
Nat. 55: 271. 1828. Oxylepis lanata Benth. PI. Hartw. 87. 1841 (type
372 FIELDIANA: BOTANY, VOLUME 24
from Cumbre de Argueta, Solola, Hartweg 593). Helenium lanatum
Gray, Proc. Am. Acad. 9: 205. 1874. Machul; quiaquen; veneno;
mirasol. Figure 121.
Subalpine meadows in the Sierra de los Cuchumatanes at 2,800-
3,700 m.; Huehuetenango; possibly also in Solola, although not seen
there by us. Mountains of central and south Mexico.
A coarse, stout perennial, 1 m. high or less, from a thick rootstock, the stems
simple below, sometimes branched above, usually densely leafy throughout, densely
lanate above; basal leaves oblanceolate or linear-oblanceolate, mostly 20-30 cm. long,
1-4 cm. broad, obtuse or acute, long-attenuate to the base, rather thick and fleshy,
entire, dilated at the base, the cauline leaves shorter, not decurrent, lanate or almost
glabrous; heads few or numerous, sometimes solitary, long-pedunculate, the
peduncles stout, lanate, bearing several linear bracts; disc 2.5 cm. broad or larger;
phyllaries oblong-linear, green, about 1.5 cm. long, attenuate, densely lanate;
receptacle very convex; ray flowers about 40, the rays bright yellow, 2-3.5 cm. long;
achenes almost 4 mm. long, densely appressed-pilose; pappus squamellae 5-7,
triangular-lanceolate, attenuate, stiff, white, somewhat shorter than the achene.
The plant is an abundant one in many places in the subalpine
meadows of the Cuchumatanes. This region is badly overgrazed by
sheep but the Helenium (along with a few other plants) seems not
to be grazed and is said to be poisonous to sheep. Whether it is
poisonous or just unpalatable to the sheep we do not know. The
plant is an attractive one when in flower, a stem often having as
many as seven heads.
Helenium mexicanum HBK. Nov. Gen. & Sp. 4: 229. 1820. H.
centrale Rydb. N. Am. Fl. 34: 125. 1915 (type from Santa Rosa,
Heyde & Lux 3403). Piretro (fide Aguilar).
Open slopes or meadows, 900-2,000 m.; Chiquimula; Jalapa;
Santa Rosa; Guatemala; El Quiche. Mexico (Chiapas); Honduras;
Costa Rica.
Plants annual, or probably often more enduring, erect, usually densely fastigiate-
branched, 1 m. high or less, puberulent and glandular-granuliferous, the branches
broadly winged; basal leaves oblanceolate, 5-10 cm. long, entire or denticulate,
puberulent and glandular-punctate, the cauline leaves linear or lance-linear,
attenuate at each end; peduncles erect, usually numerous, 3-10 cm. long; phyllaries
subulate, 4 mm. long; ray flowers pistillate, the rays 4 mm. long, yellow; disc globose,
brown, 1 cm. long in anthesis, in fruit larger; disc corollas 1.5 mm. long; achenes 1.5
mm. long, hispidulous on the angles; pappus squamellae rounded-ovate, apiculate.
Helenium scorzoneraefolium (DC.) Gray, Proc. Am. Acad. 7:
359. 1868. Hecubaea scorzoneraefolia DC. Prodr. 5: 665. 1836.
Similar to H. integrifolium but distinguished by the pappus
WILLIAMS: FLORA OF GUATEMALA 373
wanting or very much reduced, the corolla tube longer, and usually
the stems with fewer heads, usually only 1, instead of several.
The species occurs near Guatemala, in Chiapas, and may be
expected in Guatemala.
PECTIS Linnaeus
Reference: P. A. Rydberg, N. Am. Fl. 34: 194-216. 1916.
Mostly low, annual or perennial herbs, erect to prostrate, strong-scented,
branched, glabrous or pubescent; leaves opposite, narrow, small, gland-dotted,
generally entire, commonly with several pairs of marginal bristles near the base;
heads small, yellow, solitary or cymose, radiate; involucre cylindric to campanulate
or turbinate; phyllaries 3-12, free, 1-seriate, without bracts at the base, usually gland-
dotted, carinate, at least below, often gibbous at the base; receptacle small, naked;
ray flowers few, pistillate, fertile, usually as many as the phyllaries; disc flowers few,
hermaphrodite, fertile; corolla tube usually short, gradually dilated into the narrow
5-fid limb; anthers obtuse and entire at the base; style hispidulous, the short
branches obtuse, not appendaged; achenes linear, terete or angulate, pubescent or
glabrate; pappus of few or many squamellae, awns, or bristles, rarely reduced to a
low crown.
Species about 75, or perhaps fewer, all American and mostly in
North America, in tropical or warm-temperate regions.
Heads sessile or nearly so, not exceeding the leaves P. prostrata.
Heads on slender, often greatly elongated peduncles.
Heads about 1.5 cm. long; vines of the Pacific strand P. multiflosculosa.
Heads 1 cm. long or less; inland plants.
Pappus of well developed squamellae, these terminating in long bristles, but the
squamellae evidently dilated at the base, triangular.
Phyllaries 6-8, about 4 mm. long P. bonplandiana.
Phyllaries 5, 4.5-6 mm. long P. capillipes.
Pappus, at least in the ray flowers, of bristles or awns, rarely paleaceous at the
base, rarely with minute outer squamellae, sometimes reduced to a minute
crown.
Heads, from the base of the involucre to the ends of the disc flowers, about 10
mm. long, campanulate P. saturejoides.
Heads 7 mm. long or usually shorter, turbinate or oblong.
Pappus a very short crown of minute squamellae.
P. uniaristata var. holostemma.
Pappus wholly or partly of conspicuous setae or awns.
Pappus of 2-3 stout, short awns P. linifolia.
Pappus of 5-20 or more conspicuous setae P. elongata.
Pectis bonplandiana HBK. Nov. Gen. & Sp. 4: 263. 1820.
Culantrillo.
Moist or dry, brushy plains and hillsides or open banks, 250 m.
374 FIELDIANA: BOTANY, VOLUME 24
or less; Alta Verapaz; Izabal; Retalhuleu. Mexico; British
Honduras; El Salvador; Panama.
Plants slender, erect or almost prostrate, annual, dichotomously branched, the
stems often dark red or purplish, 30 cm. high or less, often densely branched; leaves
linear or lance-linear, 1-2.5 cm. long, 1-3 mm. broad, mucronate, entire or minutely
serrulate toward the apex, with 1-3 pairs of bristles near the base; heads on pedicels 3
cm. long or shorter, terminal and in the forks of the branches; involucre 4-4.5 mm.
high; phyllaries 6-8, linear-oblanceolate, subacute or obtuse, minutely puberulent
about the apex, carinate for about half their length; ray flowers 6-8, the rays almost
2 mm. long; disc flowers 10-15; achenes 2 mm. long, pilose; squamellae of the disc
flowers 5-6, those of the ray flowers 2, awn-tipped, almost equaling the disc corollas.
Called "comino" in El Salvador. The plants of this and other
species have usually a strong and distinctive odor, somewhat
suggestive of that of so-called stink bugs. It doubtless has its origin
in the numerous oil glands found on the leaves and usually also on
the involucres. In British Honduras P. bonplandiana sometimes
grows in open places along or near sea beaches.
Pectis capillipes (Benth.) Hemsl. Biol. Cent. Am. Bot. 2: 225.
1881. Lorentia capillipes Benth. in Oersted, Vidensk. Meddel. 1852:
70. 1853. Pectis erecta Fernald, Proc. Am. Acad. 40: 57. 1904.
Grassy or brushy, often rocky plains and hillsides, sometimes
on saline flats, 200-1,400 m.; Zacapa; Chiquimula; Jalapa; Jutiapa.
Honduras; El Salvador; Nicaragua; Costa Rica(?).
An erect annual or perhaps usually perennial from a slender, woody root, erect,
sparsely branched, the branches ascending or suberect, often reddish or purplish,
scaberulous or almost glabrous, dichotomous; leaves linear or subulate-linear, 1.5-3
cm. long, stiff, strongly ascending, 1-2 mm. broad, acute, with 1-2 rows of glands on
each side and 2-5 pairs of bristles toward the base; heads solitary at the ends of the
branches, on peduncles 3-6 cm. long; involucre 5-6 mm. high; phyllaries 5, oblong or
oblanceolate, ciliate at the rounded or acute apex; rays 5-7 mm. long; disc corollas 5
mm. long; achenes hispidulous; pappus of the disc flowers of 6-8 setiform, hispidulous
squamellae, more or less paleaceous and connate at the base, with 4-8 shorter
squamellae.
Pectis elongata HBK. Nov. Gen. & Sp. 4: 262, t. 392. 1820. P.
oerstediana Rydb. N. Am. Fl. 34: 213. 1916. Comillo (Peten, fide
Lundell). Figure 122.
Moist or rather dry, open or brushy slopes or plains, sometimes
in oak forest or cultivated fields, 300-1,500 m.; Peten; Jalapa;
Jutiapa; Santa Rosa; Chiquimula; Guatemala; Escuintla. Yucatan
Peninsula of Mexico; El Salvador; Honduras; Nicaragua; Costa
Rica; Panama; northern and western South America.
WILLIAMS: FLORA OF GUATEMALA 375
An erect annual, 15-60 cm. high, paniculately much branched, glabrous almost
throughout, the stems somewhat angulate, usually dark red or purplish; leaves linear,
1.5-6 cm. long, 1.5-3 mm. broad, mucronate or bristle-tipped, with 3-10 pairs of
bristles near the base, the glands numerous, in 4-6 irregular rows; heads cymose-
paniculate, very numerous, on slender pedicels 5-20 mm. long; involucre 4-6 mm.
high, glabrous; phyllaries 5, linear, purplish, acuminate, obtusely carinate near the
base; ray flowers 5, the rays 1.5-3 mm. long; disc flowers 3-7; achenes 2-2.5 mm. long,
hirsutulous on the angles or glabrate; pappus bristles of the disc flowers 8-24, of the
ray flowers 5-20, capillary, scabrous, the outer ones 4 mm. long, fulvous.
Called "hierba de talepate" in El Salvador.
The Central American forms of this species known as P.
oerstediana differ from the typical forms of P. elongata only in
minor and perhaps artificial details of the pappus.
Pectis linifolia L. Syst. Nat. ed. 10. 1221. 1759.
Moist thickets, or brushy, rocky slopes, 200-500 m.; Zacapa.
Arizona; Mexico; West Indies; Andean South America.
A slender annual, often 50 cm. high or more, sparsely leafy, sparsely
dichotomous, often glaucescent; leaves linear, 3-6 cm. long, 2-5 mm. broad, sometimes
with a few pairs of bristles at the base; pedicels 1-3 cm. long, terminal and in the
forks of the branches, glabrous or scaberulous; involucre 5-6 mm. high, glabrous or
minutely puberulent; phyllaries 4-5, linear, obtuse, with black, linear, submarginal
glands; ray flowers 4-5, the rays 1 mm. long; disc flowers 2-5, the corollas 2-3 mm.
long; achenes 4-7 mm. long, puberulent or glabrate; pappus of 2-3 stout, subulate,
divaricate, smooth awns 2-2.5 mm. long.
Pectis multiflosculosa (DC.) Sch.-Bip. in Seem. Bot. Voy.
Herald 309. 1856. Lorentea multiflosculosa DC. Prodr. 5: 102. 1836.
Pectis arenaria Benth. Bot. Voy. Sulphur 110. 1846. P. bibracteata
Klatt, Leopoldina 20: 92. 1884. P. grandiflora Klatt, Bull. Soc. Bot.
Belg. 35: 290. 1896.
Along the strand from Acapulco to Peru; at Champerico,
Retalhuleu (Molina, in lit.). Mexico; El Salvador; Honduras;
Nicaragua; Costa Rica; Colombia; Ecuador; Peru.
Prostrate vines, usually in sand along the beaches (of the Pacific Ocean), the
stem thick, reddish, to several meters long, with many short, unilateral, densely
leafed branches; leaves opposite, somewhat fleshy, linear, acute, apiculate, the
margins somewhat revolute, with one or more pairs of setae at the base, mostly 1.5-
3.5 cm. long and 0.1-0.6 cm. broad, usually profusely glandular-punctate above and in
rows along the margin below; inflorescence monocephalus, terminal, borne on a
peduncle 1-3 cm. long, the heads to about 1.5 cm. long; involucre narrowly
campanulate; phyllaries 5, elliptic-oblong, striate, acute or obtuse at the tip and
obscurely flocculose; ray flowers few, the blade narrowly oblong, about 4-5 mm. long;
disc flowers 15-40, the corollas cylindric, about 5 mm. long; achenes multistriate,
376 FIELDIANA: BOTANY, VOLUME 24
black, glabrous or obscurely setose, 6-7 mm. long; pappus of disc flowers of 10-30
bristlelike, ciliolate squamellae, some dilated at the base, about as long as the corolla,
those of the ray flowers fewer and shorter.
Possibly to be found fairly abundantly along Pacific beaches,
although no specimen is found in collections.
Pectis prostrata Cav. Icon. 4: 12, t. 324. 1797. P. multisetosa
Rydb. N. Am. Fl. 34: 198. 1916 (type from Santa Rosa, Heyde &
Lux 4232). Zacatocoche (Zacapa; zacate de cachet).
Dry or moist thickets, open fields, cultivated ground, or
sandbars along streams, 250-1,500 m.; Izabal; El Progreso; Zacapa;
Chiquimula; Santa Rosa; Guatemala; Solola; El Quiche; Suchitepe-
quez; Huehuetenango; Retalhuleu. Florida; New Mexico and
Arizona; Mexico; British Honduras; Honduras; Nicaragua; Costa
Rica; Greater Antilles.
Plants annual, usually much branched, prostrate, often forming small round
mats, the branches 5-20 cm. long, very leafy, the whole plant glabrous or nearly so,
the stems usually short-pilose in 2 lines; leaves oblanceolate-linear, 10-30 mm. long, 2-
4.5 mm. broad, acute, often mucronate, ciliate below the middle, with numerous,
scattered glands; heads axillary and terminal, sessile, usually glomerate and
subtended by several leaves; involucre oblong, 5-6 mm. long, glabrous; phyllaries 5,
oblong or linear-oblong, rounded or truncate at the apex, with a rounded keel
extending to the apex, the glands several near the apex, oblong; ray flowers 5, the
rays oblong, 3 mm. long; disc flowers 4-15, the corollas 2-2.5 mm. long; achenes 3-4
mm. long, pubescent; pappus squamellae of the ray flowers 2, narrowly lanceolate,
accompanied by 1-3 small bristles, the squamellae of the disc flowers 5, unequal.
Easily distinguished among our species by the compact
inflorescences, the keeled phyllaries and the usually compact,
prostrate habit.
Pectis saturejoides (Mill.) Sch.-Bip. in Seem. Bot. Voy.
Herald 309. 1857. Inula saturejoides Miller, Gard. Diet. ed. 8. 1768.
P. canescens var. villosior Coulter, Bot. Gaz. 20: 52. 1895 (type from
Santa Rosa, Heyde & Lux 3401). Colchon de nino (Huehuetenango).
Open, grassy slopes or plains or on brushy hillsides, rarely in
crevices of rocks, sometimes on sand, sea level to 1,700 m.; El
Progreso; Zacapa; Santa Rosa; Escuintla; Chimaltenango; El
Quiche; Huehuetenango; San Marcos. Southern Mexico; Honduras.
Perennial from a thick root, usually much branched, procumbent and often
forming dense mats, the stems 10-20 cm. long, pilose with spreading, white hairs;
leaves linear, 10-20 mm. long, 1.5-2 mm. broad, hirsutulous, the revolute margins
often denticulate, spinose- tipped, with 3-6 pairs of marginal bristles near the base, the
glands scattered, inconspicuous; peduncles solitary in the upper leaf axils, very
slender, 4-7 cm. long, few or numerous; involucre campanula te, 8-10 mm. high;
phyllaries 7-12, linear, subacute, hispidulous; rays oblong, 5-6 mm. long; disc flowers
WILLIAMS: FLORA OF GUATEMALA 377
numerous, the corollas 5 mm. long; achenes 4 mm. long, hirsute; pappus of the disc
flowers of about 20 unequal, hispidulous bristles, the longest 4-5 mm. long; pappus of
the ray flowers none or a mere border.
Pectis uniaristata DC. var. holostemma Gray, Proc. Am.
Acad. 19: 46. 1883. P. dichotoma Klatt, Leopoldina 20: 92. 1888. P.
flava Standl. & Steyerm. Field Mus. Bot. 23: 106. 1944.
Grassy plains or fields, 900-1,000 m.; Jutiapa; El Quiche;
Huehuetenango. Southern Mexico; El Salvador; Honduras; Costa
Rica; Colombia.
Plants annual, erect or spreading, 20 cm. high or less, dichotomously much
branched, glabrous or nearly so, the stems often dark red or purplish; leaves linear or
oblanceolate-linear, 1-3 cm. long, 0.1-0.4 cm. broad, mucronate, with 2-4 pairs of long
bristles near the base, bearing 2-4 rows of conspicuous glands; heads terminal and in
the forks of the branches, forming leafy cymes, the pedicels 1-2 cm. long; involucre 3-
4 mm. high, purplish; phyllaries 5, linear, acute, glabrous, only slightly carinate; ray
flowers 5, the rays 1.5-2 mm. long; disc flowers 3-5; achenes 2 mm. long, hispidulous
or glabrate; pappus a short crown of minute, united or divided squamellae.
Called "comino silvestre" in El Salvador.
In recent studies by Keil, some Honduran and Guatemalan
specimens that seem to differ only in having the minute squamellae
divided have been separated out as Pectis swartziana Less.
(Linnaea 6: 711. 1831) which they may or may not be. We have
written our description to cover both forms. How these two forms
may differ from the typical variety is questionable since the only
difference seems to be the presence of one (or more?) elongated
squamellae in the typical variety. Pectis flava Standl. & Steyerm. is
based on an immature and poor specimen. It seems to differ no way
from this species.
POROPHYLLUM Adanson
Reference: R. Roy Johnson, Monograph of the plant genus
Porophyllum, Univ. Kansas Sci. Bull. 48: 225-267. 1969.
Annual or perennial herbs or low shrubs, usually glabrous; leaves opposite or
alternate, broad to filiform, entire or crenate, usually petiolate, with conspicuous
pellucid glands along the margins and sometimes also on the surfaces; inflorescence
cymose, corymbiform or paniculate, heads medium-sized or rather small, discoid,
homogamous; involucre cylindric to campanulate; phyllaries 5-9, linear to obovate, 1-
seriate, distinct or connate only at the base, usually with 2 vertical rows of glands;
receptacle small, naked; flowers all hermaphrodite and fertile; corolla yellow or
purplish, the limb funnelform or cylindric, shorter or longer than the tube; anthers
rounded at the base; style branches elongate, filiform-subulate, hirtellous; achenes
narrow, often attenuate to the apex, multistriate, generally hispidulous; pappus of
numerous scabrous or hispidulous bristles.
378 FIELDIANA: BOTANY, VOLUME 24
Species all in tropical and warm- temper ate regions of America
and most numerous in Mexico. Johnson recognizes 28 species.
Involucre 20-25 mm. high P. ruderale subsp. macrocephalum.
Involucre 10-14 mm. high P. punctatum.
Porophyllum punctatum (Mill.) Blake, Contr. Gray Herb. 52:
58. 1917. Eupatorium punctatum Mill. Gard. Diet. ed. 8. no. 11. 1768.
P. nummularium DC. Prodr. 5: 649. 1836. P. millspaughii Robinson
in Millsp. Field Mus. Bot. 2: 109. 1900 (type from Yucatan). P.
pittieri Rydb. N. Am. Fl. 34: 188. 1916 (type from Cuesta de
Quililha, near Purulha, Baja Verapaz, H. Pittier 146). P. guatema-
lense Rydb. torn. cit. 190 (type from Nenton, Huehuetenango, E. W.
Nelson 3523). Ruda de monte. Figure 123.
Moist or dry, open or brushy, often rocky slopes or plains,
sometimes in pine or oak forest, 100-2,000 m.; Peten; Baja Verapaz;
Zacapa; Jutiapa; Santa Rosa; El Quiche; Huehuetenango. Southern
Mexico; Honduras; El Salvador.
Plants slender, erect, 2 m. high or less, sometimes weak or even subscandent,
herbaceous, probably perennial, glabrous, sparsely or much branched, the branches
often spreading; leaves mostly opposite, on petioles 1 cm. long or shorter, oval to
elliptic or obovate-oblong, 1-3 cm. long, rounded to subacute at the apex and base,
subcrenate, with 3-5 glands along each margin and a few scattered over the surface;
inflorescence usually cymose, heads few or numerous, the peduncles slender, often
spreading in age; involucre 1 cm. high; phyllaries linear, obtuse to acuminate, with 2
rows of glands, often purplish; corollas mostly dark bronze, sometimes purplish or
yellowish, 1 cm. long, the tube several times as long as the limb; achenes 6-7 mm.
long, hispidulous; pappus stramineous or often purplish, 7 mm. long, the bristles
hispidulous.
Collectors often call this plant a shrub, but it seems more likely
to be a suffrutescent annual or perennial.
Porophyllum ruderale (Jacq.) Cass. subsp. macrocephalum
(DC.) R. R. Johnson, Univ. Kansas Sci. Bull. 48: 223. 1969. P.
macrocephalum DC. Prodr. 5: 648. 1936. Ruda cimarrona; oreja de
monte.
Brushy rocky slopes or plains, most often in sandy soil,
frequently on sandbars along streams, 200-1,200 m.; Zacapa;
Chiquimula; Jutiapa. Arizona; Mexico; El Salvador; Honduras;
Nicaragua; Costa Rica; Panama; north and western South America.
A glabrous, erect, strong-scented annual, 1 m. high or less, simple or usually
rather sparsely branched; leaves alternate, on slender petioles 1-2 cm. long,
membranaceous, broadly oval or obovate, 1-3 cm. long, usually rounded at each end,
subcrenate, glaucescent, the glands oblong, conspicuous, 1 in each sinus along the
WILLIAMS: FLORA OF GUATEMALA 379
margins; heads few, the peduncles 2-7 cm. long, conspicuously thickened above;
involucre at first cylindric, about 2 cm. high; phyllaries usually 5, linear, acute or
obtuse, with 2 rows of linear glands; corollas 12 mm. long, yellowish, the tube many
times longer than the limb; achenes 12-13 mm. long, hispidulous; pappus brown or
brownish, 1 cm. long, the bristles minutely scabrous.
The Maya names of Yucatan are reported as "xpechukil" and
"ukche"; "hierba de venado" (Yucatan).
SCHKUHRIA Roth
References: P. A. Rydberg, N. Am. Fl. 34: 44-46. 1914 (as
Tetracarpum}. Charles B. Reiser, Jr., A revision of the genus
Schkuhria, Ann. Mo. Bot. Gard. 32: 265-278. 1945.
Slender, erect annuals, rarely perennials, paniculately branched; leaves alternate
or the lower ones opposite, pinnately divided into linear or filiform lobes, impressed-
punctate; heads small, discoid or radiate, slender-pedicellate; involucre turbinate, the
phyllaries 4-6 (ours), usually scarious, colored, and petaloid at the apex, carinate
dorsally, often subtended at the base by bractlets; receptacle small; ray flowers 1-3,
the rays ascending or spreading or sometimes wanting; disc flowers usually few
(ours); tube of the corolla glandular, shorter than the campanulate limb, the 5 lobes
short; achenes narrowly obpyramidal, 4-angulate, hispidulous on the angles; pappus
of 8 scarious squamellae, these with callous-thickened bases or a prominent costa,
those on the angles sometimes larger than the others.
Species about six, all American, in tropical and warm-
temperate regions. Only the following is found in Central America.
The generic name is conserved over Sckuhria Moench.
Schkuhria virgata (La Llave & Lex.) DC. Prodr. 5: 654. 1836.
Mieria virgata La Llave & Lex. Nov. Veg. Descr. 2: 9. 1825.
Hopkirkia anthemoidea DC. Prodr. 5: 660. 1836. Tetracarpum
anthemoideum Rydb. N. Am. Fl. 34: 45. 1914. T. guatemalensis
Rydb. I.e. (type from Guatemala, Bernoulli 135). T. virgatum Rydb.
I.e. Schkuhria guatemalensis Standl. & Steyerm. Field Mus. Bot.
22: 319. 1940. S. pinnata (Lam.) O. Kuntze var. virgata Reiser, Ann.
Mo. Bot. Gard. 32: 271. 1945. S. anthemoidea var. guatemalensis
Reiser, torn. cit. 273. Conchalagua; escoba amarga; flor de
mosquito. Figure 124.
Dry rocky thickets or open places, sometimes in oak forest,
often weedy, 600-2,400 m.; Jalapa; Jutiapa; Chiquimula; Guate-
mala; Huehuetenango. El Salvador; Honduras; Nicaragua.
A slender, erect annual, 25-75 cm. high, usually much branched, the branches
erect or ascending, sparsely puberulent or almost glabrous; leaves small, alternate,
with 3-5 linear-filiform lobes, or the upper leaves entire, 2-3 cm. long, impressed-
punctate; heads very numerous, slender-pedicellate, dull yellow; involucre 5 mm.
380 FIELDIANA: BOTANY, VOLUME 24
high, 4 mm. broad; phyllaries obovate, glandular-punctate, glabrous, purplish at the
apex; ray flower 1 or wanting, the ray oblong, 1-2 mm. long; disc flowers 5-7, the
corollas 2 mm. long; achenes 3 mm. long, obpyramidal, densely villous on the angles;
squamellae subequal, lanceolate, aristulate, about equaling the corolla.
TAGETES Linnaeus
Reference: P. A. Rydberg, N. Am. Fl. 34: 148-159. 1915.
Annual or perennial herbs, strong-scented, glabrous or pubescent, with abundant
and usually conspicuous oil glands; leaves opposite, or the upper ones alternate,
dentate or more often pinnate or pinnatifid; heads small or large, pedunculate,
solitary at the ends of the branches or in leafy cymes; involucre cylindric, fusiform,
or campanula te; phyllaries 3-10, narrow, 1-seriate, united almost to the apex, bearing
2 vertical rows of glands, not calyculate at the base; receptacle favose; ray flowers
usually present, few, pistillate, fertile, the ligules often large and showy, usually
yellow, sometimes brown or dark red, rarely whitish; disc flowers few or numerous,
hermaphrodite, fertile; corolla tube cylindric, narrow, usually shorter than the limb;
achenes elongate, clavate or linear, compressed, angulate, or terete, rarely striate,
glabrous or pubescent; pappus squamellae 3-10, very unequal, often more or less
united, 1-2 of them linear, subulate, or setiform, acute, scabrous- hispidulous, the
others much shorter, linear to obovate, usually truncate or rounded at the apex.
Species 35 or perhaps more, on the basis of Rydberg's account
of the genus, all American, mostly in mountains of the tropical
regions, a few extending into the warmer temperate areas. The true
number of species is probably considerably lower, for in North
America there probably are scarcely half as many species as
Rydberg recognized.
Robert T. Neher, in 1957-58, studied and annotated the
material of Tagetes in our herbarium and other major herbaria. The
results of his studies have never been published.
Leaves simple, dentate or serrate but not at all lobate T. lucida.
Leaves pinnatifid.
Inflorescences corymbiform with crowded heads on peduncles shorter than the
involucres.
Pales of the pappus all less than 2 mm. long; ray flowers large and conspicuous,
the blade mostly more than 10 mm. long T. nelsonii.
Pales of the pappus, at least 2 of them, more than 5 mm. long; ray flowers
inconspicuous, the blade 2 mm. long or shorter T. foetidissima.
Inflorescences not corymbiform or if appearing so then the pappus pales much
more than 2 mm. long; heads usually on peduncles longer than the involucres.
Involucres mostly 6-9 mm. long; leaves small with filiform or narrowly linear,
entire divisions T. filifolia.
Involucres more than 10 mm. long, often to 20 mm. or more long; divisions of the
leaves usually serrate to incised.
Involucre 10-12 mm. long and 5-7 mm. in diameter; leaves and stems
puberulent T. nelsonii.
WILLIAMS: FLORA OF GUATEMALA 381
Involucre usually more than 12 mm. long; leaves and stems glabrous.
Segments of the leaves deeply incised, mostly linear or very narrow;
involucres 15-18 mm. long, borne on long slender peduncles, usually only
one on each branch T. subulata.
Segments of the leaves merely serrate, mostly broader than linear.
Ray flowers small, the blades mostly 2 mm. long or shorter; inflorescence
dense, the peduncle usually as short as the involucre.
T. foetidissima.
Ray flowers conspicuous, the blades mostly 4-10 mm. long; inflorescence
open; peduncles all or mostly longer than the involucre.
Ray flowers less than 1 cm. long; involucre 4-5 mm. broad.
T. tenuifolia.
Ray flowers more than 1 cm. long; involucre 6-10 mm. broad.
T. erecta.
Tagetes erecta L. Sp. PI. 887. 1753. T. patula L. I.e. T.
remotiflora Kunze, Linnaea 20: 23. 1847. Flor de muerto; tutz
(Coban, Quecchi); chus (El Quiche, fide Tejada); coxua (Totonica-
pan, fide Tejada); cotzij caminiac (El Quiche, fide Tejada); ixtupug,
sanpuel (Peten, Maya, fide Lundell); subay tus; Kaqi tus (red
ornamental); q'an tus or cot tus (yellow ornamental used in
religeous decoration, Alta Verapaz, Quecchi).
Moist or rather dry thickets or open fields, often a weed in
cultivated or waste ground; grown commonly for ornament; 1,850
m. or less, but often cultivated at even higher elevations; Peten;
Alta Verapaz; Izabal; Chiquimula; Jalapa; Jutiapa; Santa Rosa;
Escuintla; Guatemala; Sacatepequez; Chimaltenango; Huehue-
tenango; El Quiche; Retalhuleu. Mexico; British Honduras; El
Salvador; Honduras; Nicaragua; Costa Rica; doubtfully native
south of Guatemala; often more or less naturalized in other parts of
tropical America and in the Old World tropics.
An erect annual, sometimes 1 m. high but usually much lower and often not
more than 25 cm., glabrous throughout, branched; leaves pinnatisect, the leaflets 11-
17, lanceolate or linear-lanceolate, mostly 1-3 cm. long, acute or acuminate at each
end, serrate, bearing scattered oil glands; heads solitary at the ends of the branches,
the peduncles elongate, more or less dilated and fistulose at the apex; involucre
narrowly campanulate, usually 14-20 mm. long, 7-12 mm. thick; phyllaries 4-8, with
obtuse or acute tips, with usually 2 rows of linear glands; rays 5-8 (often very
numerous in cultivated forms), broadly obovate, bright yellow, 1-2 cm. long; disc
flowers numerous, the corollas 8-10 mm. long, glabrous; achenes black, 7-8 mm. long,
glabrous or pubescent; longer pappus squamellae as much as 10 mm. long, the others
half as long.
The Maya name of Yucatan is reported as "xpuhuc."
Throughout Mexico and Central America the garden marigold is
well known by the name "flor de muerto," whose derivation is not
altogether clear. It might arise from the strong odor of the plant,
382 FIELDIANA: BOTANY, VOLUME 24
although the reason for such an application is not obvious. The
plant is common in most cemeteries, where it is planted because it
grows easily, and the flowers are much used for decorating graves,
as well as for other decorative purposes. More significantly, it is
reported that in Honduras the plant is placed in bath water to
remove the effects of the emanations from dead bodies (ejido de
muerto), and this is perhaps the real explanation of the vernacular
name. The plant doubtless has been in cultivation for many
centuries, and it is not improbable that it had formerly some
important religious significance. The marigold is one of the
commonest garden flowers of all parts of Guatemala, and in some
regions the more primitive forms are apparently native. Many of
the more or less wild plants are derivatives of garden plants, some
of them with double flowers and in no way inferior to garden
plants. Wild plants have bright yellow rays, but in cultivated forms
the rays often are wholly or partly brown or deep red. To T. erecta
we refer all or most Guatemalan material that has been determined
as T. patula L. and T. remotiflora Kunze. Just how these three, if
recognized as distinct species, are to be separated, we do not know.
The characters used in Rydberg's key are rather useless, and do not
in fact exist. It is suspected that T. remotiflora, as interpreted by
Rydberg, is the wild and original type, while his names erecta and
patula pertain to cultivated forms, or those of cultivated origin. At
any rate, better characters will have to be found for their separation
if two or three species are to be recognized.
Intergression with Tagetes tenuiflora Cav. seems to occur, as
we have noted under that species.
Tagetes filifolia Lag. Blench. 1805; Gen & Sp. Nov. 28. 1816.
T. pusilla HBK. Nov. Gen. & Sp. 4: 194. 1820. T. scabra Brandegee,
Zoe 1: 314. 1890 (type from Antigua, the collector unknown). Anis;
anisillo; anisillo de monte; ariis de chucho; r-anis c'o [= rat's anise]
(Alta Verapaz, Quecchi).
Moist, open or brushy, often rocky plains and slopes, common
in pine-oak forest, often a weed in cornfields, 900-2,500 m.; Alta
Verapaz; Zacapa; Santa Rosa; Chiquimula; Jalapa; Jutiapa;
Guatemala; Sacatepequez; Chimaltenango; Huehuetenango. Mex-
ico; El Salvador; Honduras; Nicaragua; Costa Rica; Panama.
A glabrous, erect annual, 8-50 cm. high, usually much branched, often diffuse;
leaves small, mostly opposite, pinnatifid or bipinnatifid, the divisions linear-filiform,
sometimes scaberulous; heads small, very numerous, slender-pedicellate or almost
sessile; involucre fusiform, 6-9 mm. high, often 5-angulate at the base; phyllaries 5,
WILLIAMS: FLORA OF GUATEMALA 383
rounded and mucronate at the apex, with 2 vertical rows of numerous small glands;
ray flowers 1-3 or sometimes none, the rays oval, 1.5 mm. long, pale yellow or
whitish; disc corollas 3 mm. long, glabrous; achenes 4.5 mm. long, striate; 2 of the
pappus squamellae setiform, 3 mm. long, hispidulous, the other 3 oblong, 1 mm. long.
Known in El Salvador as "anisillo" and "anis de monte." The
plant has a strong odor similar to that of anise or licorice.
Guatemalan material has been reported as T. micrantha Cav., a
more northern species that probably is distinct, but has not been
found in Guatemala.
Tagetes foetidissima DC. Prodr. 5: 645. 1836. Flor de muerto.
Moist, open banks or at the edge of forest, often in open or
dense forest of pine, oak, or Cupressus, sometimes on limestone,
1,800-3,400 m.; Jalapa; Chimaltenango; Solola; Huehuetenango; El
Quiche; Quezaltenango; San Marcos. Southern Mexico; Costa Rica.
A glabrous, erect or spreading annual, 1 m. high, or more often, half as tall (at
3,400 m. in Guatemala as small as 5 cm. tall), branched above, the stems often tinged
with dark red or purple; leaves alternate or the lower ones opposite, 3-9 cm. long, the
leaflets 11-29, the upper ones confluent, oblong or lance-oblong, obtuse, incised-
serrate; heads densely cymose-paniculate, numerous, slender-pedunculate; involucre
subcylindric, acute at the base, about 15 mm. long and 3-4 mm. thick; phyllaries 5,
dark reddish, deltoid at the apex, with 2 vertical rows of oblong glands; rays usually
5, pale yellow, inconspicuous, 2 mm. long or shorter; disc flowers about 7, the corollas
greenish yellow, 7 mm. long, glabrous below, the limb viscid-pilose; achenes hirsute, 6
mm. long; 2 of the pappus squamellae awnlike, 7 mm. long, the other 3 oblong, 1 mm.
long.
It is possible that T. foetidissima is not distinguishable from T.
multiflora HBK. (1820), a common species in Andean South
America. However, Mr. Neher, who studied but did not publish on
this genus, maintained them separately.
Tagetes lucida Cav. Ic. 3: 33. t. 264. 1794. Pericon (sometimes
written Hipericon); liya (Totonicapan, fide Tejada); iya, jolomocox,
uca (El Quiche, fide Tejada); hierba de San Juan (Quezaltenango,
fide Tejada).
Mostly in open grassy fields, frequently in oak forest,
sometimes on dry rocky hillsides, 1,000-2,000 m.; Peten; Jalapa;
Guatemala; Sacatepequez; Chimaltenango; El Quiche; Huehue-
tenango; San Marcos. Mexico; El Salvador; Honduras.
A glabrous, erect perennial commonly 30-75 cm. high, strong-scented, arising
from a short, thick, woody base, cymosely branched above; leaves opposite, sessile,
linear or narrowly oblong, 5-10 cm. long, obtuse or acute at each end, densely serrate,
with numerous small, scattered glands; heads small, in dense or open, flat-topped
cymes; involucre cylindric, 9-10 mm. long; phyllaries 5-7, subulate at the apex, with
384 FIELDIANA: BOTANY, VOLUME 24
numerous small scattered glands; rays commonly 3, flabelliform, 3 mm. long,
truncate; disc flowers 5-7, the corollas 5-6 mm. long; achenes 6-7 mm. long, striate;
pappus squamellae 5-6, two of them setiform, 3 mm. long, the others one-third as
long, oblong, obtuse.
Sometimes called "periquillo" in Mexico. The name "pericon" is
well established in Guatemala, and there are even villages bearing
the name El Pericon in El Quiche and Huehuetenango. Throughout
its range the plant has a high reputation as a remedy for digestive
disturbances. Many of the foreign residents of Guatemala praise it
quite as highly as the native people, so that there is every reason for
believing that it does have definite remedial properties. Bunches of
the dried stems and leaves may be found in every Guatemalan
market. They are used to prepare the tea that is taken whenever
there is a serious upset of the stomach or intestines. The plants
grow abundantly during the wet months but wither as soon as the
rains cease. It is unusual to see a single flowering plant during most
of the dry season. The Indians who gather the plants insist that
they must be cut before June 24, Dia de San Juan, because after
that day the devil rolls in them. In Mexico the powdered leaves are
often used to repel or destroy mosquitoes, hen lice, and other
insects, also as a remedy for malaria and various other diseases.
Guatemalan material has been referred to T. florida Sweet and T.
schiedeana Less., Mexican species that probably do not differ in
any appreciable or constant character from T. lucida. It is worthy
of note that presumably authentic material of T. lucida, received
from the Botanic Garden of Madrid, from the Cavanilles herbarium,
includes both T. lucida and T. schiedeana.
Tagetes nelsonii Greenm. Proc. Am. Acad. 39: 117. 1903;
Contr. Gray Herb. n. s. 25: 117. 1903. T. sororia Standl. & Steyerm.
Field Mus. Bot. 23: 146. 1944 (type from Quezaltenango, Standley
85228). Figure 125.
Moist or dry forests and open slopes, perhaps sometimes
cultivated by the highland Indians, 2,500-3,300 m. but somewhat
lower in Chiapas; Totonicapan; Huehuetenango; Quezaltenango;
San Marcos.
An erect herb, 1 m. high or often more, perennial, sometimes suffruticose at the
base, branched, the stems slender, glabrous to sparsely short-pilose or villosulous;
leaves opposite or the upper ones alternate, pinnate, the leaflets 5-15, lanceolate,
oblanceolate or narrowly elliptic (l-)3.5-7 cm. long, mostly acute or acuminate at
each end, coarsely serrate, glabrate to puberulent above, mostly densely pilosulous
below, the glands numerous, small, scattered; heads few or numerous, the peduncles
short and becoming elongate, naked or often bearing a few small pinnatisect bracts;
WILLIAMS: FLORA OF GUATEMALA 385
involucre narrowly campanulate, 5-8-dentate, 10-12 mm. long, glabrous, 5-6 mm.
broad, obtuse at the base; phyllaries 5-7, deltoid and obtuse at the apex, with 2
vertical rows of conspicuous, short, oblong glands; rays yellow, oblong, to 15 mm.
long; disc corollas 6 mm. long; puberulent; achenes black, about 5 mm. long, almost
glabrous; 2 of the pappus scales subulate, 3 mm. long, the other three oblong, scarcely
1 mm. long.
Perhaps the good forms of this are the most attractive of the
Tagetes of Guatemala, those of the high Cuchumatanes being
especially fine. The plants are probably not palatable to sheep and
goats for they apparently are not eaten by these animals.
Tagetes subulata Cervantes ex La Llave & Lex. Nov. Veg.
Descr. 1: 31. 1824. T. multiseta DC. Prodr. 5: 645. 1836. Flor de
muerto.
Moist fields or open banks, often on dry, brushy slopes or in
pine-oak forest, 800-1,900 m., rarely near sea level (in Izabal;
perhaps introduced); Izabal; Zacapa; Chiquimula; Jalapa; Jutiapa;
Santa Rosa; Guatemala; Sacatepequez; Chimaltenango; Huehue-
tenango. Mexico; El Salvador; Honduras; Costa Rica; Colombia.
A slender, glabrous, erect annual, 20-50 cm. high, usually corymbosely branched
above; leaves opposite and alternate, 2-3 cm. long, pinnate, the leaflets 5-13,
pectinately dissected into linear bristle- tipped divisions, the glands few, large; heads
usually numerous, each on a very long and slender, naked peduncle; involucre
cylindric-fusiform, 15-18 mm. long, 4-5 mm. broad; phyllaries 4-5, deltoid at the apex,
with irregular rows of linear glands; rays 3-5, suborbicular, 2 mm. long, yellow,
inconspicuous; disc flowers 4-8, the corollas 8 mm. long; achenes 5 mm. long,
scabrous on the angles; 1-2 of the pappus squamellae subulate, 10 mm. long, the
other 3-4 linear, truncate, 7-8 mm. long.
Easily distinguished among the Central American species by
the much dissected leaves and the elongated, slender peduncles.
Tagetes tenuifolia Cav. Ic. 2: 54, t. 169. 1793. IT. peduncularis
Cav. Descr. 201. 1802. Flor de muerto.
Moist, open or brushy plains or hillsides, frequent in sand or
gravel along streams, sometimes in oak forests, often a weed in
cultivated fields, 300-2,700 m.; Alta Verapaz; Baja Verapaz;
Chiquimula; Jalapa; Jutiapa; Santa Rosa; Escuintla; Guatemala;
Sacatepequez; Chimaltenango; Solola; El Quiche; Huehuetenango;
Quezaltenango; San Marcos. Mexico; El Salvador; Honduras;
Nicaragua; Costa Rica; perhaps South America.
A slender, erect, glabrous annual, commonly 30-75 cm. high, branched above;
leaves opposite or the upper ones alternate, pinnate, the leaflets 13-23, linear to
linear-lanceolate, 1-2 cm. long, serrate, often sharply so, with a row of oil glands on
each side; heads few to usually numerous, cymose, long-pedunculate; involucre
386 FIELDIANA: BOTANY, VOLUME 24
tubular-campanulate, 11-22 mm. long, 4-6 mm. broad; phyllaries about 5, with
triangular, acute tips, with usually elongate lateral glands; rays 4-8 mm. long, yellow,
retuse; disc corollas puberulent, 7 mm. long; achenes about 7 mm. long, strigose; 1-2
of the pappus squamellae subulate, about 6 mm. long, the others half as long, linear,
truncate.
Tagetes tenuifolia belongs in the complex with T. erecta, the
most perplexing group in the genus in our region. The species
probably intergrades with what we consider to be the more
primitive forms of T. erecta, and it is possibly this intergression
which makes their determination so difficult. The many specimens
from our region, determined as T. microglossa Benth., by Mr.
Neher, seem to belong in this complex also.
EXCLUDED SPECIES
TAGETES HETEROCARPHA Rydb. N. Am. Fl. 34: 155. 1914.
The type, Pringle 2488, and all other Mexican specimens seen,
are from the state of Jalisco, Mexico. Mr. Neher has determined
Carlson 850 from the Department of Santa Ana, El Salvador, as
this species, apparently correctly so. Normally, we would include in
this flora species known from Mexico and El Salvador, but in this
case we have not done so, since only one collection is recorded from
Central America, and the disjunct distribution is most unusual.
TRIBE VII. ANTHEMIDEAE
By DOROTHY L. NASH
Annual or perennial herbs or sometimes shrubs, often strong-scented; leaves
alternate (in ours), the blades simple and entire or dentate, or often incised or
dissected, frequently 1-3-times pinnatisect with linear or filiform segments;
inflorescences commonly more or less corymbose but sometimes spiciform, race-
miform, or paniculiform, or of solitary heads; heads heterogamous and radiate or
rarely homogamous and discoid; receptacle paleaceous, naked, or pilose; involucres
mostly hemispheric or campanulate; phyllaries 2-multiseriate, at least the inner ones
more or less scarious; ray flowers, when present, commonly conspicuously ligulate,
rarely narrowly cylindric and inconspicuous, the ligules often white but may be
variously colored; disc corollas tubular, usually fertile; anthers usually obtuse or
entire at the base, rarely subsagittate; style branches truncate and penicillate;
achenes small, often angular and truncate at the apex; pappus none or reduced to a
very short crown of inconspicuous scales.
The tribe of more than 50 genera is represented in Guatemala
by only four genera (six species); only one of these is native, the
others naturalized species from other areas, chiefly temperate and
arctic regions of the Old World.
NASH: FLORA OF GUATEMALA 387
Receptacle paleaceous Achillea.
Receptacle naked or pilose.
Inflorescence (in ours) a dense, narrow, elongated panicle; ray flowers inconspic-
uous, with narrow, cylindric corollas Artemisia.
Inflorescence corymbose or of solitary heads; ray flowers (when present)
conspicuously ligulate.
Achenes equally or irregularly 5-10-costate Chrysanthemum.
Achenes 3-5-costate on the inner surface, ecostate dorsally Jdatricaria.
ACHILLEA Linneaus
Reference: P. A. Rydberg, N. Am. Fl. 34: 219-227. 1916.
Perennial herbs, strong-scented, usually villous, with leafy stems; leaves
alternate, serrate to tripinnatifid, often appearing plumelike; inflorescences
corymbose or corymbiform-paniculate; heads few to many, usually radiate;
involucres campanulate to hemispheric; phyllaries 3-4-seriate, graduate; receptacle
conic or convex; pales membranaceous; ray flowers 5-12, pistillate and fertile, the
ligules short and broad, spreading, white or pink; disc flowers numerous,
hermaphrodite, fertile, the corollas yellowish, the limb 5-cleft; anthers entire at the
base, with an ovate, obtuse apex; style branches of the disc flowers with truncate,
fimbriate tips; achenes oblong or obovate, obcompressed, glabrous; pappus none.
About 100 species, mostly in temperate and arctic regions of
the Old World, a single one extending into Central and South
America (an introduced weed). While Rydberg reports 24 species
from North America, the valid species are probably a fraction of
that number.
Achillea millefolium L. Sp. PL 899. 1753. A. lanulosa Nutt.
Journ. Acad. Phila. 7: 36. 1834. A. pecten-veneris Pollard, Bull.
Torrey Bot. Club 26: 371. 1899. Alhucema (Huehuetenango). Figure
126.
Open, grassy hillsides, sometimes in rocky places, sometimes a
weed along roadsides or about cultivated ground, occasionally
cultivated for ornament or for medicinal use, 1,800-2,500 m.;
Chimaltenango; Huehuetenango; Jalapa. Mexico; Honduras; Costa
Rica; South America. Native of Europe and Asia, widely natural-
ized as a weed in temperate North America and perhaps in part of
native origin.
Strong-scented, perennial herbs from creeping rootstocks, the stems erect, stout,
30-50 cm. high, simple or branched, more or less villous; leaves petiolate, the blades
10-20 cm. long, plumelike, villous 2-3-times pinnatifid, in outline oblanceolate or
linear, the ultimate divisions very numerous and small, linear or lance-linear,
spinu lose- tipped; heads numerous, disposed in dense, corymbiform panicles; in-
volucres 4-5 mm. high; phyllaries 4-seriate, the outer ones ovate, obtuse, the inner
ones oblong, the margins pale brown; ray flowers usually 5, the ligules white, rarely
388 FIELDIANA: BOTANY, VOLUME 24
pink, orbicular, 2.5-3 mm. long; disc flowers 20-25, the corollas 2.5-3 mm. long;
achenes about 2 mm. long.
In Yucatan, where the plants are much cultivated, they are
known by the names "alcanfor" and "mil en rama." Although
widespread in Jalapa and Huehuetenango, it is probable that all the
Guatemalan plants have been introduced accidentally or are
escapes from cultivation.
ARTEMISIA Linneaus
Reference: P. A. Rydberg, N. Am. Fl. 34: 244-285. 1916.
Annual or perennial herbs or sometimes shrubs, usually bitter-aromatic; leaves
alternate, usually dissected, lobate, or dentate, often tomentose; inflorescences
spiciform, racemiform or paniculiform ; heads disciform, discoid or radiate, nutant
when young; involucres campanulate to hemispheric; phyllaries 2-4-seriate, at least
the inner ones somewhat scarious; receptacle flat to hemispheric, naked or pilose;
marginal or ray flowers (when present) usually pistillate and fertile, the corollas
cylindric, the limb commonly 2-3-dentate; styles more or less exserted, the branches
linear-filiform and somewhat flattened, truncate, penicillate; disc flowers hermaphro-
dite, fertile or sterile, their corollas tubular-campanulate or funnelform, the limb 5-
dentate; anthers longer than the short filaments, obtuse or subsagittate at the base,
the connective forming a lanceolate or subulate tip; styles included or short-exserted;
achenes ellipsoid, terete, usually glabrous; pappus none or reduced to a very short
crown.
One of the largest genera of Compositae, with about 200
species, mostly in temperate regions and in the northern hemis-
phere, in tropical regions confined to the mountains, and with only
one in Guatemala. A few species of the genus are sometimes planted
for ornament. The dried leaves of A. dracunculus L., "tarragon," are
sometimes employed for flavoring beverages or food. A. absinthium
L., of the Old World, is a source of the well-known beverage,
absinthe.
Artemisia mexicana Willd. ex Spreng. Syst. 3: 490. 1826.
Incienso. Figure 127.
Dry, often rocky fields and slopes, 2,000-2,600 m.; Huehue-
tenango; Jalapa; Totonicapan; sometimes planted in gardens at
high or low elevations. Southwestern United States; Mexico.
Erect perennial herbs, the stems usually less than 1 m. high, simple or branched,
striate, floccose- tomentose, at least when young; leaves numerous, often crowded on
the stem, mostly 5-10 cm. long, glabrate above but when young floccose-tomentose,
densely white-tomentose beneath, the cauline leaves ternately or pinnately lobate
into linear or lanceolate segments 2-4 mm. wide, or the upper leaves entire, the basal
leaves usually with more numerous lobes; heads numerous, inconspicuously radiate,
greenish, disposed in a dense, narrow, elongated panicle; involucres campanulate to
NASH: FLORA OF GUATEMALA 389
hemispheric, about 3 mm. high; phyllaries about 3-seriate, sparsely arachnoid-
tomentose, the outermost ovate, acute or subacute, the inner ones oval, obtuse,
broadly scarious-margined; ray flowers 5-10, the corollas 1-1.5 mm. long; disc flowers
7-15, yellowish; achenes about 1 mm. long.
It is somewhat uncertain whether this plant is native in
Guatemala, but it probably is in Huehuetenango. Elsewhere it has
the appearance of having escaped from gardens, as it is persistent
about old dwelling sites. It is grown for medicinal purposes, used as
a domestic remedy for colic and other afflictions of the digestive
organs. A. vulgaris 1., "wormwood," or in Spanish, "ajenjo," native
of Europe, is also sometimes planted in Guatemalan gardens for
medicinal use. It is similar to A. mexicana but has thinner, broader
leaves with broader, usually dentate or lobate leaf segments.
CHRYSANTHEMUM Linneaus
Annual or perennial herbs, glabrous or laxly pubescent; leaves alternate, simple
and entire or dentate, often incised or dissected; heads heterogamous, radiate, long-
pedunculate at the ends of the branches, or sometimes disposed in corymbs;
involucres usually hemispheric; phyllaries multiseriate, imbricate, broad, appressed,
the innermost scarious at the apex, the outer ones shorter, often with scarious or
fuscous margins; receptacle flat to convex or hemispheric; ray flowers uniseriate,
fertile or sometimes partly sterile, the ligules white or yellow, rarely pink or purple,
spreading, entire or dentate; disc flowers hermaphrodite, fertile or sometimes partly
sterile, the corolla regular, the tube terete or 2-winged, the limb little dilated,
shallowly 4-5-cleft; anthers obtuse and entire at the base; styles of the hermaphrodite
flowers truncate, penicillate; achenes subterete or variously angulate, equally or
irregularly 5- 10-costate, glabrous or sometimes glandular, apex truncate; pappus none
or some or all with a pappus consisting of a crown of few or numerous, small scales.
About 200 species, mostly in temperate regions of the Old
World; only a very few species native in America, and these chiefly
in arctic regions. Although several species are widely grown in
Guatemalan gardens, only two have become naturalized and are
therefore treated here.
Leaves crenate, dentate, or pinnatifid; heads solitary or few, the disc 1-2 cm. broad.
C. leucanthemum.
Leaves mostly bipinnatifid; heads disposed in corymbs, the disc about 0.7 cm. broad.
C. parthenium.
Chrysanthemym leucanthemum L. Sp. PI. 888. 1753. Marga-
rita.
Native of Europe and Asia, often grown in temperate and
tropical regions for its handsome flowers; extensively naturalized as
a weed over large areas of the United States; a common garden
390 FIELDIANA: BOTANY, VOLUME 24
plant of Guatemala, more or less naturalized in some localities,
particularly in hedgerows.
Erect perennials, glabrous or nearly so, simple or sparsely branched, commonly
30-50 cm. high; lowest leaves incised or pinnatifid, the upper ones oblong or linear,
dentate, clasping; heads few or solitary; disc about 1.5 cm. broad; ray flowers in a
single series, the ligules large, white.
One of the popular garden plants of Guatemala. The flowers
are often sold in the markets to be used for decorating houses and
churches. In the high mountains of Costa Rica it has become
thoroughly naturalized, but elsewhere in Central America it shows
less tendency to spread from cultivation.
Chrysanthemum parthenium (L.) Pers. Syn. PL 2: 462. 1807.
Matricaria parthenium L. Sp. PL 890. 1753. Aphanostephus
pinulensis Coulter ex Donn.-Sm. Enum. PI. Guat. 2: 35. 1891,
nomen; Bot. Gaz. 16: 98. 1891 (type from Pinula, Guatemala, J. D.
Smith 2407). Chusita; artemisa; margarita; altamisa. Figure 128.
A native of Europe but planted extensively in other regions and
often naturalized; a frequent plant in Guatemalan gardens at
almost all elevations and sometimes more or less naturalized in
waste ground, hedgerows, roadsides, etc.
Strong-scented perennials, rather finely pubescent almost throughout, the stems
erect, often much branched, to about 50 cm. high; leaves thin, slender-petiolate,
bipinnatifid, the segments or lobes small, obtuse; inflorescences corymbose; heads
often numerous; disc about 7 mm. broad; ray flowers uniseriate, the ligules short,
white.
The English name is "feverfew." In Central America, as well as
in remote regions, the plants are often employed in domestic
medicine.
MATRICARIA Linneaus
Reference: P. A. Rydberg, Chamomilla and Matricaria, N. Am.
Fl. 34: 229-234. 1916.
Annual or perennial herbs, often strong-scented, pubescent or glabrate; leaves
alternate, 1-3-times pinnatisect, the segments linear or fililorm; heads usually
heterogamous and radiate, rarely homogamous and discoid, pedunculate at the ends
of the branches, solitary or disposed in corymbs; involucres hemispheric; phyllaries 2-
3-seriate (in ours), imbricate, appressed, often scarious-marginate, graduate; recep-
tacle usually conic, sometimes hemispheric, naked; ray flowers fertile or sterile, the
ligules white, spreading, subentire; disc flowers hermaphrodite, all or mostly fertile,
the corollas regular, yellow or greenish yellow, the limb little ampliate, shallowly 4-5-
cleft; anthers obtuse and entire at the base; style branches of the disc flowers
NASH: FLORA OF GUATEMALA 391
truncate at the apex; achenes oblong, usually incurved, truncate, 3-5-costate on the
inner face; pappus none or reduced to a short crown.
About 50 species, almost all in the Old World, a very few native
in North America. Two are treated here, both naturalized, one from
the Mediterranean region, the other from the western United
States.
Heads radiate; pappus a short crown M. courrantiana.
Heads discoid; pappus obsolete M. matricarioides.
Matricaria courrantiana DC. Prodr. 6: 52. 1837. Chamomilla
courrantiana C. Koch, Linnaea 24: 338. 1851. Manzanilla. Figure
129.
Native of the Mediterranean region, but naturalized at many
places in America from the southwestern United States south
through South America; in Guatemala much planted in gardens to
3,900 m., frequently naturalized in fields and cultivated or waste
ground, 1,300-2,500 m.; Alta Verapaz; Chimaltenango; Jalapa;
Quezaltenango; Sacatepequez; San Marcos; Zacapa.
Annuals, with a rather sweet odor, usually much branched, 15-40 cm. high,
glabrous or nearly so; leaves 3-6 cm. long, bipinnatifid, the segments filiform; heads
radiate, solitary at the ends of the branches; involucres about 2.5 mm. high, 6-8 mm.
broad; phyllaries biseriate, oblong or elliptic, rounded at the apex, with broad,
scarious margins; receptacle conic, acute; ray flowers 10-15, the ligules white, about 6
mm. long; disc corollas yellow, about 2 mm. long; achenes ellipsoid, 5-costate on the
inner side; pappus a membranous, white, erose or fimbriate crown.
One of the best known plants of Guatemala and other parts of
Central America and the West Indies, grown in almost every
garden, and perhaps the most used of all domestic remedies, with
fantastic properties sometimes ascribed to the plant. Small bunches
of the dried or fresh plants are on sale in practically every market,
and a tea made from it is administered for almost every kind of
affliction. Closely related plants are official in some of the
pharmocopoeias of Europe, used as mild tonics and for various
other purposes. The closely related M. chamomilla L., distinguished
from M. courrantiana by the absence of pappus, has been reported
from Central America but it is questionable whether it is grown
there.
Matricaria matricarioides (Less.) Porter, Mem. Torrey Bot.
Club 5: 341. 1894. Artemisia matricarioides Less. Linnaea 6: 210.
1831. Chamomilla suaveolens Rydb. N. Am. Fl. 34: 232. 1916, not
M. suaveolens L. (1755). M. discoidea DC. Prodr. 6: 50. 1837.
392 FIELDIANA: BOTANY, VOLUME 24
Weeds near houses, probably escaped from cultivation, 1,800-
2,400 m., San Marcos. United States; Mexico; Europe.
Annuals, with an odor of pineapple when bruised, the stems branching, leafy,
glabrous; leaves mostly 2-5 cm. long, 1-3-pinnately parted, the ultimate segments
short, linear to filiform; heads discoid, on short peduncles; receptacle conic, acute;
phyllaries essentially oval, the broad margins scarious; disc corollas greenish yellow,
the limb 4-dentate; achenes obscurely costate; pappus obsolete.
TRIBE VIII. SENECIONEAE
By Louis O. WILLIAMS
Herbs, shrubs or weak trees; leaves alternate, sessile, usually petiolate,
occasionally peltate; the inflorescence from a single, nodding head to multicapitate,
compound inflorescences; heads homogamous or heterogamous, the disc flowers
rarely heterogamous (in 2 species of Senecio), or usually if the ray flowers lacking,
then the disc flowers all perfect; receptacle naked; involucre usually consisting of a
relatively consistent number of phyllaries in a given species, the phyllaries uniseriate
or sub-biseriate and mostly subequal in length, the involucre often subtended by
bracts and then said to be calyculate; achenes relatively short, ridged or smooth,
most often glabrous; pappus (in all of ours) setose, the setae arranged in one or more
than one series; disc corollas tubular or cylindric below but occasionally enlarged at
the base, the throat slightly to greatly expanded, the lobes very short and dentiform
or almost as long as the throat (in PsacaUwn); anthers ecaudate at the base and
usually appendaged at the apex; styles of the disc flowers usually truncate and often
penicillate, rarely narrowed and terminated by triangular, acute or acuminate,
dorsally hispidulous appendages (Senecio chenopodioides).
One travelling through Guatemala would not suspect that
there are a relatively large number of species present, as members of
the tribe are only occasionally conspicuous in the make-up of the
vegetation.
Most species occupy quite definite niches ecologically. The
greatest number are found in wet forest situations well above the
hot tropical lowlands. These are in flower most often from
December to July, but are occasional throughout the year. Species
that inhabit the open slopes of the highlands or the high volcanoes
are usually in flower toward the end of the rainy season, November
to February, the normal flowering season for highland Compositae.
The tribe is abundant in Mexico. A surprising number,
especially of Senecio, are found in Guatemala, with a large portion
of these endemic to the Guatemalan highlands and to the Central
American floristic region, which extends as far north as Veracruz.
Southeastward in Central America the tribe diminishes in species
and individuals, to increase slightly in Costa Rica with elements
related to the Andean kinds.
NICOLSON: FLORA OF GUATEMALA 393
Several generic segregates, especially of the senecios of the
Central American floristic region, have been proposed, and all have
been rejected in this flora. Psacalium, an allied genus, has been
accepted, as it obviously should be.
Several genera, traditionally placed in the Senecioneae, have
been placed in other tribes in this flora: Liabum to Vernonieae;
Schistocarpha, Neurolaena and Alepidocline to the Heliantheae.
Corolla throat divided almost to its base into narrow lobes; basal leaves peltate.
Psacalium.
Corolla throat divided les than half way to its base; basal leaves peltate only in two
species of Senecio.
Pistillate flowers in several series, without rays; plants herbaceous; corollas
greenish to pink or purple Erichtites.
Pistillate flowers in a single series or none; rays present or none.
Style branches with long subulate, hirtellous appendages; escapes or cultivated
plants Gynura.
Style branches truncate or with short appendages.
Plants acaulescent perennial herbs with long white rays Werneria.
Plants caulescent herbs, shrubs or trees, never with long white rays.
Heads pink, red or orange; small annual herbs Emilia.
Heads yellow, white, orange-yellow or reddish yellow; perennial herbs,
shrubs or vines, or trees Senecio.
EMILIA Cassini
By DAN H. NICOLSON
Diffuse annual herbs; leaves alternate, the margins subentire to lyrate, often
dimorphic, the basal leaves oblanceolate to orbicular and petiolate or tapering to the
base, the upper leaves sagittate and broadest at the base, sessile or semiamplexicaul;
inflorescence corymbose, erect, terminal, with few capitula; phyllaries few, uniseriate,
united below, reflexing in maturity; receptacle naked, becoming convex and weakly
rugose; florets discoid, bisexual, 5-merous; corollas slender, orange through red to
lilac; stamens with filaments thickened above, the apex of the thecae appendaged;
style branches elongate, each with a penicillate appendage with two strigose lines;
pappus abundant, deciduous, scabridulous; achenes 5-angled, each angle with an
internally puberulent groove, sometimes with intermediate ribs.
Florets equalling the narrowly cylindric involucre; corolla pale purple; corolla lobes
0.5-0.7 mm. long; lower leaves lyrate E. sonchifolia.
Florets clearly exceeding the broadly cylindric to urceolate involucre; corolla red to
orange; corolla lobes longer than 1 mm.; lower leaves entire to dentate.
Florets 1.5 times as long as the involucre; corolla light to dark red (no yellow
overtone); corolla lobes 1.1-1.5 mm. long; involucre twice as long as broad;
lower leaves strongly dentate E. fosbergii.
Florets two times as long as involucre; corolla yellowish red; corolla lobes 1.7-2
mm. long; involucre as long as broad; lower leaves entire to weakly dentate.
E. coccinea.
394 FIELDIANA: BOTANY, VOLUME 24
A paleotropical genus of 45 species of which two have become
common weeds in the neotropics and one escapes from occasional
cultivation.
Emilia coccinea (Sims) G. Don in Sweet's Hort. Brit., ed. 3, p.
382. 1839. Cacalia coccinea Sims in Curtis Bot. Mag. 16: t. 564.
1802. Cacalia sagittata Willd., Sp. PL 3(3): 1731. 1803, non Vahl
(1794). Emilia flammea Cass. in Cuvier, Diet. Sci. Nat. 14: 406.
1819. Emilia sagittata DC., Prodr. 6: 302. 1838 (the latter two
binomials based on Cacalia sagittata Willd., non Vahl).
A garden ornamental not yet collected in Guatemala but
included because of the probability of its occurence in disturbed
areas, 1,000-2,000 m. Probably of East African origin: Mexico;
Honduras; El Salvador; Costa Rica; West Indies; Colombia;
Venezuela; Brazil.
Lower leaves entire to shallowly dentate, 9-12 cm. long, 2-4 cm. broad, peduncles
15-30 cm., terminating in 2-3 capitula, each capitulum on a 1-3 cm. pedicel; involucre
urceolate, 0.5 cm. by 0.5 cm.; phyllaries 10-13; florets 50-60; corollas red and tending
toward yellow (scarlet, orange), 7.5-9.5 mm. long, much exceeding the involucre (5-6
mm.); corolla lobes elongate (1.6-2 mm.); stamen with thickened portion of filament
0.5 mm. long, thecae 1.7 mm., anther appendages 0.4 mm.; achenes 3 mm. long, 0.5
mm. broad, intermediate and paler ribs present between grooves.
In the neotropics the binomial E. sagittata has often been
applied to specimens of E. coccinea and E. fosbergii. The original
Cacalia sagittata Vahl (1794) was an illegitimate renaming of
Hieracium javanicum N. Burm. (1768), included in synonymy (Art.
63). Willdenow (1803) redefined Vahl's taxon by excluding Vahl's
synonymy, but created a new taxon with a later homonyn (Art. 64).
Emilia fosbergii Nicolson, Phytologia 32: 33. 1975. Emilia
javanica sensu auctt., non (N. Burm.) C. B. Robinson: Fosberg,
Occas. Papers Bishop Mus. 23: 136. 1966; Vuilleumier, J. Arnold
Arb. 50: 122. 1969; Adams, Fl. PL Jamaica 757. 1972. Emilia
coccinea sensu auctt., non (Sims) G. Don: Britton & Wilson, Sci.
Surv. Porto Rico & Virgin Isl. 6: 321. 1926; Baldwin, Bull. Torrey
Bot. Club 73: 18. 1946; Gooding et al., Fl. Barbados 436. 1965 (fig.
27, mislabelled E. sonchifolia, is this species). Emilia sagittata
sensu auctt., pro parte, non DC.; Standley, Field Mus. Bot. 18:
1454. 1938; Alain, Fl. Cuba 5: 239. 1962.
Common weed of wetter and disturbed habitats, 100-1,500 m.;
Peten, Chiquimula, Alta Verapaz. Possibly of hybrid origin in
western Pacific islands; Florida, West Indies and south through
Brazil.
WILLIAMS: FLORA OF GUATEMALA 395
Lower leaves coarsely dentate, 8-13 cm. long, 3-5 cm. broad; peduncles 15-30 cm.,
terminating in 2-4 capitula, each capitulum on a 2-5 cm. pedicel; involucre 11-15 mm.
long, 2-3 mm. broad; corolla lobes 1.1-1.4 long; stamen with thickened portion of
filament 0.5 mm. long, thecae 1.5-1.6 mm. long, anther appendages 0.3-0.4 mm. long;
achenes 4 mm. long, 0.6 mm. broad, with intermediate ribs between grooves not
differentiated by color.
This species has been consistently misidentified as an African
taxon (E. sagittata, E. coccinea) or a southeast Asian taxon (E.
javanica). It does not occur in either area, although it may be
related to E. javanica.
Emilia sonchifolia (L.) DC. ex Wight, Contrib. Bot. India 24.
1834; DC., Prodr. 6: 302. 1838. Cacalia sonchifolia L., Sp. PL 834.
1753. Figure 130.
Common weed in dry, open or disturbed habitats, 0-500 m.;
Izabal. From Asia, now pantropical; Florida; Mexico; West Indies
and south into Brazil.
Basal leaves petiolate, dentate, obovate to spatulate, often with a pair of lobes
below blade, the central cauline leaves stongly lyrate, the upper leaves sessile to
clasping (subsagittate), dentate to lyrate and widest at base; peduncles 5-15 cm.,
terminating in 2-5 capitula, each capitulum on a 2-4 cm. pedicel; involucre 1 mm.
long by 0.3 mm., with 8-9 bracts; florets ca. 45; corolla pale purple to pinkish, 7-9
mm. long, equalling or slightly (0.1 mm.) exceeding the involucre; corolla lobes short
(0.5-0.7 cm.), stamens with thickened portion of filament 0.4 mm. long, thecae 0.4-0.7
mm. long, anther appendages 0.1-0.2 mm.; achenes 3 mm. long by 0.4 mm., not or
only weakly ribbed between grooves.
ERECHTITES Rafinesque
Reference: Robert O. Belcher, A revision of the genus
Erechtites (Compositae), with inquiries into Senecio and Arrhe-
nechthites, Ann. Mo. Bot. Gard. 43. 1-85. 1956.
Annual herbs (ours), erect, simple or branched; leaves alternate, dentate or
lobate, usually petiolate, or sessile and decurrent or subamplexicaul; inflorescence
corymbose-paniculate, with many rather large, discoid, calyculate heads; flowers
whitish to greenish or purple; involucre uniseriate, cylindric, often enlarged at the
base; phyllaries equal, linear or lanceolate, erect and connivent becoming deflexed at
maturity; marginal florets filiform, 4-5-fid, usually pistillate, the style branches
without marginal fringe but with apex conic-appendaged; central florets in-
fundibuliform, 5-fid, perfect, style branches with crown of fused papillose hairs;
achenes subcylindric to subfusiform, about 10-ribbed; pappus in several series,
capillary.
A small genus of about five species.
Leaves sessile, or alate-petiolate; heads half as broad as long; pappus white.
E. hieracifolia var. cacalioides.
396 FIELDIANA: BOTANY, VOLUME 24
Leaves petiolate or inconspicuously alate at the base; heads one-third as broad as
long; pappus usually more or less reddish E. valerianaefolia.
Erechtites hieracifolia (L.) Raf. var. cacalioides (Fisch. ex
Spreng.) Griseb. Fl. Brit. W. Indies 381. 1861, emend Belcher, Ann.
Mo. Bot Gard. 43: 19. 1956. Senecio cacalioides Fisch. ex Spreng.
Nov. Prov. 37. 1819; Syst. Veg. 3: 565. 1826. Sonchus agrestis Sw.
Prodr. 110. 1788. Erechtites agrestis Standl. & Steyerm. Field Mus.
Bot. 23: 265. 1947.
Moist or wet thickets, sometimes in marshes or pine forest,
often a weed in waste or cultivated ground, 350-2,400 m.; Peten;
Alta Verapaz; Izabal; Zacapa; Jalapa; Jutiapa; Santa Rosa;
Escuintla; Guatemala; Sacatepequez; Suchitepequez; Retalhuleu;
Quezaltenango; San Marcos; Huehuetenango. Mexico; British
Honduras; Honduras and El Salvador to Panama; West Indies;
tropical South America. Asia.
A stout, erect annual, usually 1 m. high or lower, simple or sparsely branched,
the stems stout and thick, striate, somewhat succulent, the branches and leaves
sparsely or often densely sordid-villous; leaves thin, the lower ones petiolate,
narrowly or broadly lanceolate, coarsely and irregularly dentate and often incised-
dentate, the upper cauline leaves much smaller, lanceolate, dentate or incised-lobate,
sessile, usually amplexicaul; heads usually numerous, forming a short or elongate,
narrow, often thyrsiform, leafy-bracteate panicle, slender-pedicellate or sometimes
subsessile and densely clustered; involucre green, 1.5 cm. high; phyllaries numerous,
linear, with white scarious margins, glabrous, the outer basal bracts short, linear or
subulate, densely villous: corollas yellowish green; pappus white, soft, very abundant;
achenes linear, almost 3 mm. long, brownish.
This plant is often a weed, plentiful in many parts of
Guatemala and other regions of Central America.
Erechtites valerianaefolia (Wolf) DC. Prodr. 6: 295. 1838.
Senecio valerianaefolia Wolf, Ind. Sem. Hort. Berol. 1825, teste
Reichb. Icon. Bot. Exot. 59: t. 85. 1827. Figure 130.
Moist or wet thickets, pine-oak forest, or often a weed in coffee
plantations, 1,200-2,500 m.; Alta Verapaz; Zacapa; Guatemala;
Sacatepequez; Chimaltenango; Suchitepequez; Solola; El Quiche;
Huehuetenango; Quezaltenango; San Marcos. Southern Mexico; El
Salvador; Nicaragua; Costa Rica; Panama; tropical South America.
Old World.
Plants stout, erect, annual, 1 m. high or less, the stems striate, simple or
branched, villous or glabrate; lower leaves long-petiolate, ovate to elliptic or
lanceolate, acuminate, somewhat runcinate-lobate and coarsely and unequally
dentate, usually narrowed and decurrent at the base, glabrate above, sparsely or
rather densely villous beneath, the upper cauline leaves deeply pinnate-lobate, often
WILLIAMS: FLORA OF GUATEMALA 397
almost to the costa, the segments few or numerous, unequal the lateral ones mostly
lanceolate, laciniate-dentate; heads rose-purple, usually very numerous, in mostly
elongate and narrow, leafy panicles, on short or elongate, slender pedicels; involucre 1
cm. high, glabrous; phyllaries green, linear, the basal bracts very short, subulate,
glabrous or nearly so; pappus very copious, purplish above.
An agressive weed, perhaps sometimes hybridizing with E.
hieracifolia var. cacalioides. The variety found in our region is var.
valerianaefolia.
GYNURA Cassini, Velvet plant
Annual or perennial herbs with soft, often purplish pubescence; the leaves entire
or serrate-dentate (ours), petiolate; heads discoid, calyculate; involucre cylindric or
campanulate, the phyllaries in a single series; achenes 5-10-ridged; pappus abundant,
multiseriate, about as long as the florets.
Twenty species or more of Asia and Africa, one cultivated or
escaped in Central America. Related to Emilia.
Gynura aurantiaca (Blume) DC. Prodr. 6: 300. 1837. Cacalia
aurantiaca Blume, Bidjr. 908. 1826. Hoja tornasol.
Annual or perhaps perennial herbs to 1 m. tall, stout, branched, covered with
segmented, violet or purple hairs; leaves lanceolate-ovate to ovate, acute, dentate,
the uppermost sessile and amplexicaul; heads to 2 cm. tall in a lax terminal raceme;
florets orange to yellow.
Native of Java. Cultivated and perhaps as escape, probably not
enduring as an escape.
PSACALIUM Cassini
Reference: Richard W. Pippen, Mexican "Cacalioid" genera
allied to Senecio (Compositae), Contr. U. S. Nat. Herb. 34: 365-447.
1968.
Perennial herbs from a thick, fleshy root, to 3 m. tall; stems erect, annual,
slender or stout, glabrous or pubescent (ours); basal leaves 2-several, centrally
peltate and palmately nerved, the blades lobate or rarely subentire, cauline leaves
peltate or the upper reduced or bractlike and not peltate; inflorescence paniculate or
corymbose, glandular-puberulent or not; heads many (ours), calyculate, the bracts as
long as the phyllaries or shorter; involucre cylindrical to campanulate; phyllaries 5-17
(5 in ours), subequal, sub-biseriate, florets all discoid, 5-50; corollas with narrow
cylindrical tubes, the limb deeply divided into 5 narrow lobes, these recurved when
mature; anthers exserted; style branches spreading or at maturity to doubly
recurved; achenes 10-16, costate; pappus a single row of barbellate bristles.
A genus of about 15 species in Mexico and Guatemala, all
Mexican except the two mentioned below. Most have been referred
to Cacalia at one time or another.
398 FIELDIANA: BOTANY, VOLUME 24
Phyllaries 8-9 mm. long, glandular-puberulent with very short hairs; cauline leaves
presumed to be sessile p. guatemalense.
Phyllaries 10-11 mm. long, villous with rather long, lax, eglandular hairs; cauline
leaves mostly long petiolate p. pinetorum.
Psacalium guatemalense (Standl. & Steyerm.) Cuatr. Brit-
tonia 8: 157. 1955. Cacalia guatemalensis Standl. & Steyerm. Field
Mus. Bot. 23: 99. 1944.
Known only from the type, summit of Buena Vista, Montana
Miramundo, between Miramundo and Buena Vista, between Jalapa
and Lago de Ayarza, Jalapa, 2,000-2,200 m., Steyermark 32823.
Plants very stout, about 1 m. high, the stems terete, striate, glandular-puberulent
throughout; basal and lowest cauline leaves unknown, the middle and upper ones
sessile, ovate-oblong, 25 cm. long or less, 10-13 cm. broad, obtuse, cordate-amplexicaul
at the base, rather shallowly about 9-lobate or coarsely and irregularly dentate,
puberulent and glandular above, densely whitish lanate beneath; inflorescence
paniculate, large, the individual panicles 8-10 cm. long, densely glandular-puberulent,
the pedicels 2-4 mm. long; heads turbinate-cylindric, 5-9-flowered, 3.5-6 mm. broad;
involucre 8 mm. high; phyllaries 5, linear-lanceolate, acute, glandular-puberulent;
corollas 9-10 mm. long, the tube cylindric, the limb somewhat ampliate, divided
nearly to the base into linear-lanceolate, recurved or recoiled lobes; achenes glabrous,
6 mm. long, about 16-costate; pappus ample, white.
Pippen, in his monograph (p. 434) excludes this species from
Psacalium since he thought the leaves not to be peltate. Basal
leaves are unknown but doubtless will prove to be peltate, since the
floral morphology is exactly that of Psacalium.
Psacalium pinetorum (Standl. & Steyerm.) Cuatr. Brittonia
8: 157. 1955. Cacalia pinetorum Standl. & Steyerm. Field Mus. Bot.
23: 142. 1944. Figure 131.
Mixed woods, open slopes and ravines, 2,600-3,200 m.; Quezal-
tenango (type, Steyermark 34933). Endemic.
Viscid herbs to 1 m. tall, stems puberulent with multicellular hairs, these often
glandular-viscid; leaves toward the base of the stem large, peltate and petiolate, those
above much reduced and bractlike, sessile and not peltate, the leaves peltate near the
center of the suborbicular, deeply incised blade, the blade about 8-lobed, 7-18 cm.
across, the lobes 2-to-several lobulate, the lower surface densely puberulent with
short, crisped hairs, the upper surface sparsely puberulent; inflorescence of several
rather strict several-many headed panicles from the axils of upper leaves; heads
homogamous, discoid; involucre cylindric, calyculate; phyllaries 5, glandular-
puberulent, about 7-8 mm. long, 2-3 mm. broad, lanceolate to oblong-lanceolate,
obtuse, 3 with scarious margins, the others not, puberulent with crisped hairs; florets
5; the corollas about 9 mm. long, tubular below, expanded and deeply lobed above,
the lobes linear- lanceolate, acute, about 3.5 mm. long; pappus abundant, shorter than
the corollas; achenes 10-ridged, about 2.5-3 mm. long.
WILLIAMS: FLORA OF GUATEMALA 399
Known from two collections but closely related to P. pippenii
L. Wms. from Mexico.
EXCLUDED
CACALIA CALOTRICHA Blake, Contr. Gray Herb. 52: 58. 1917. =
Senecio heterogamus (Benth.) Hemsl.
SENECIO (Tournefort) Linnaeus
References: Jesse More Greenman, Monographic der nord- und
centralamerikanischen Arten der Gattung Senecio, Engler's Bot.
Jahrb. 32: 1-33 1902 (synopsis only); Monograph of the North and
Central American species of the genus Senecio, Part II, Ann. Mo.
Bot. Gard. 2: 573-626. 1915, op. cit. 3: 85-194. 1916, op. cit. 4: 15-36.
1917; op. cit. 5: 37-103. 1918. Incomplete.
Annual or perennial herbs, shrubs, vines or weak trees; leaves alternate or
radical, pinnately or palmately veined, entire or variously dentate or divided; heads
heterogamous and radiate, or discoid; involucre cylindrical or campanulate, usually
subtended by one or more bracteoles; phyllaries uniseriate or by overlapping
sub-biseriate, variable in number (but usually subdefinite in a given species); ray
flowers when present dispersed in a single row, fertile, the rays sometimes reduced;
disc flowers perfect or heterogamous with the corollas tubular to ampliate and
campanulate above, the 5 teeth or lobes of the limb usually short but sometimes
nearly as long as the throat; anthers obtuse or subsagittate at the base; style
branches subterete, truncate, rounded-obtuse or with a tuft of hairs, recurved or
spreading, rarely terminated (section Convolvuloides) by an acute or acuminate
appendage; achenes subterete, usually ribbed, glabrous or hirtellous especially on the
angles; pappus of numerous, usually white (rarely purple) setae.
A large and diverse genus distributed throughout much of the
world, with perhaps as many as 1,500 species. It is found from
deserts and cold arctic regions to deep tropical rain forests. Many
are herbs, but shrubs and weak trees occur in the tropics. The genus
is represented by many species in Guatemala, but rarely are great
numers of individuals seen together, so as to appear to be a
prominent part of the vegetative cover. A small number of
additional species occur in other Central American countries. The
genus is represented by a much larger number of species in Mexico.
Dr. Jesse More Greenman, of fond memory, studied Senecio for
a lifetime, but only a portion of his "Monograph of the North and
Central American species of the genus Senecio" was published
before his death. Greenman divided the genus into two subgenera,
and these into 22 sections. The subgeneric treatment which follows
is essentially that proposed and used by Dr. Greenman.
400 FIELDIANA: BOTANY, VOLUME 24
Several segregate genera of Senecio have been proposed or
resurrected by Robinson and Brettell. These and allies are treated
in Phytologia 27: 402-439. 1974.
Richard W. Pippen's paper on the "Mexican 'Cacalioid' genera
allied to Senecio (Compositae)," in Contr. U. S. Nat. Herb. 34: 365-
447, tt. 1968, is basically a better account of the Senecio allies than
Robinson and Brettell's. It does not treat the species here
considered to belong to Senecio, but does account for the Cacalioid
genera related to Senecio.
The following segregates are considered to be synonymous with
Senecio for purposes of this flora; Barcleyanthus, Nelsonianthus,
Psacaliopsis, Roldana, Telanthophora, and Pseudogynoxys. The
author feels that these segregates have but slight importance in the
account of Senecio.
KEY TO THE SUBGENERA AND SECTIONS
Style branches truncate, rounded-obtuse or terminated with a penicillate tuft of
hairs SUBGENUS I, SENECIO, with 7 sections in Guatemala, p. 400.
Style branches terminated by triangular, acute or acuminate, dorsally hispidulous
appendages. SUBGENUS II, PSEUDOGYNOXIS, with a single section in
Guatemala, p. 401.
SUBGENUS I, SENECIO.
Plants epiphytic vines or shrubs, the stems pendent or climbing; leaves often fleshy,
the nerves obscure or in 2 species conspicuous; phyllaries 8; heads radiate or not.
Section 1, Streptothamni, p. 401.
Plants not epiphytic, all terrestrial, herbs, shrubs or weak trees; phyllaries 5, 8, or 10
or more; heads radiate or not; in two species the disc flowers heterogamous.
Mature leaves about as broad as long or usually broader than long, subentire to
deeply lobate and usually subcordate and palmately nerved; plants usually
weak shrubs, some herbaceous.
Inflorescence a single nodding head; subscapose herbs with peltate leaves.
Section 3, Psacaliopsides, p. 402.
Inflorescence usually with many erect heads; caulescent shrubs or coarse herbs;
leaves peltate (1 species) or not Section 2, Palmatinervii, p. 401.
Mature leaves obviously much longer than broad, entire to variously lobate or
dissected; herbs, shrubs or trees.
Plants herbaceous.
Stems scapose; the inflorescences with fewer than 5 heads.
Section 8, Tomentosi, p. 403.
Stems not scapose; the inflorescences with many heads.
Leaves floccose-tomentose below; heads radiate.
Section 6, Amplectentes, p. 403.
Leaves nearly glabrous below; heads usually discoid.
Section 7, Mulgedifolii, p. 403.
WILLIAMS: FLORA OF GUATEMALA 401
Plants woody, or suffrutescent, at least at the base.
Stems woody to the top, shrubs or weak trees Section 4, Fruticosi, p. 402.
Stems suffrutescent, at least at the base.
Lateral nerves of the leaves parallel-arcuate. ...Section 5, Multinervii, p. 403.
Lateral nerves of the leaves not parallel-arcuate; leaves auriculate or
subamplexicaul at the base Section 6, Amplectentes, p. 403.
SUBGENUS II, PSEUDOGYNOXIS Greenm.
A single section with a single variable species Section 9, Convolvuloides, p. 403.
KEY To THE SPECIES IN THE VARIOUS SECTIONS
Section 1, STREPTOTHAMNI. Epiphytic vines or weak shrubs or subherbaceous, the
stems erect, climbing or pendent; leaves usually fleshy, the nerves obscure or in two
species obvious; phyllaries 8; heads radiate or not, if not the heads homogamous.
Leaves 4 cm. or more broad (and 1 l/2 times as long); pinnate nerves 7-8 pairs and
prominent S. epidendrus.
Leaves much less than 4 cm. broad, nerves subpalmate, subtriplinerved or nerves
obscure.
Leaf venation obviously subpalmate or triplinerved; upright epiphytic herbs or
suffrutescent herbs; ray flowers none S. armentalis.
Leaf venation obscure; pendent epiphytic vines or shrubs; ray flowers present.
Phyllaries linear-oblong, 8, about as long as the corollas; a species of the Atlantic
forests at middle elevations, from Veracruz to Alta Verapaz.
S. parasiticus.
Phyllaries much shorter than the heads, lanceolate or ovate, 8 or connate into 5-
6; plants of the Atlantic slope S. phorodendroides.
Section 2, PALMATINERVII. Mostly large shrubs or small weak trees of the moist
forests; leaves mostly palmately nerved and as broad as long or broader (if longer
than broad then semipenninerved), one species has peltate leaves; phyllaries 5, 8, or
(10 or more) 13; heads radiate or not, 2 species with heterogamous disc flowers.
Phyllaries 8 or fewer, often 5.
Leaves mostly longer than broad S. steyermarkii.
Leaves mostly broader than long.
Ray flowers present, usually 2-5, often soon deciduous.
Lobes of the leaves rounded or obtuse, leaves usually densely floccose-
tomentose beneath S. petasioides.
Lobes of the leaves acute or acuminate, leaves sparsely floccose to glabrous
beneath.
Phyllaries glandular-hispidulous, the pubescence often reddish.
S. jurgensenii.
Phyllaries glabrate, not at all glandular-hispidulous S. acutangulus.
Ray flowers absent.
Heads heterogamous, about 8 pistillate florets around the margin of each head,
the others perfect; leaves large, 25-45 cm. broad, the lobes deeply incised.
S. greenmanii.
402 FIELDIANA: BOTANY, VOLUME 24
Heads homogamous, all florets perfect; leaves smaller, 4-20 cm. broad,
relatively shallowly lobate S. cristobalensis.
Phyllaries 10 or more, often 13.
Leaves mostly peltate; ray flowers none but 2-6 disc florets around the margin of
each head pistillate, the others perfect S. heterogamus.
Leaves never peltate; ray florets 2-8, soon deciduous.
Heads large, 12-20 mm. long, with about 30 disc florets and 7-8 ray florets.
S. gilgii.
Heads smaller, 8-12 mm. long, with 10-20 disc florets and 2-8 ray florets.
Leaves penninerved, often longer than broad.
Lower leaf surface densely floccose-tomentose, the pubescense completely
covering the surface; leaves shallowly lobate; heads with 2-8 ray florets.
S. barba-johannis.
Lower leaf surface sparsely floccose; leaves more prominently lobate; heads
with 6-8 ray florets S. quezalticus.
Leaves palmately nerved, never longer than broad, heads with 6-8 ray florets.
S. lanicaulis.
Section 3, PSACALIOPSIDES. Subscapose herbs of subalpine regions; heads one (ours)
or few; leaves peltate.
One species in Guatemala S. nubivagus.
Section 4, FRUTICOSI (including section TERMINALES of Greenman). A group of
species, largely of convenience, that are shrubs or weak trees with penninerved leaves.
Phyllaries either 12-15, 8 or 5 in various species. Certain species seem to be related to
section PALMATINERVII and two of the species are keyed under that section.
Lower leaf surface densely floccose-tomentose.
Leaves linear-lanceolate to lanceolate; phyllaries 12-15 S. deppeanus.
Leaves elliptic-ovate to ovate or oval.
Phyllaries about 8 S. steyermarkii.
Phyllaries 12-13 S. barba-johannis.
Lower leaf surface glabrous or nearly so, or if sparsely floccose then appearing
stellate-floccose.
Leaves linear to elliptic-linear, 0.2-1 cm. broad S. salignus.
Leaves narrowly lanceolate to ovate, mostly always more than 3 cm. broad.
Phyllaries 5.
Leaves elliptic-oblanceolate to oblanceolate, long attenuate-cuneate to the
base, on short petioles, entire or somewhat dentate-serrate above; florets
usually 5, 1-2 radiate or all discoid S. cobanensis.
Leaves lanceolate or lanceolate-oblong to ovate, obtuse or abruptly attenuated
to the base, not long cuneate, entire or lobate-dentate.
Mature leaves entire, lanceolate-oblong, mostly 15-30 cm. long and 4-10 cm.
broad; heads radiate, mostly less than 8 mm. long; phyllaries about as
long as the head S. uspantanensis.
Mature leaves entire to sinuate-dentate to lobate-dentate, mostly ovate to
ovate-lanceolate; heads mostly radiate with more than 5 florets;
phyllaries much shorter than the heads S. schaffneri.
Phyllaries 8 (-7).
WILLIAMS: FLORA OF GUATEMALA 403
Leaf margins prominently lobate-dentate to lacerate-lobate; common forest
bog species from Mexico to Panama S. arborescens.
Leaf margins entire or shallowly and sparsely dentate to dentate-lobulate;
uncommon species of Mexico, Guatemala and British Honduras.
Leaves oblanceolate to oblanceolate-ovate, entire or denticulate, fleshy, the
nerves obscure S. montidorsensis.
Leaves lanceolate to oblong or oblong-ovate, the nerves conspicuous.
Heads 7-8 mm. long; leaves obscurely dentate to dentate-lobulate; known
from adjacent Mexico S. grandifolius.
Heads mostly about 6 mm. long; leaves subentire to dentate-lobate; young
stems densely floccose-tomentose; occasional in Alta Verapaz and
Suchitepequez S. serraquitchensis.
Section 5, MuLTiNERVii. Perennial herbs of the subalpine regions, with thick heavy
rootstalks, white-tomentose almost throughout; nerves of the leaves parallel-arcuate.
One species S. oerstedianus.
Section 6, AMPLECTENTES. Perennial herbs with the stems leafy to the inflorescence,
the leaves floccose-tomentose below and auriculate or semiamplexicaul at the base.
A single species S. warszewiczii.
Section 7, MULGEDIFOLII Greenm. Perennial or occasionally flowering as biennial
herbs, the stems leafy with the leaves reduced upward; leaves petiolate, variously
dentate to deeply incised; heads discoid or rarely radiate; phyllaries 8-13; florets
purplish or whitish to yellow. The three species credited to Guatemala are often
difficult to distinguish. Monographic studies are needed.
Basal leaves long petiolate and hastate or obtuse at the base; phyllaries on mature
heads about two-thirds or less the length of the heads S. doratophyllus.
Basal leaves runcinate-pinnatifid; phyllaries nearly as long as the heads.
Lower cauline leaves runcinate-pinnatifid S. rhyacophilus.
Lower cauline leaves dentate to lobulate, not runcinate S. callosus.
Section 8, TOMENTOSI. Subscapose or scapose terrestrial herbs; inflorescence with
fewer than 5 heads; heads radiate; plants of the subalpine region.
Leaves linear-lanceolate to oblanceolate, acute S. gerberaefolius.
Leaves oblong-ovate to suborbicular, obtuse S. cuchumatanensis.
Section 9, CONVOLVULOIDES.
One species S. chenopodioides.
Senecio acutangulus (Bertol.) Hemsl. Biol. Cent.-Am. Bot. 2:
235. 1881. Cineraria acutangula Bertol. Fl. Guat. 35. 1840 (type:
Sacatepequez, Velasquez}. Roldana acutangula Robinson & Bret-
tell, Phytologia 27: 415. 1974.
Moist forests and thickets, fairly common in the western
departments, principally in December and January, (1,200-) 2,000-
404 FIELDIANA: BOTANY, VOLUME 24
3,600 m.; Sacatepequez; Chimaltenango; Solola; Huehuetenango;
Totonicapan; Quezaltenango; San Marcos. Endemic, but to be
expected in Chiapas.
Weak, mostly glabrous shrubs or trees to 4 m. tall, the stem to 10 cm. in
diameter, araneose pubescent or glabrous; leaves 5-7 lobate, orbicular to broadly
reniform, the margins callose-dentate, long to short petiolate, glabrous to sparsely
puberulent to floccose below, the lobes mostly acute, the blades subpenninerved,
obtuse to cordate at the base, 6-25 cm. long and as broad or broader, petioles almost
none to 20 cm. long, slender; inflorescence a multicapitate, corymbose panicle with
slender, usually glabrous pedicels, usually a little longer than the leaves; heads
heterogamous, mostly about 1 cm. long, subcylindric but spreading in age; involucre
mostly with 8 phyllaries, these about 8 mm. long, as long as the pappus, glabrous,
linear-oblong, acute, margins of half of them scarious; flowers 10-14, 2-5 radiate, the
others discoid; achenes glabrous; pappus equaling the disc corollas.
Distinguishable from S. jurgensenii principally by the lack of
glandular-hispidulous pubescence in the inflorescence. Perhaps S.
jurgensenii should be included in this species.
Senecio arborescens Steetz in Seem. Bot. Voy. Herald 162, t.
31. 1854. S. chicharrensis Greenm. Field Mus. Bot. 2: 285. 1907.
Telanthophora arborescens Robinson & Brettell, Phytologia 27:
426. 1974. T. chicharrensis Robinson & Brettell, I.e. Estrella, mano
de leon.
Commonly in ravines and boggy places in wet forests, mostly
1,200-1,600 m. but as low as 600 m. in Guatemala. Alta Verapaz;
Guatemala; Escuintla; El Quiche; Quezaltenango; San Marcos.
Mexico (Chiapas) to Panama.
Weak, simple or branched shrubs or trees, 1-12 m. tall, sometimes said to be
herbs, the young stems floccose or in age nearly glabrous, pithy; leaves from
sinuately lobate to deeply incised, mostly oblong to ovate in outline, obtuse, acute or
even acuminate, the base cuneate to subcordate. glabrous above at maturity, usually
sparsely floccose along the veins below, the blades 10-40 (-60) cm. long and 5-25 (-30)
cm. broad, the petiole 3-9 cm. long; inflorescences terminal or in the axils of
uppermost leaves, multicapitate, usually long pedunculate, corymbose panicles about
half as long as the subtending leaves; heads radiate, campanulate, about 7-8 mm.
high, subtended by linear bracteoles or ecalyculate; involucre about two-thirds as
long as the heads, phyllaries usually 8, linear or narrowly lanceolate, acute or obtuse,
about 4-6 mm. long; flowers yellow, 5-11 in a head, usually with 2-3 ray flowers and
3-9 disc flowers, ray corollas about 5 mm. long, the lamina 2 mm. the disc corollas
tubular, about 5 mm. long, the lobes narrow; anthers exserted.
A variable species commonly with deeply incised leaves but
sometimes sinuate-lobate. Perhaps not abundantly distinct from
Senecio grandifolius or only a variety of it.
Senecio armentalis L. Wms. Phytologia 28: 225. 1974.
WILLIAMS: FLORA OF GUATEMALA 405
Nelsonianthus epiphyticus Robinson & Brettell, Phytologia 27: 54.
1973, not Senecio epiphyticus O. Kuntze.
On cold, wet, mossy banks or more usually epiphytic in mossy
montane forests, 2,600-4,000 m., Solola; Huehuetenango; Totonica-
pan; Quezaltenango; San Marcos. Endemic.
Small shrubs or suffrutescent herbs to about 50 cm. long, the stems somewhat
fleshy, occasionally leafless at time of flowering, glabrous; leaves lanceolate to ovate,
acute or acuminate, obtuse at the base, petiolate, entire or sparsely callose-
denticulate, principal nerves from above the base of the blade, blade 4-10 cm. long
and 2-6.5 cm. broad, the petiole slender, to 2.5 cm. long; inflorescences terminal, few-
many-flowered panicles; heads narrowly campanulate, mostly 12-15 mm. long, the
florets yellow; involucre calyculate, glabrous, phyllaries usually 8, purple, linear to
linear-oblong, acute, glabrous, shorter than the florets, 9-12 mm. long and 1.2-2 mm.
broad; florets homogamous, discoid, 9-12; corolla subcylindric, slightly, expanded
from below the middle, the throat somewhat wider than the tube, about 10 mm. long,
the lobes lanceolate and about 1.5 mm. long; achenes glabrous, ridged, 2-4 mm. long;
pappus abundant, as long as the florets.
A rare but attractive Senecio, usually to be found epiphytic on
oaks in the montane forests.
Senecio barba-johannis DC. Prodr. 6: 430. 1837. S. donnell-
smithii Coulter, Bot. Gaz. 16: 100. 1891 (type from Volcan de Agua,
Donnell- Smith 2362). Roldana barba-johannis Robinson & Brettell,
Phytologia 27: 415. 1974. R. donnell-smithii Robinson & Brettell,
torn. cit. 418.
Occasionally abundant in mixed forests or coniferous forests at
2,000-3,700 m., or higher in Mexico; El Progreso; Sacatepequez;
Chimaltenango; Huehuetenango; Quezaltenango. Mexico.
Weak and brittle shrubs to about 7 m. tall, the stems floccose-tomentose,
becoming glabrescent, pithy; leaves elliptic-ovate to broadly oval, cordate, subcoriac-
eous, petiolate, the blade 6-20 cm. long and 4-10 cm. broad, densely floccose-
tomentose below, glabrous above, the petiole to about 6 cm. long; inflorescence
terminal and from the axils of the uppermost leaves, of multicapitate, dense, long-
pedunculate corymbose cymes; heads radiate, about 6 mm. high; involucre obscurely
calyculate, campanulate; phyllaries about 12-13, alternately linear and oblong-
lanceolate, acute, about 3 mm. long, floccose or glabrescent dorsally, half as long as
the heads; ray flowers about 2 in each head, the lamina oblong-elliptic, about 2 mm.
long; disc flowers about 10, corollas subcylindric, about 4 mm. long, the lobes linear-
lanceolate about 1 mm. long; achenes about 2 mm. long, glabrous.
The only broad-leafed, penninerved, weak shrub in which the
under surface of the leaf is tomentose and there are 12-13
phyllaries. The similar S. steyermarkii has eight phyllaries.
Senecio callosus Sch.-Bip. Flora 28: 498. 1845. S. decorus
406 FIELDIANA: BOTANY, VOLUME 24
Greenm. Field Mus. Bot. 2: 279. 1907 (type, Sacatepequez, Donnell
Smith 2361).
Most often found in coniferous or coniferous-oak forest; 2,500-
4,000 m.; Sacatepequez; Chimaltenango; Solola; Huehuetenango;
Totonicapan; San Marcos. Mexico.
Erect, perennial herbs, 1 m. or less tall; leaves basal and cauline, the basal leaves
runcinate-pinnatifid, to 25 cm. long and 14 cm. broad, the terminal lobe somewhat
triangular, acuminate, blades arachnoid-pubescent below to nearly glabrous, petioles
often elongate and sometimes dentate-alate, cauline leaves smaller, usually sessile
and amplexicaul, subentire and dentate to pinnatifid, arachnoid-pubescent below and
sometimes above; inflorescence a terminal few-several-flowered panicle or corymb;
heads discoid, 8-14 mm. long and half as broad; involucre campanulate, calyculate;
phyllaries about 12-13, linear to narrowly oblong, obtuse or acute, glabrous except at
the tip, mostly 8-10 mm. long; corollas exceeding the involucre, pale yellow to
purplish; achenes striate, glabrous.
A rather uncommon plant in the western highlands and
vegetatively a variable one, especially in foliage.
Senecio chenopodioides HBK. Nov. Gen. & Sp. PL 4: 179.
1820; L. Wms. Phytologia 31: 437. 1975. Gynoxys haenkei DC.
Prodr. 6: 326. 1836 (basionym for Senecio kermesinus). G.
berlandieri DC. I.e. G. fragrans Hook. Bot. Mag. 76: t. 4511. 1850
(grown from seed sent from Guatemala by Skinner). Senecio
berlandieri Hemsl. Biol. Cent.-Am. Bot. 2: 236. 1881, not Sch.-Bip.
1845. S. calocephalus Hemsl. torn. cit. 237. S. kermesinus Hemsl. I.e.
S. skinneri Hemsl. torn. cit. 247 (based on Gynoxys fragrans). S.
chinotegensis Klatt, Leopoldina 24: 125. 1888. S. hoffmannii Klatt,
Leopoldina 25: 106. 1889. S. confusus Britten, Journ. Bot. 36: 260.
1898. S. convolvuloides Greenm. Monog. nord. und centralam. Art.
Gattung Senecio, pt. 1, 26. 1901; Engler Bot. Jahrb. 32: 22. 1902,
nomen, S. trixioides Greenm. ll.cc., nomen. S. rothschuhianus
Greenm. ll.cc., nomen. Pseudogynoxys haenkei Cabrera, Brittonia 7:
54. 1950. P. hoffmannii Cuatr. Brittonia 8: 156. 1955.
Thickets and forest edges, commonly at 700-1,200 m. but found
from near sea level to 2,200 m.; known from all departments except
Izabal, El Progreso and Totonicapan and to be expected in all.
Mexico; El Salvador; Honduras; Nicaragua; Costa Rica.
Small herbaceous to large suffrutescent vines, the stems multistriate, sparsely
hispidulous to almost glabrous, up to 5 m. and perhaps longer; leaves alternate,
petiolate or nearly sessile, lanceolate to lanceolate-ovate, rarely subcordate, acute or
acuminate, densely hispidulous to glabrous, serrate-dentate, the teeth sometimes
glandular, the blade 3-12 cm. long and 1-7 cm. broad; inflorescence a single head to a
multicapitate panicle; heads heterogamous, variable in size, 1-3 cm. high (commonly
WILLIAMS: FLORA OF GUATEMALA 407
1.5 cm.) and as broad; involucre uniseriate, subtended usually by several conspicuous
bracteoles, mostly about 1 cm. long; phyllaries numerous, linear, acuminate, densely
hispidulous to glabrous, the mid-nerve thickened, phyllaries half as long as the head
or usually longer; ray corollas orange (usually) to reddish, uniseriate; disc corollas
yellow to orange or red, mostly about 1 cm. long, tubular to narrowly campanulate
above; achenes hispidulous, cylindric, obscurely ridged, to about 4 mm. long; pappus
abundant, white, slightly shorter than the disc corollas.
An attractive and quite common vine at the beginning of the
dry season. A variable species in most all parts. Heads in
multicapitate inflorescences are often smaller than those in few- or
unicapitate inflorescences. This is the only terrestrial vining
Senecio, at middle to low elevations, in Central America. A few
vine-like epiphytes occur in the cloud forests.
Senecio cobanensis Coulter, Bot. Gaz. 16: 101. 1891. S.
cobanensis var. sublaciniatus Greenm. Ceiba 1: 120. 1950 (type
from Sacatepequez, Standley 63689). Telanthophora cobanensis
Robinson & Brettell, Phytologia 27: 427. 1974. T. molinae Robinson
& Brettell, torn. cit. 428 (type, Williams, Molina & Williams 26271,
from San Marcos). Figure 132.
Wet forest or forest edges, 1,300-2,800 m.; Alta Verapaz (type
(type, Tuerckheim 1158); El Progreso; Chimaltenango; Solola;
Huehuetenango; San Marcos; Suchitepequez. Mexico (Chiapas);
Honduras; perhaps El Salvador.
Shrubs or weak trees to 9 m. tall but mostly 2-3 m. tall, glabrous; leaves crowded
toward the apex of the branches, slightly to conspicuously fleshy, glabrous entire to
sometimes dentate, petiolate, the blades narrowly elliptic-oblanceolate to oblanceo-
late, acuminate, 8-30 cm. long and 1.5-10 cm. broad (commonly 15-20 cm. long and 2-
2.5 cm. broad), the petiole 1.5-15 cm. long; inflorescences few to many long-
pedunculate, often massive, compound corymbs from the apex of the branches,
usually exceeded in length by the leaves; heads in an inflorescence radiate, discoid or
mixed, small, mostly 6-8 mm. long and half as broad; when radiate corollas present,
then usually the head with one radiate flower and four disc flowers (see illustration)
or all five flowers in a head discoid (the common situation in most collections);
involucre ecalyculate but subglandular at the base, glabrous; phyllaries 5, lance-
oblong, acute, the margins of 2 or 3 obviously scarious, 5-6 mm. long and 1-2 mm.
broad, about as long as the corollas; ray flowers pistillate and perhaps usually sterile,
the lamina elliptic, to about 5 mm. long; disc corollas about 5-6 mm. long, the lobes
lanceolate, recurved, about 2 mm. long; achenes about 5-ridged, glabrous; pappus
about as long as the corollas, minutely barbellate.
A complex group of plants and a variable one. There seems to
be no difference between discoid and radiate individuals except the
obvious ones in the flowers. Some discoid flowers may be
functionally only staminate since the style and stigmas are much
408 FIELDIANA: BOTANY, VOLUME 24
reduced. There are no obvious geographical and ecological differ-
ences in the population. All Honduran collections, except one, are
discoid. Most radiate specimens are from the mountains of adjacent
Mexico and Guatemala, but not all from that region are radiate.
Senecio cristobalensis Greenm. ex Loesener, Bull. Herb.
Boiss. ser. 2, 6: 867. 1906. Roldana cristobalensis Robinson &
Brettell, Phytologia 27: 417. 1974.
Moist forests or thickets, 1,300-1,900 m.; Alta Verapaz;
Huehuetenango. Mexico (Chiapas).
Weak shrubs 1-2 m. tall, the stems slender, hispidulous; leaves suborbicular or
subcordate-reniform in outline, sparsely callose-denticulate or entire, 5-7-lobate, the
lobes rounded and obtuse, subpenninerved, hispidulous-puberulent, less so above to
almost glabrous, the blades 4-18 cm. long and 4-20 cm. broad, petioles from nearly
none on the uppermost reduced leaves to 20 cm. long on the lower leaves;
inflorescence a multicapitate corymbose panicle, usually much exceeding the leaves,
hispidulous or reddish glandular-hispidulous; heads discoid, 8-14 mm. long; involucre
subcampanulate; phyllaries about 8, 4-7 mm. long, about half as long as the head,
lanceolate or linear-lanceolate, acute, hispidulous-pubescent dorsally, the innermost
of the series scarious margined and broader than the outer ones; corollas all discoid,
to 10 mm. long, cylindric but somewhat expanded toward the apex, the lobes linear-
lanceolate, about 1 mm. long; pappus copious, about as long as the corollas.
A relatively rare plant. It is questionable if the species is
distinct from S. petasioides Greenm., from which it differs in the
lack of ray flowers, and usually less floccose leaves, especially on
the lower surface.
Senecio cuchumatanensis Williams & Molina, Phytologia 31:
438. 1975.
Subalpine, limestone summits of the Cuchumatanes Moun-
tains, probably only 3,300-3,400 m. (type, Steyermark 50170).
Endemic.
Subscapose perennials to about 30 cm. tall; the scapes slender, purple, with
reduced leaves or bracts; the long-petiolate leaves crowded at the crown of the thick
roots, the blade oblong-ovate to suborbicular, obtuse, revolute, sparsely araneose and
purple beneath, glabrescent, 1-3.5 cm. long and 1-1.5 cm. broad, petioles to 3 cm.
long; inflorescence one or two heads at the end of the scape; heads calyculate,
heterogamous, campanulate, the florets yellow; involucre campanulate, sparsely
puberulent to glabrous; phyllaries about 20, linear-lanceolate, acute, the tips
penicillate, about 7 mm. long and 1.5 mm. broad; receptacle slightly alveolate; florets
about 60, 10-12 radiate and about 50 discoid; ray corollas tubular with a conspicuous
ray, the tube about 5 mm. long, the rays about 10-12 mm. long and 3 mm. broad,
elliptic, tridentate; disc corollas 8-9 mm. long, tubular below, throat narrowly
campanulate, about 3.5 mm. long, the lobes narrowly triangular, about 1 mm. long;
anthers and styles slightly exserted at anthesis; immature achenes about 2 mm. long,
pubescent; pappus abundant, caducous, about three-fourths as long as the corolla.
WILLIAMS: FLORA OF GUATEMALA 409
Allied to Senecio gerberaefolius Sch.-Bip., and found in the
same subalpine region of the Cuchumatanes in Huehuetenango.
The suborbicular to oblong-ovate leaves abruptly attenuated to the
petiole, the greater number of phyllaries and the less pubescent
undersurfaces of the leaves will assist in separating the two.
Senecio deppeanus Hemsl. Biol. Cent. Am. Bot. 2: 239. 1881.
S. thomasii Klatt, Abh. Naturf. Ges. Halle 15: 332. 1882. Papelillo;
ho/a de ceniza.
Thickets and mixed forests, rarely on disturbed roadsides, 600-
2,200 m.; Baja Verapaz; Jalapa; Zacapa; Guatemala; Escuintla;
Chimaltenango; El Quiche; Huehuetenango. Southern Mexico from
whence the type; El Salvador; Honduras.
Erect, branching shrubs or weak trees to 5 m. tall, the stems ridged, densely
tomentose or glabrescent with age; leaves linear-lanceolate to lanceolate, acute or
acuminate, (3-) 5-20 cm. long and 1-4 cm. broad, margins slightly revolute, obscurely
dentate or entire, densely white-floccose-tomentose below, glabrous or nearly so
above, petiolate or the uppermost leaves nearly sessile, petioles to 3 cm. long;
inflorescences terminal or in the axils of reduced uppermost leaves, of multicapitate
corymbose panicles to about 10 cm. high and as broad; heads radiate, relatively
small, yellow- flowered, mostly 6-8 mm. long and 4-5 mm. broad; involucre narrowly
campanulate; phyllaries about 12-15, linear, acute, glabrous or nearly so, 5-6 mm.
long; ray corollas about 5 mm. long, with the elliptic blade about 2 mm. long or
more; disc corollas cylindric, 3.5-4.5 mm. long; pappus shorter than the corolla.
Occasionally abundant in pine-oak forests and thickets and
sometimes found on disturbed soil. Called panza de burro in El
Salvador and amargosa, hoja blanca, and Molul in Honduras. Most
botanists, including Dr. Greenman, have used the name S. thomasii
but there seems no justification for so doing.
Senecio doratophyllus Benth. PL Hartw. 87. 1841 (type:
Hartweg 594, Acatenango Volcano, either Dept. Chimaltenango or
Dept. Sacatepequez); L. Wms, Phytologia 31: 438. 1976. Cacalia
cuspidata Bertol. Fl. Guat. 32. 1840, not S. cuspidatus DC., 1837
(type, Velasquez, Volcan de Agua, Sacatepequez). S. guatimalensis
Sch.-Bip. Flora 28: 499. 1845. S. godmanii Hemsl. Biol. Cent. Am.
Bot. 2: 240. 1881 (type, Salvin & Godman 327, Volcan de Agua,
Sacatepequez); Blake, Bull. Torr. Bot. Club 53: 218. 1926.
Montane forests and wet banks, 2,200-3,400 m.; Jalapa; El
Progreso; Chimaltenango; Sacatepequez; El Quiche; Solola;
Totonicapan; Quezaltenango; Huehuetenango; San Marcos. Mexico
(Chiapas).
Glabrous, perennial herbs to 2 m. tall, the stems striate, often purplish; leaves
410 FIELDIANA: BOTANY, VOLUME 24
membranaceous to subcoriaceous, the basal leaves usually long petiolate and hastate
or obtuse at the base of the blade, prominently bidentate-serrate, narrowly oblong to
triangular-ovate, acute or obtuse, the blade to 30 cm. long and 10 cm. broad, the
petiole to 20 cm. long, often dentate-alate, cauline leaves linear to lanceolate to
ovate-lanceolate, petiolate, becoming sessile, amplexicaul and much reduced above,
serrate, acute or acuminate, the base often hastate, the blade to 30 cm. long;
inflorescence an ample, diffuse, terminal corymb often with hundreds of heads; heads
discoid or rarely radiate, narrowly campanulate, about 1 cm. high; phyllaries about 8-
13, the common number being 13, acute, margins chartaceous, mostly 5-8 mm. long;
corollas pale yellow to yellow and (rarely) purple, slightly ampliate above at anthesis,
about 6 mm. long; style obtuse and obscurely penicillate; anthers exserted from the
corolla about 2 mm.; achenes black, glabrous, about 4 mm. long.
Common along streams in the montane or cloud forests on the
outer slopes of Tajumulco volcano, and other western volcanoes.
The species is a variable one in number of phyllaries in the
involucre. Two specimens with radiate heads were seen in this
normally discoid species.
Senecio epidendrus L. Wms. Phytologia 31: 440. 1975.
Cerro de Monos and Monte Virgin, Zacapa. Mexico (Chiapas).
Large epiphytic vines with araneose-floccose stems becoming; glabrous leaves
subcoriaceous, pubescent below, the blades oval, acute, 8-12 cm. long and 4.5-9 cm.
broad, petioles to 3 cm. long; inflorescence an axillary, multicapitate, corymbose
raceme; heads heterogamous, about 10-12 mm. long, with about 28 florets of which
about 5 are radiate; involucre with 8 narrowly oblong, acute phyllaries to about 5-6
mm. long; ray florets 10-12 mm. long with the small rays half that length; disc
corollas 8-9 mm. long, gradually expanded upward into a subcampanulate throat, the
lobes lanceolate and about 1 mm. long.
The single Guatemalan specimen assumed to be this species is
immature. It could prove to be distinct from S. epidendrus when
adequate material is at hand. Prominent nerves in the leaves help
to distinguish the plant from most other epiphytic species of
Senecio. Only one other species among the epiphytes has radiate
heads, that is S. morazensis Greenman from El Salvador and
Honduras, which has obscure nerves in the much smaller leaves.
Senecio gerberaefolius Sch.-Bip. in Hemsley, Biol. Cent. Am.
Bot. 2: 240. 1881.
Known in Guatemala only from the limestone summits of
Sierra de Cuchumatanes, 3,300-3,700 m. Huehuetenango. Mexico.
Scapose or subscapose, perennial herbs from a thick woody root, to about 20 cm.
tall, the scapes with much reduced leaves or bracts, densely floccose at the base;
leaves linear-lanceolate to oblanceolate, acute, attenuate into a long, winged petiole,
the margins revolute, densely silky-tomentose below, glabrous above, 5-15 cm. long
and 0.5-1.5 cm. broad; inflorescence of 1-3 heads at the apex of the scape; heads
WILLIAMS: FLORA OF GUATEMALA 411
calyculate, radiate and heterogamous, broadly campanulate, yellow; involucre 12-14
mm. long, with about 13 phyllaries, about as long as the disc flowers at anthesis;
phyllaries linear-lanceolate, acute to acuminate, glabrous, about 10-12 mm. long and
2 mm. broad; florets about 50, of which about 9 are radiate and pistillate, the
remainder perfect disc florets; ray corollas with a slender tube about 6 mm. long, the
ligule about 15 mm. long and 3 mm. broad, tridentate at the apex; disc corollas 8-9
mm. long, tubular at the base, expanded below the middle into a narrowly
campanulate throat, the lobes lanceolate, acute, about 1.5 mm. long; pappus
abundant, about as long as the disc corollas; achenes seen were immature, about 3
mm. long, pubescent.
There are only two small scapose Senecios in Guatemala, the
other is S. cuchumatanensis with oblong-ovate leaf blades. The
description above is based on Guatemalan specimens.
Senecio gilgii Greenm. Field Mus. Bot. 2: 282. 1907. Roldana
gilgii Robinson & Brettell, Phytologia 27: 419. 1974. Mano de Icon
de tierra fria; sivit.
Thickets and moist forests, 1,400-3,000 m.; Guatemala; Chimal-
tenango; El Quiche; Solola; Suchitepequez; Totonicapan; Quezal-
tenango; San Marcos. Mexico (Chiapas).
Tall, often much branched herbs or perhaps weak shrubs, 1.5-4 m. tall, the stems
densely lanate-tomentose; leaves suborbicular to broadly reniform, cordate, sinuately
many lobate, the lobes often triangular and acute, the margins mucronate-
denticulate, the blades palmately 5-11 nerved, hirtellous-pubescent on both sides to
densely lanate below, 9-25 cm. long and 15-40 cm. broad, the petioles to 22 cm. long;
inflorescence a several- multicapitate corymbiform panicle, densely reddish puberulent
with segmented hairs; heads large, 12-20 mm. long and 8-15 mm. broad,
heterogamous, usually with 7-8 ray florets and about 30 disc florets; involucre with
13 oblong or oblong-lanceolate, acute phyllaries, these densely puberulent dorsally,
the inner ones of the series with scarious margins, mostly about 1 cm. long at anthesis
and somewhat shorter than the heads; ray corollas to 10-15 mm. long including the 6-
7 mm. long ligule; disc corollas 7-9 mm. long, the lobes linear-lanceolate; pappus as
long as the disc corollas.
The relatively large heads, with 13 phyllaries and relatively
large number of florets, make this species easy to recognize. It is
probably almost always an herb.
Senecio grandifolius Less. Linnaea 5: 162. 1830. Roldana
grandifolia Robinson & Brettell, Phytologia 27: 427. 1974.
A wet forest species, mostly near 1,200-1,400 m.; southern
Mexico, known in Chiapas from Lago Tsiskaw on the Guatemalan
border.
Shrubs or trees to 12 m. tall; leaves lanceolate-oblong to oblong-ovate, acute or
acuminate, subentire to dentate-lobate or lobate, the blade to 30 cm. long and 20 cm.
412 FIELDIANA: BOTANY, VOLUME 24
broad, f loccose-tomentose to subglabrous below, usually glabrous above;
inflorescence and detail of heads and flowers much like those of S. arborescens.
This species and S. arborescens form an intergrading popu-
lation extending from southern Mexico to Panama. These and
related species need careful study to see whether only one species
and variations is present or whether two or more species are
represented.
Senecio greenmanii (Rob. & Brettell) L. Wms. Phytologia 31:
441. 1975. Roldana greenmanii Robinson & Brettell, Phytologia 27:
419. 1974.
Montane forests of the western slopes of the volcanoes of the
western highlands, mostly 1,400-1,800 m.; Retalhuleu (type, Strieker
394); Quezaltenango; San Marcos. To be expected in Mexico
(Chiapas).
Large herbs or weak treelike herbs to 8 m. tall, the upper stems hollow, striate,
puberulous; leaves largest of the genus (in Guatemala), long-petiolate or the
uppermost nearly sessile and reduced upward to bracts in the inflorescence, the blade
suborbicular to reniform in outline, usually cordate, deeply 5-7, incised-lobate, lobes
sometimes lobulate, the margins callose-denticulate, puberulous, especially below, to
quite glabrous, mature blades 17-25 cm. long and 25-45 cm. broad, the nerves palmate
from the apex of the petiole, petiole to 30 cm. or perhaps longer; inflorescence
puberulous throughout, a very large, multicapitate series of terminal and lateral
panicles, each panicle to about 30 cm. long; heads heterogamous, the involucre
mostly about 1 cm. long and 0.4-0.5 cm. broad at anthesis; phyllaries 8, linear or
linear-lanceolate, about 10 mm. long and 1.5-2 mm. broad, acute or obtuse,
puberulous on the midline dorsally, in 2 subseries, the inner broader and scarious
margined; florets discoid, mostly 22-24, about 8 marginal ones pistillate and the
remainder polygamous, at anthesis slightly longer than the phyllaries; corollas
obscurely puberulent on the tube, pistillate florets tubular and slightly expanded
above, mostly 8 mm. long with lobes 0.5 mm. long, corollas of polygamous florets
tubular but campanulate toward apex, about 10 mm. long with lobes about 1 mm.
long; anthers and styles exserted; pappus abundant, about as long as the corollas;
mature achenes not seen, glabrous, perhaps about 2 mm. long.
Easily distinguished among the Guatemalan species by the
narrow heads with eight phyllaries, the discoid heterogamous heads
and the enormous leaves.
Senecio heterogamus (Benth.) Hemsley, Biol. Cent. Am. Bot.
2: 242. 1881. Cacalia heterogama Benth. in Oersted, Medd. Vid.
Kjoebenhavn 1852: 107. 1853. Senecio heterogamus var. keller-
mannii Greenman, Field Mus. Bot. 2: 282. 1907 (type, Kellerman
4706 from Sacatepequez). Roldana heterogama Robinson &
Brettell, Phytologia 27: 420. 1974.
WILLIAMS: FLORA OF GUATEMALA 413
A species most often found in the wet and cold montane forests
of the western highlands, (1,800-) 2,300-3,600 m.; Guatemala;
Chimaltenango; Sacatepequez; El Quiche; Solola; Huehuetenango;
Totonicapan; Quezaltenango; San Marcos. Mexico (Chiapas).
Weak shrubs, 1-2.5 m. tall, often diffusely branched, the stems sparsely
pubescent to densely short-villous-hirsute; leaves usually peltate, suborbicular to
reinform in outline, mostly 7-9 lobate, the lobes subtriangular, acute or acuminate,
margins callose-denticulate, blades 6-25 cm. long and 8-30 cm. broad, densely
araneose-puberulent below to nearly glabrous, subglabrous or glabrous above;
petioles to 17 cm. long, but small upper leaves nearly sessile or with an alate petiole
and usually not peltate; inflorescences borne in axils of upper leaves or subterminal,
often diffuse, multicapitate panicles; heads mostly 1-1.5 cm. long at anthesis,
campanulate, heterogamous, involucres campanula te; phyllaries 10-13, linear-
lanceolate, acute, dorsally short-villous-hirsute about 7-10 mm. long; florets 20-30, all
discoid, 2-6 pistillate, the others perfect; corollas mostly about 10 mm. long, tubular,
the upper half in perfect flowers subcampanulate, in pistillate flowers narrower;
achenes 3-4 mm. long, mostly 10 ridged, glabrous; pappus about as long as the
corollas, abundant.
A variable species showing considerable variation in foliage and
in the heads. The leaves are usually peltate with the attachment
fairly near the base of the leaf, or the petiole is attached at the base
of the leaf, in ample specimens both conditions are usually found.
The known range of the species is curious. It is found on the
volcanoes of Costa Rica and adjacent Panama, and then skips to
the western highlands of Guatemala and adjacent Mexico, where it
is sometimes abundant.
Senecio jurgensenii Hemsley, Biol. Cent. Am. Bot. 2: 242.
1881. Roldana jurgensenii Robinson & Brettell, Phytologia 27: 421.
1974. R. breedlovei Robinson & Brettell, torn. cit. 415.
In moist mixed or coniferous forests, 2,000-3,500 m.; El
Progreso; Chimaltenango; El Quiche; Solola; Huehuetenango;
Totonicapan; Quezaltenango; San Marcos; Suchitepequez. South-
ern Mexico; El Salvador. Honduras.
Weak shrubs to 5 m. tall, the stems obscurely striate, hispidulous or glabrous
below, often with muricate processes which are caused by or invaded by a fungus;
leaves suborbicular to subreniform, 5 (-7) lobate, the margins callose-denticulate, the
lobes usually acute or acuminate, subpenninerved, hispidulous to glabrous, long-
petiolate and usually subcordate, the uppermost leaves sometimes sessile or with an
alate petiole, becoming bractlike in the inflorescence, the blades 5-15 cm. long and 6-
15 cm. broad, petioles from none to usually 7-15 cm. long; inflorescence a diffuse,
multicapitate, corymbose panicle from axils of uppermost leaves or bractlike leaves,
glandular-hispidulous, usually much exceeding the leaves in length; heads hetero-
gamous, clear yellow, 8-12 mm. tall; involucre mostly with 7-8 phyllaries,
campanulate, usually glandular-hispidulous; phyllaries linear-oblong to linear, acute,
414 FIELDIANA: BOTANY, VOLUME 24
about half in each head scarious margined, the others not, 5-8 mm. long, either about
half as long as the heads or nearly as long as the heads; flowers mostly 10-14 in a
head, 2-5 radiate, the others discoid; disc corollas cylindric, 6-8 mm. long, the lobes
lanciform; achenes striate, glabrous; pappus equaling the disc corollas.
Two forms or phases of this species are found, one with
phyllaries one-half to two-thirds as long as the heads and these
campanulate; the other with phyllaries narrower and about as long
as the subcylindric heads. The species is not abundantly distinct
from S. acutangulus (Bertol.) Hemsl., it is distinquished by the
glandular-hispidulous inflorescence, in contrast to glabrous or
slightly araneose ones of the latter.
The conspicuous muricate processes often on the stems are
perhaps insect and /or fungus caused.
Senecio lanicaulis Greenm. Field Mus. Bot. 2: 283. 1907.
Roldana lanicaulis Robinson & Brettell, Phytologia 27: 421. 1974.
Spetch.
Wet, mixed montane or cloud forests; 2,400-3,000 m.; El
Quiche; Huehuetenango (type, Heyde & Lux 3377). Mexico
(Chiapas).
Weak shrub, 2-7 m. tall, the stems densely lanate-tomentose, becoming glabrous;
leaves suborbicular to broadly reniform, cordate, palmately 5-7 nerved, 20-25 cm. long
and 20-28 cm. broad, shallowly lobulate and with prominent, mucronulate processes,
tomentulose becoming glabrescent above, densely and permanently lanate-tomentose
beneath, petioles stout, densely floccose, to 15 cm. long; inflorescence a multicapitate,
compound, terminal corymbose panicle usually exceeding the leaves; heads yellow,
heterogamous, to about 10 mm. long; involucre narrowly campanulate, calyculate,
tomentulose at the base, half or more as long as the head; phyllaries about 13, lance-
oblong or lanceolate, acute, the inner ones broadest and with scarious margins,
glabrous or nearly so, about 4-5 mm. long; ray flowers 6-8; disc flowers 10-20;
achenes striate, glabrous.
One of the most distinctive of the palmate nerved species in
Guatemala. The leaves are large and densely lanate-tomentose
below. Rare.
Senecio montidorsensis L. Wms. Phytologia 31: 441. 1975.
Telanthophora bartlettii Robinson & Brettell, Phytologia 27: 426.
1974, not Senecio bartlettii Greenm., 1950.
Probably a forest inhabiting species; British Honduras (type,
Bartlett 11852). Endemic.
A weak shrub to 3 m. tall, the stems at first pubescent becoming glabrescent;
leaves crowded on new growths, fleshy, the nerves obscure, oblanceolate or
oblanceolate-ovate, entire or denticulate, short-petiolate, the blades 9-20 cm. long and
WILLIAMS: FLORA OF GUATEMALA 415
3-8 cm. broad; petiole 2-3 cm. long; inflorescences terminal, long-pedunculate,
subumbellate-corymbose; heads subcylindric, 7-10 mm. long; involucre glabrous;
phyllaries 8, linear to linear-lanceolate, the inner with scarious margins, acute, 7-8.5
mm. long; florets 10-12, ray flowers about 3; disc corollas about 6-7 mm. long,
cylindric, the throat slightly expanded, the lobes linear, 3 mm. long, about as long as
the throat; achenes glabrous, about 2 mm. long; pappus ample.
A distinctive Senecio long since recognized by Greenman but
not published by him.
Senecio nubivagus L. Wms. Phytologia 31: 441. 1975. Cacalia
pudica Standl. & Steyerm. Field Mus. Bot. 23: 255. 1947, not
Senecio pudicus Greene, 1900. Pericalia pudica Cuatr. Brittonia 8:
157. 1955. Psacaliopsis pudica Robinson & Brettell, Phytologia 27:
408. 1974. Figure 133.
Moist or wet, often cloud-covered, alpine meadows, 3,400-3,700
m. Huehuetenango (type, Steyermark 48344). Endemic.
An erect, scapose, perennial herb to 30 cm. tall, the rootstock short, thick, with
many fibrous roots, the crown with sordid, floccose pubescence, the stem simple,
scapiform; leaves mostly radical, occasionally with reduced leaves on the scape, on
slender petioles to 14 cm. long, the petioles sparsely villous or glabrate, the blades
orbicular, 3-5 cm. broad, peltate near the middle, shallowly 7-lobate or the lobes
nearly obsolete, glabrous or sparsely villosulous below, the lobes short and rounded,
usually apiculate, the cauline leaf usually only one, reduced, more deeply lobate and
perhaps sometimes not peltate; peduncles thickened at the apex and somewhat
pubescent with segmented hairs; heads calyculate, nutant; involucre 10-12 mm. high
and 2-2.5 cm. broad, hemispheric; phyllaries 10-12, biseriate, lance-oblong to broadly
ovate, acute or obtuse, subequal or the outer ones narrower and shorter, villous or
glabrate; florets all discoid, about 50-60; corollas 7-8 mm. long, the tube cylindric and
about 2.5 mm. long, the throat narrowly campanulate, the lobes, short, triangular;
achenes glabrous, about 3 mm. long at anthesis; pappus bristles about 7 mm. long.
Distinctive among the Guatemalan Senecios by its nodding
heads and the relatively broad biseriate phyllaries. It is known from
only two collections, both from the high Cuchumatanes.
Senecio oerstedianus Benth. ex Oersted, Vid. Medd. Kjoeb.
1852: 109. 1853.
Open and forested slopes of the western volcanoes, 2,500-4,600
m.; San Marcos; Quezaltenango. Costa Rica; Panama; Mexico
(Chiapas).
Large, erect, terrestrial herbs from a thick root-stalk, the stems striate, 1-2 m.
tall, white-tomentose almost throughout, the upper leaf surfaces (rarely the lower
surfaces also) and stems becoming glabrate; basal leaves petiolate, ovate or oblong-
ovate, acute or acuminate, the base subcordate or attenuate, the blade serrate with
usually glandular-tipped teeth and to about 20 cm. long and 9 cm. broad, cauline
leaves reduced upward, the uppermost sessile and clasping, ovate to lanceolate, acute,
416 FIELDIANA: BOTANY, VOLUME 24
serrate, 25 cm. long or less; inflorescence a multicapitate corymbose panicle; heads 1-
1.5 cm. high and half as broad, heterogamous; involucre 6-10 long, glabrous or nearly
so; phyllaries about 8, linear-oblong with scarious margins and a thickened mid-rib,
acute or obtuse; corollas yellow, ligules or ray flowers small; disc corollas tubular, 7-
10 mm. long, subcylindric; styles subtruncate and penicilate; pappus about as long as
the disc corollas.
The range of the species in Guatemala is limited, as it is in
Costa Rica and Panama. The description above is based on
Guatemalan specimens, leaves on Costa Rican specimens are often
much larger. This is the only densely tomentose, herbaceous
Senecio in Guatemala.
Senecio parasiticus Hemsl. Biol. Cent. Am. Bot. 2: 224. 1881;
L. Wms. Phytologia 31: 444. 1975. Cacalia parasitica Sch.-Bip. ex
Hemsl. ll.cc., in synon.
Epiphytic vines in the Atlantic forests at middle elevations,
mostly about 1,500 m. Alta Verapaz. Mexico (Veracruz).
Epiphytic vines (not parasitic!) usually pendent; stems glabrous or nearly so,
somewhat fleshy; leaves lanceolate to elliptic-lanceolate, petiolate, somewhat fleshy,
the blades 6-9 cm. long and 1.5-2.5 cm. broad, petioles slender, to about 1 cm. long;
inflorescences mostly terminal on branches, racemose or paniculate, heads few to
many; heads discoid, monogamous, campanulate, about 10-12 mm. long; phyllaries
usually 8, linear-oblong, acute, coriaceous with scarious margins, 6-9 mm. long;
corollas with cylindric tube more than half the length of the whole, throat slightly
expanded and narrowly campanulate, lobes lanceolate, about 1 mm. long; achene
about 2 mm. long, glabrous; pappus about as long as the corollas.
Known from few collections but formerly thought to include
most of the epiphytic Senecios of the region. Senecio phoroden-
droides is a closely related Pacific slope species.
Senecio petasioides Greenm. in Donn.-Sm. Bot. Gaz. 37: 419.
1904; L. Wms., Phytologia 31: 444. 1975 (type from Santa Rosa,
Heyde & Lux 4522). Roldana chiapensis Robinson & Brettell,
Phytologia 27: 416. 1974. Hoja de queso.
Moist or wet forests or thickets, most often from December
through February but to May in Alta Verapaz, 1,400-3,200 m.; Alta
Verapaz; Baja Verapaz; Zacapa; Jutiapa; El Progreso; Guatemala;
Escuintla; Chimaltenango; Zacatepequez; Huehuetenango; Solola;
Totonicapan; Quezaltenango; San Marcos. Mexico; El Salvador;
Honduras; Nicaragua.
Weak shrubs to 2 m. tall, the stems subtomentose to almost glabrous; leaves
suborbicular to subcordate-reniform, the margins callose-denticulate, 5-9 lobate, the
lobes rounded and obtuse, subpenninerved, usually densely floccose-tomentose below,
rarely nearly glabrous, the upper surface hispidulous-puberulent to quite glabrous.
WILLIAMS: FLORA OF GUATEMALA 417
the blade 6-20 cm. long and 7-24 cm. broad, the petiole to 17 cm. long, the upper
leaves sometimes nearly sessile; inflorescence a diffuse, multicapitate, corymbose
panicle usually much exceeding the leaves, hispidulous or glandular-hispidulous;
heads heterogamous, 10-15 mm. long, conspicuously shorter than the disc flowers;
involucre with about 8 phyllaries; phyllaries 5-9 mm. long, linear-lanceolate to
lanceolate, acute, the outer 4 narrower than the inner scarious-marginate series, all
hispidulous dorsally on the cartilaginous portion; ray florets 2-4, with corollas to 15
mm. long but usually less, the lamina conspicuous and about 5 mm. long or much
reduced, fewer and inconspicuous, disc florets 10-14, the corollas subcylindric, slightly
expanded above, to 10 mm. long, the lobes linear-lanceolate, acute, about 1 mm. long;
mature achenes 10-ridged, glabrous, about 4 mm. long; pappus copious, about as long
as the disc corollas.
The name S. petasitis (Sims) DC. has often been applied to this
taxon, but the application is questionable. S. cristobalensis is
perhaps at most a variety of this.
Senecio phorodendroides L. Wms. Phytologia 31: 445. 1975.
Epiphytes in forests at 1,400-2,800 m. Escuintla (type, Standley
64554); El Quiche; Solola; Quezaltenango; San Marcos. Mexico
(Chiapas).
Epiphytic vines or shrubs, glabrous or nearly so, the stems striate; leaves
alternate, fleshy, petiolate, the veins inconspicuous, the blade lancolate to lanceovate,
acute or acuminate, 4.5-8 cm. long and 1.5-3 cm. broad; petioles slender, to 1.5 cm.
long; inflorescences terminal or lateral few-many-headed racemes; heads
homogamous, discoid, campanula te, 7-8 mm. long, florets about (10-) 20; involucre
calyculate, basically with 8 phyllaries, but these sometimes connate into fewer
unequal phyllaries lanceolate to ovate, glabrous, the tips obscurely penicillate, mostly
4-5 mm. long, much shorter than the heads; corollas 6-7 mm. long, tubular below and
somewhat expanded at the base, the throat narrowly campanulate, the lobes
lanceolate, about 1 mm. long; anthers and stigmas exserted; achenes ridged, glabrous,
about 2 mm. long; pappus ample, as long as the corollas.
Related to Senecio parasiticus Hemsl. from the Atlantic forests
of Veracruz and Alta Verapaz. S. parasiticus differs in having
phyllaries about as long as the heads.
Senecio quezalticus L. Wms. Phytologia 31: 446. 1975.
Damp mixed or Cypress forests, flowering well into the dry
season, 2,000-3,000 m.; Jalapa; Chimaltenango; Sacatepequez;
Quezaltenango (type, Standley 84286). Mexico (Veracruz, to be
expected in Chiapas).
Weak, simple or branched shrubs, 1-4.5 m. tall, stems sparsely floccose, becoming
glabrous, striate or obscurely ridged; leaves large and long-petiolate, broadly ovate-
cordate to suborbicular-cordate, mostly 5-7 lobate or lobulate and short mucronate-
denticulate, araneose-lanate below to glabrescent, penninerved but principal nerves
near the base, blades 5-23 cm. long and 5-15 cm. broad, the petioles 2-11 cm. long,
418 FIELDIANA: BOTANY, VOLUME 24
floccose to glabrescent; inflorescences multicapitate, pedunculate, corymbose panicles
from the axils of the upper leaves, about as long as the subtending leaves at anthesis
or slightly longer; heads heterogamous, 8-12 mm. long, campanula te, calyculate;
involucre with about 12 phyllaries; phyllaries linear-oblong, acute, canaliculate and
thickened at the base, about 5-6 mm. long; flowers yellow; ray flowers 6-8, pistillate,
the tube cylindric, about 4 mm. long, the lamina linear-oblong, obtuse, 5-6 mm. long;
disc flowers about 14-16, corolla subcylindric, somewhat ampliate above, about 7 mm.
long, the lobes linear-lanceolate, acute, about 2 mm. long; style about as long as the
anthers, the stigmas truncate; immature achenes ridged, glabrous, 1.6-2 mm. long;
pappus abundant, slightly shorter than the corollas.
Closely related to Senecio barba-johannis, but to be dis-
tinguished by the more prominently lobate, sparsely floccose leaves,
which are larger on the average.
Senecio rhyacophilus Greenm. Field Mus. Bot. 2: 280. 1907.
Wet montane forests, 2,400-3,000 m.; Sacatepequez (type,
Heyde & Lux 4502); El Quiche. Endemic.
Glabrous perennial herbs to about 1 m. tall, stems fragile, striate; leaves
runcinate-pinnate with unequal and irregular, mucronate-subdentate or sublobate
segments, paler beneath, membranaceous, basal leaves petiolate, 15-30 cm. long,
cauline leaves reduced upward, becoming sessile and amplexicaul; inflorescence a
terminal, few-headed panicle; heads discoid, 12-15 mm. high; involucre narrowly
campanula te, calyculate; phyllaries about 12-14, linear, acute, about 1 cm. long;
corollas about 7 mm. long, cylindric and slightly ampliate above, purplish; styles
obtuse and subpenicillate; anthers exserted from the corolla about 3 mm; achenes
glabrous, about 2-3 mm. long.
Closely related to Senecio callosus and perhaps not really
distinct.
Senecio salignus DC. Prodr. 6: 430. 1837. Barkleyanthus
salicifolius Robinson & Brettell, Phytologia 27: 407. 1974. Cineraria
salicifolia HBK. Nov. Gen. & Sp. 4: 189. 1820, not Senecio
salicifolius Pers., nor Wallr. Chilco; chilca; ch'homp.
Open slopes, thickets or open oak forests, 1,300-3,100 m.; Baja
Verapaz; Jalapa; Guatemala; Sacatepequez; Chimaltenango; Es-
cuintla; El Quiche; Huehuetenango; Totonicapan; Quezaltenango;
San Marcos. Mexico; El Salvador; to be expected in Honduras.
Erect, much branched, glabrous or nearly glabrous shrubs to 2.5 m. tall, leaves
linear to elliptic-linear, acute or acuminate, subplinerved, entire or obscurely
denticulate, tapered to a subpetiolar base, 3-13 cm. long and 0.2-1 cm. broad;
inflorescence a multicapitate, terminal and subterminal, corymbose panicle to about
10 cm. high; heads radiate, relatively small, yellow, narrowly campanulate, mostly 6-
8 mm. high at anthesis; involucres ebracteolate, glabrous, subglandular and usually
darker at the base; phyllaries about 8, lanceolate-ovate, acute, glabrous, scarious,
mostly about 4-5 mm. long; ray flowers about 5-6 mm. long, the lamina narrowly
WILLIAMS: FLORA OF GUATEMALA 419
oblanceolate; disc flowers about 15-20, the corollas subcylindric, mostly 5-6 mm. long;
pappus abundant, barbellate, about as long as the disc corollas.
Relatively common in the mountains at about 2,000 m. The
very narrow leaves, shrubby habit, small heads, and mostly open
habitat are distinctive among Guatemalan Senecios. The species
was first discovered in Guatemala by John Donnell Smith in 1890,
and found in recent years to be occasional to common during Field
Museum exploration of Guatemala.
Senecio schaffneri Sch.-Bip. Leopoldina 24: 126. 1888. S.
grandifolius var. glabrior Hemsl. Biol. Cent. Am. Bot. 2: 240. 1881.
S. ghiesbreghtii var. pauciflorus Coulter, Bot. Gaz. 16: 101. 1891
(type from Guatemala, Donnell-Smith 2359). S. santarosae Greenm.
Field Mus. Bot. 2: 281. 1907 (type from Santa Rosa, Heyde & Lux
4520). Roldana schaffneri Robinson & Brettell, Phytologia 27: 423.
1974. Quesillo de monte.
Damp, humid forests or thickets, 1,100-2,300 m.; Alta Verapaz;
Jalapa; Zacapa; El Progreso; Guatemala; Santa Rosa; Chimal-
tenango; Sacatepequez; El Quiche; Solola; Huehuetenango; Quezal-
tenango; Suchitepequez; San Marcos. Mexico; El Salvador;
Honduras.
Shrubs or weak trees 1-4.5 m. tall, the stems arachnoid-tomentulose, becoming
glabrous, leafy toward the apices of the new growths; leaves coriaceous, narrowly
elliptic-lanceolate to lanceolate-ovate, acute or acuminate, rarely shallowly sinuate-
dentate to usually entire, glabrous or nearly so, blade 8-20 (-30) cm. long and 2.5-8
(-12) cm. broad, petioles 2-6 cm. long; inflorescence of terminal or subterminal, long-
pedunculate, multicapitate corymbs; heads radiate or discoid, 5- flowered, 6-10 mm.
high, mostly about 8 mm., yellow; involucres glabrous to arachnoid-pubescent;
phyllaries 5, usually three oblong, obtuse and scarious margined, two narrower, acute
and not scarious margined; ray corollas pistillate, (2, 1 or none in each head), about 2
mm. long, the lamina elliptic-oblong, about 3 mm. long; disc corollas 6-7 mm. long,
cylindric, the lobes linear-lanceolate, about 2 mm. long; stamens exserted about 1
mm. at anthesis; achenes about 2 mm. long, glabrous; pappus slightly shorter than
the corolla.
A fairly distinctive species occasional in the montane forests, or
forest edges along open slopes. It is most commonly in flower from
January to April. The coriaceous leaves and phyllaries much
shorter than the heads help to distinguish it.
Senecio serraquitchensis Greenm. Field Mus. Bot. 2: 286.
1907. S. ghiesbreghtii var pauciflora Coulter, Bot. Gaz. 20: 52. 1895,
in part. Telanthophora serraquitchensis Robinson & Brettell,
Phytologia 27: 428. 1974.
420 FIELDIANA: BOTANY, VOLUME 24
A wet forest species, 800-1,500 m.; Alta Verapaz (type, Donnell
Smith 1598)', Suchitepequez. Endemic.
Weak shrubs to about 5 m. tall, the stems tawny-tomentose above; leaves
oblong-lanceolate to oblong-ovate, from somewhat sinuate- denticulate to dentate-
lobate, acute or acuminate, petiolate, the blade 9-25 cm. long and 3-15 cm. broad,
glabrous; petiole somewhat tomentose, 5-6 (-20) cm. long; inflorescences terminal, 1
or more long-pedunculate, compound, multicapitate corymbs; heads radiate,
narrowly campanulate, 8-10 mm. high at anthesis; florets 10-12, rays mostly 3-4; disc
corollas about 6 mm. long, with the tube cylindric, about 3 mm. long, the narrowly
expanded throat about 3 mm. long and the lobes linear and about 1 mm. long;
achenes glabrous, about 2 mm. long; pappus shorter than the disc corollas.
Related to Senecio arborescens Steetz, and perhaps only a form
with more nearly entire leaves.
Senecio steyermarkii Greenm. Ceiba 1: 124. 1950.
A forest species, 1,150 m.; Huehuetenango (type, Steyermark
49556). Endemic.
Shrubs to 6 m. or more tall, the young branches floccose-tomentulose, doubtless
glabrous with age; leaves elliptic-ovate to ovate, acute, long petiolate, floccose-
tomentose below, glabrescent above, the blade 6-15 (-20) cm. long and 4-10 cm. broad,
entire or obscurely denticulate, the petiole to about 8 cm. long; inflorescence
terminal, corymbose-cymose, multicapitate; heads heterogamous, yellow, about 6-8
mm. high; involucre campanulate, calyculate; phyllaries about 8, linear to linear-
oblong, 3-4 mm. long, half as long as the head; ray florets about 3, disc florets about
10; achenes glabrous.
Related to S. barba-johannis. It is known only from the type
collection, which was made in the vicinity of Barillas, botanically a
little known region of Guatemala.
Senecio uspantanensis (Coulter) Greenm. Field Mus. Bot. 2:
286. 1907. Senecio ghiesbreghtii var. uspantanensis Coulter, Bot.
Gaz. 20: 52. 1895. Telanthophora uspantanensis Robinson &
Brettell, Phytologia 27: 428. 1974.
Wet montane forests, 1,800-3,000 m.; Jalapa; El Quiche (type,
Heyde & Lux 3368); San Marcos. Mexico.
Weak shrubs or trees to 6 m. tall, the stems glabrous or nearly so; leaves elliptic-
oblong to lanceolate-oblong or oblong, acute or acuminate, long-petiolate,
prominently penninerved, the nerves to 18 pairs but usually fewer, the blades
glabrous, 15-30 cm. long and 3.5-11 cm. broad, petioles to 10 cm. long; inflorescences
terminal, long-pedunculate multicapitate, subumbellate corymbs; heads cylindric,
small, 6-8 mm. long, heterogamous; phyllaries 5, linear, acute, glabrous, about 6-7
mm. long; florets about 5, of which 1-2 are radiate; disc corollas about 6 mm. long,
the tube cylindric, 3-3.5 mm. long, the throat slightly expanded, about 3 mm. long,
the lobes linear, about 1 mm. long; achenes about 2 mm. long; pappus ample, slightly
shorter than the disc corollas.
WILLIAMS: FLORA OF GUATEMALA 421
A rather uncommon plant, closely allied to and part of the
complex surrounding Senecio cobanensis.
Senecio warszewiczii A. Br. & Bouche, Linnaea 25: 298. 1852.
Margarita de monte.
Open slopes, thickets, mixed or coniferous forests; 2,000-4,100
m.; Chimaltenango; Solola; Huehuetenango; Totonicapan; Quezal-
tenango; San Marcos. Mexico (Tacana volcano, Chiapas).
Herbs or subshrubs, 0.5-1.5 m. tall, the stems floccose to glabrate, striate, leafy to
the apex but the leaves reduced above; leaves linear to elliptic-linear to elliptic, acute
or acuminate, denticulate or entire, the margins usually somewhat revolute, petiolate
or not, the base of at least the upper leaves auriculate and often subamplexicaul,
blades 2-15 cm. long and 0.2-3 cm. broad, densely floccose-tomentose below, less so to
glabrate above; inflorescence a multicapitate, corymbose panicle, usually densely
floccose-pubescent, mostly 10-20 cm. high and nearly as broad; heads radiate, yellow,
about 8-13 mm. high and 5-8 mm. broad; involucres campanulate, floccose to
glabrate; phyllaries about 20, linear, acute, 6-8 mm. long, the margins often scarious;
ray flowers 7-8 mm. long, the lamina 3-4 mm. long, elliptic-oblong; disc flowers many,
5-6 mm. long, tubular; achenes terete and obscurely ridged, 2-3 mm. long, covered
with short, thick hairs ( ± 0.1 mm. long); pappus as long as the disc flowers.
Abundant especially at upper middle elevations ( ± 2,500 m.) in
the western highlands. The foliage is exceeding variable but the
plant may be distinguished easily from the nearest ally, S.
deppeanus, by the combination of auriculate leaves, scantier
pubescence on the lower leaf surface, and usually herbaceous or
only suffrutescent stems.
EXCLUDED SPECIES
SENECIO ERIOPHYLLUS Greenm. Field Mus. Bot. 2: 282. 1907.
Sterile and inadequate specimens from Guatemala have been
determined questionably by Dr. Sam Gibson as this species. Since
the species is known otherwise only from the state of Oaxaca in
Mexico, its extention to Guatemala is questionable.
SENECIO PETASITIS (Sims) DC. Prodr. 6: 431. 1837. Cineraria
petasitis Sims, Bot. Mag. 37: t. 1536. 1813.
This name, often applied to Guatemalan specimens, is of
dubious application. Most specimens so determined are Senecio
petasioides Greenm.
WERNERIA HBK.
Low, caespitose, perennial herbs glabrous or pubescent; leaves radical or crowded
on the very short scape, usually entire; heads heterogamous, radiate, large or
422 FIELDIANA: BOTANY, VOLUME 24
medium -sized, sessile among the leaves or rarely pedunculate; disc yellow, the rays
variously colored, white in the Guatemalan species; ray flowers uniseriate, pistillate;
disc flowers hermaphrodite, fertile; involucre broadly campanulate or hemispheric;
phyllaries uniseriate, equal, concrete at the base or higher, receptacle flat or convex,
naked; rays spreading, entire or nearly so; disc flowers regular, tubular, the limb
ampliate, cylindric, shallowly 5-fid; anthers entire at the base or with minute obtuse
auricles; style branches of the disc flowers truncate, penicillate, or with short
appendages; achenes oblong to turbinate, glabrous or villosulous; pappus bristles
numerous, slender, scabrous or rarely barbellate.
Species 30 or more, mostly in the Andes of South America, only
one in North America, two to three in the high mountains of Asia
and Africa.
Werneria nubigena HBK. Nov. Gen. & Sp. 4: 193. 1820. W.
mocinniana DC. Prodr. 6: 324. 1837 (type from Sierra de los
Cuchumatanes, Huehuetenango, Sesse & Mociho). Oribasia acaulis
Moc., Sesse & Cerv. ex DC. I.e., as synonym. Margarita. Figure 134.
Moist or wet alpine meadows, often in rocky places, frequently
on limestone, 3,300-4,300 m.; Huehuetenango (Sierra de los
Cuchumatanes); Quezaltenango (Volcan de Santa Maria); San
Marcos (volcanoes of Tajumulco and Tacana). Mexico; (Chiapas)
high mountains of the Andes from Ecuador to Bolivia.
Plants perennial from a thick, deep taproot, solitary or usually caespitose, the
caudex densely covered by the long-persistent leaf bases, these very densely covered
with long, white or yellowish wool; leaves broadly linear or liguliform, coriaceous, 10
cm. long or shorter, 5-10 mm. broad, rounded at the apex, entire, dilated and clasping
at the base, the blades glabrous; heads mostly solitary, appearing sessile in the middle
of the dense rosette of leaves, the yellow disc 2.5 cm. broad; flowers very numerous;
phyllaries numerous, linear-lanceolate, 1.5 cm. long or more, pubescent at the
narrowed obtuse apex, elsewhere glabrous, pale-marginate; rays white, 2.5 cm. long, 4
mm. broad, spreading.
This is perhaps the most handsome, and certainly one of the
most conspicuous, of the subalpine plants of Guatemala. It is
apparently not palatable to the sheep that are grazed in these
subalpine meadows. The great abundance of the plant may be due,
in part, to the overgrazing of the meadows by sheep, which results
in the reduction of other native plants.
On the wide meadows of Chemal, on the summit of the
Cuchumatanes, it is very abundant and showy, the large heads with
their long white rays, scarcely rising above the surface of the
ground, appearing like so many great daisies or stars through the
meadows. The flowers may be found, apparently, at almost any
season of the year. In the herbarium of Field Museum there is a
photograph of a copy of the original drawing of W. mocinniana,
NASH: FLORA OF GUATEMALA 423
which is a rather good representation of the plant. This species,
strangely enough, is not mentioned by Hemsley in Biologia
Central-Americana. De Candolle based his description upon the
drawing mentioned, and the plant apparently is not represented by
specimens in the Sesse and Mocino Herbarium. These collectors are
known to have visited Guatemala, but since their herbarium
specimens do not usually bear locality data, it is impossible to
determine which of their specimens were obtained there. Only in
the case of this species have we been able to determine a
Guatemalan locality at which specimens were collected by Sesse
and Mocino, near the end of the eighteenth century. The locality of
W. mocinniana is given as "in Mexico ad cacumen montis
Cuchumatanes." This is, of course, the alpine meadow now crossed
by the automobile road, a well-known locality on the long and
excessively arduous trail formerly followed by travelers on their
way from Mexico City to Guatemala City, perhaps the most
difficult route that it would have been possible to find by which to
proceed from one region to the other.
TRIBE IX. CYNAREAE
By DOROTHY L. NASH
Coarse, perennial or biennial herbs (in ours), commonly monoecious but rarely
dioecious, the stems simple or branched; cauline leaves alternate, often pinnatifid or
incised the margins usually dentate and more or less spinose and prickly, the basal
leaves usually numerous, often densely tomentose or lanate; inflorescences of solitary
heads or more or less cymose and sometimes paniculate; heads usually homogamous,
discoid (rarely with a single series of usually neutral ray flowers); involucres ovoid or
globose; phyllaries in several series, often spine-tipped; receptacles flat to subconic,
usually setiferous, rarely naked; marginal flowers enlarged or not, all corollas
tubular, 5-dentate, variously colored; anthers sagittate or caudate at the base, the
apices long-appendaged; style with a thickened ring at the base of the branches, this
often pubescent, the branches connate or bifid, short, obtuse; achenes (in ours)
glabrous, more or less angulate; pappus a crown of multiseriate, plumose bristles.
About 40 genera in both the Old and New World but only one,
Cirsium, is known in Guatemala.
CIRSIUM Miller. Thistle.
Reference: Fr. Petrak, Die mexikanischen und zentralameri-
kanischen Arten der Gattung Cirsium, Bot. Centralbl. Beih. 27 (2):
207-255. 1910.
Perennial or biennial herbs, simple or branched, usually tall and coarse, rarely
acaulescent; cauline leaves alternate, often decurrent at the base, serrate or
424 FIELDIANA: BOTANY, VOLUME 24
pinnatifid and with dentate lobes, the lobes and teeth usually tipped with sharp
spines, the basal leaves often numerous and conspicuous, the pubescence usually
lanate; plants commonly monoecious but rarely by abortion dioecious; heads
homogamous, large, solitary or aggregate at the ends of the branches or sometimes
disposed in large panicles, sometimes sessile among the basal leaves; involucres ovoid
or glabose; phyllaries multiseriate, imbricate, narrow, appressed at least at the base,
graduate, usually spine-tipped, the innermost sometimes colored and muticous, the
outermost sometimes spinose-pectinate; receptacle flat to subconic, densely setose;
corollas purplish, white, pinkish, or rarely yellowish, the tube slender, the limb
symmetrical or oblique, the 5 lobes narrow; stamens with filaments somewhat
papillose-pilose or rarely glabrous, the anthers sagittate at the base, the auricles
caudate, the apices long-appendaged; style branches linear or filiform, short or long,
subobtuse; achenes glabrous, somewhat compressed or 4-angulate, smooth or costate,
truncate or umbonate at the apex; pappus bristles multiseriate, plumose, affixed to a
basal annulus and deciduous with it.
Perhaps 200 species, widely distributed in both hemispheres,
most numerous in temperate regions; in the tropics usually
confined to the mountains. Seven are in Guatemala and two other
species are known from southern Central America.
True involucre surrounded by erect, bractlike, spinose leaves.
Plants often to 1 m. tall; outer phyllaries spinose-pectinate C. horridulum.
Plants acaulescent or nearly so; outer phyllaries not spinose-pectinate.. ..C. skutchii.
True involucre not surrounded by erect, bractlike leaves.
Leaves deeply pinnatifid, often almost or quite to the costae; outer phyllaries not
spinose-pectinate.
Leaf blades decurrent at the base as wings on the stem C. mexicanum.
Leaf blades clasping to auriculate at the base but not decurrent as wings on the
stem.
Phyllaries glabrous or nearly so, commonly blackish when dried C. nigriceps.
Phyllaries densely and laxly arachnoid-tomentose with long, spreading hairs,
at least on the margins, not blackish when dried.
Heads 3.5-4.5 cm. high, 6-9 cm. broad; corolla lobes 11-14 mm. long.
C. consociatum.
Heads 2.5-4 cm. high but only 3-5 cm. broad; corolla lobes 5-8 mm. long.
C. radians.
Leaves pinnatifid but not close to the costae; outer phyllaries spinose-pectinate.
C. subcoriaceum.
Cirsium consociatum Blake, Brittonia 2: 355. 1937.
Open alpine meadows or limestone hillsides, sometimes in rocky
forest of Abies or Juniperus, 3,300-3,700 m.; Huehuetenango (type
collected in the Sierra de los Cuchumatanes, along trail between
Huehuetenango and Soloma, probably about Chemal, Skutch 1211).
Perennials from a thick root, the stems very thick, mostly 20-60 cm. tall but
sometimes to 1.5 m., or rarely dwarfed and only about 6 cm. high, simple below,
bearing few or rather numerous heads, densely arachnoid-lanate; basal and lowest
cauline leaves lanceolate, mostly 10-50 cm. long and as much as 10 cm. wide, deeply
NASH: FLORA OF GUATEMALA 425
pinnatisect, often almost or quite to the costae, with up to 15 pairs of segments, these
broadly triangular, to 5 cm. long, usually 2-3-furcate, the segments tipped with a
stout, stramineous spine to 1 cm. long, the blades thinly or densely arachnoid-
tomentose above, glabrate in age, densely grayish-tomentose beneath; heads nutant,
solitary or aggregate at the ends of the branches; involucres hemispheric, 3.5-4.5 cm.
long, 6-9 cm. broad; phyllaries maroon, straight, stiff, gradually acuminate into a
spine 2-4 mm. long, with narrow, pale margins, glabrous or nearly so except for the
very abundant, arachnoid ciliation, this more or less covering the involucre like wool;
corollas whitish to lavender, about 27 mm. long, the lobes 11-14 mm. long; achenes
oblong, about 5 mm. long, glabrous; pappus about 2.5 cm. long, whitish.
Cirsium horridulum Michx. Fl. Bor. Am. 2: 90. 1803. Cnicus
horridulus Pursh, Fl. Am. Sept. 2: 507. 1814. Cirsium horidulus var.
elliottii Torrey & Gray, Fl. N. Am. 2: 460. 1843. Cnicus chrismarii
Klatt, Leopoldina 25: 108. 1889. Carduus vittatus Small, Bull. N. Y.
Bot. Gard. 3: 349. 1905. Cirsium chrismarii Petrak, Bot. Centralbl.
Beih. 27 (2): 215. 1910. C. horridulum subsp. chrismarii Petrak, Bot.
Tidssk. 31: 58. 1911. C. pinetorum Small, Fl. Miami 199. 1913 (non
C. pinetorum Greenm. 1905). C. vittatum Small, loc. cit. C.
horridulum subsp. megacephalum Petrak, Bot. Centralbl. Beih. 35
(2): 384. 1917. C. horridulum f. elliottii Fern. Rhodora 45: 353. 1943.
C. horridulum var. vittatum R. W. Long, Rhodora 72: 45. 1970.
Cardosanto.
Open, marshy hillsides, swampy valleys, or dry, rocky hills,
sometimes in pine-oak forest, 300-1,750 m.; Alta Verapaz; Baja
Verapaz; Guatemala; Huehuetenango; Peten. Eastern and southern
United States; Mexico.
Stout biennial herbs to about 1 m. tall, the stems thick, striate, arachnoid-
tomentose, glabrate, or essentially glabrous, rather densely leafy, usually simple,
bearing 1-few heads; basal leaves oblong-lanceolate, shallowly incised-lobate and
sinuate-dentate, the margins with rather stout, pale or yellowish spines to about 1
cm. long, both surfaces laxly arachnoid-tomentose or essentially glabrous, to about 45
cm. long, the cauline leaves similar but smaller, sessile and clasping at the base, not
decurrent on the stem, with longer marginal spines to 1.5 cm.; heads 1-5, crowded,
short-pedunculate; involucres campanulate, commonly 3-5 cm. long, 4-8 cm. broad,
surrounded by erect, bractlike, lanceolate leaves, the margins densely beset with long,
stout spines; outer phyllaries lanceolate or linear-lanceolate, scaberulous, spine-
tipped and pectinate-ciliate, the next inner ones merely spinose-ciliate, the innermost
ones lance-linear, attenuate into a weak, soft tip; corollas rose-purple, creamy white,
or yellowish, the tube about 28 mm. long, the limb 15-17 mm. long; style long-
exserted; achenes pale yellowish brown, lustrous, glabrous, 4-5 mm. long.
A variable complex; Petrak's alleged differences between C.
horridulum and his var. chrismarii do not hold true. Both glabrous
and densely arachnoid-tomentose forms have been collected in some
of the southeastern United States, as well as in Mexico and
426 FIELDIANA: BOTANY, VOLUME 24
Guatemala, and there is wide variation in length of spines and in
corolla color.
Cirsium mexicanum A. DC. in DC. Prodr. 6: 636. 1837. Cnicus
mexicanus Hemsl. Biol. Centr. Am. Bot. 2: 251. 1881. Carduus
mexicanus Greene, Proc. Amer. Acad. 1892: 363. 1893. Cirsium
mexicanum var. bracteatum Petrak, Bot. Centralbl. Beih. 27 (2):
231. 1910. Cardosanto; cardosanto macho (Peten); alcachofa,
suctzun (Coban, Alta Verapaz: Quecchi). Figure 135.
Damp or wet fields or thickets, frequent in pastures, sometimes
in open pine forest, often on limestone, near sea level to 2,100 m.,
but mostly at 900-1,700 m.; Alta Verapaz; Baja Verapaz;
Chiquimula; Guatemala; Huehuetenango; Izabal; Jalapa; Peten;
Quezaltenango; El Quiche; Retalhuleu; Sacatepequez; Santa Rosa;
Zacapa. Southern Mexico; British Honduras to El Salvador;
Nicaragua; Costa Rica.
Tall, coarse herbs, probably biennial but perhaps perennial, 1-2.5 m. high, simple
or branched, the stems thick, striate, at least partially winged by the decurrent
petiole bases; basal leaves often 40 cm. long or more, on broadly winged, very spiny
petioles, the lower cauline leaves similar, amplexicaul and decurrent on the stem,
oblong or lance-oblong, deeply pinnatifid, often almost or quite to the costae, thinly
villous or glabrate above, whitish or grayish beneath and covered with usually dense,
lax or appressed, arachnoid tomentum, the lobes oblong or triangular, usually
coarsely dentate, the lobes and teeth tipped with slender, yellowish, short or
elongated spines, the upper leaves much smaller and narrower, less conspicuously
decurrent; heads few or numerous, aggregate, short-pedunculate, sometimes disposed
in large, broad panicles; involucres hemispheric, about 3.5 cm. high; phyllaries linear-
lanceolate, subcoriaceous, closely arachnoid-tomentose or almost glabrous, greenish,
the outer ones short, usually somewhat spreading or reflexed, tipped with a slender
spine 4-8 mm. long, this usually somewhat curved and spreading, the inner phyllaries
much longer, long-acuminate but scarcely spine-tipped; corollas pink or pale purple,
about 30 mm. long, the linear lobes about 5 mm. long; achenes glabrous, dark brown,
3-3.5 mm. long; pappus bristles dirty white, long and very abundant.
Although widely distributed in Guatemala, these thistles are
seldom abundant in a given locality.
Cirsium nigriceps Standl. & Steyerm. Field Mus. Bot. 23: 258.
1947.
Damp, mixed forest, 3,300-3,700 m.; Huehuetenango (type from
Sierra de los Cuchumatanes, between Tejquia and Caxin bluff,
Steyermark 50157). Mexico (Chiapas).
Large, coarse, erect herbs commonly 1-2 m. tall, the stems about 1 cm. thick
above, not winged, rather densely and laxly arachnoid-tomentose above; cauline
leaves sessile, clasping by a dilated base, not decurrent on the stem, linear-oblong in
NASH: FLORA OF GUATEMALA 427
outline, mostly 10-18 cm. long, viscid-villosulous or puberulent or glabrate above,
paler beneath and laxly arachnoid-tomentose on costae and veins, elsewhere glabrous
or nearly so, the blades deeply pinnatifid, often almost to the costa, the lobes about
15 pairs or fewer, triangular or triangular- lanceolate, often trilobate and usually
spinose-serrate, acute or acuminate, terminated by a slender but rigid, stramineous,
erect spine to about 7 mm. long; heads sessile or short-pedicellate and several
clustered at the ends of the few short branches, often disposed in a leafy panicle to
about 22 cm. long, involucrate by a few inconspicuous, deeply spinose-lobulate,
narrow bracts; involucres broadly campanulate or hemispheric, blackish when dried,
about 2 cm. high and broad; phyllaries numerous, appressed, graduate, lance-linear,
as much as 1.3 mm. wide, long-attenuate, ending in an erect, pale, stiff spine scarcely
2 mm. long, almost wholly glabrous, minutely ciliolate, glabrous dorsally or
practically so, eglandular; corolla pale purple, glabrous outside, the tube almost
filiform, about 12 mm. long, slightly dilated in the short throat, the lobes linear,
spreading, about 3.5 mm. long; achenes subcompressed, glabrous, cuneate-oblong,
about 4 mm. long; pappus bristles plumose, sordid, about 15 mm. long.
Cirsium radians Benth. PI. Hartweg. 77. 1841. Cnicus radians
Hemsl. Biol. Centr. Am. Bot. 2: 252. 1881. Cirsium guatemalense
Blake, Journ. Wash. Acad. Sci. 24: 442. 1934 (type from Chimal-
tenango, Skutch 660). Cardon (Chimaltenango).
Open meadows, fields, roadsides, or sometimes in open pine-oak,
Alnus, or Cuppressus forest, 2,400-3,300 m.; Alta Verapaz; Chimal-
tenango; Huehuetenango; Quezaltenango (type from Quezal-
tenango, Hartweg 538); San Marcos; Totonicapan.
Coarse, stout herbs to about 1.5 m. tall, simple below or sparsely branched, the
stems very thick, arachnoid-tomentose or glabrate, not winged; cauline leaves sessile
and somewhat clasping but not decurrent on the stem, the blades narrow, mostly 5-15
cm. long, deeply pinnatifid, often nearly or quite to the rachis, the segments short,
oblong or very narrowly triangular, often 2-3-lobate, the lobes acuminate,
terminating in a long, sharp, stout, yellowish spine, laxly arachnoid-tomentose above
or in age glabrate, white or grayish beneath with very dense arachnoid tomentum;
heads solitary or several at the ends of the branches, nutant, 2.5-4 cm. high 3-5 cm.
broad; phyllaries purplish, the outer ones arachnoid-lanate, subcoriaceous, rigid,
linear-lanceolate, entire, usually more or less recurved, tipped with a long, spreading,
slender spine, the inner phyllaries mostly shorter, erect, linear-attenuate, tipped with
a short, flexible spine; corollas rose-purple or lavender, the lobes linear, 5-8 mm. long;
pappus very abundant, dirty white.
Cirsium skutchii Blake, Brittonia 2: 357. 1937.
Damp or wet alpine meadows, or on limestone rocks in
Juniperus forest, 3,150-3,750 m.; endemic; Huehuetenango (type
collected along trail between Huehuetenango and Soloma, Sierra de
los Cuchumatanes, probably in the region of Chemal, Skutch 1264).
Plants perennial from short, thick rhizomes, acaulescent or nearly so, the stems
commonly less than 6 cm. high; leaves basal, very numerous and forming large, dense
rosettes, petiolate, mostly 5-20 cm. long, narrowly oblong in outline, pinnate-lobate,
428 FIELDIANA: BOTANY, VOLUME 24
the segments numerous, triangular or quadrangular, terminated by a slender,
yellowish spine 7-14 mm. long, glabrous on the upper surface, sparsely arachnoid-
tomentose beneath or in age glabrous or nearly so; plants dioecious; heads short-
pedunculate, few or numerous, crowded at the base of the leaves, about 3.5 cm. high;
involucres broadly campanulate or hemispheric, surrounded by a few erect, bractlike
leaves; phyllaries about 6-seriate, strongly graduate, glabrous or nearly so, the outer
ones tipped with a stout, yellowish spine 3-5 mm. long, the inner ones sometimes
inconspicuously hispidulous, the innermost with narrowly lanceolate, stramineous,
serrulate tips; corollas purplish white, the tube about 18 mm. long, the limb 4 mm.
long, the teeth 2.5-3 mm. long; achenes 4-5 mm. long, dark brown, glabrous, lustrous;
pappus brownish, copious, about 3 cm. long.
One of the characteristic alpine plants of the Chemal region.
Cirsium subcoriaceum (Less.) Sch. Bip. in Seem. Bot. Voy.
Herald 312. 1856. Carduus subcoriaceus Less. Linnaea 5: 130. 1830.
Cirsium heterolepis Benth. PL Hartweg. 87. 1841 (type from
mountains of Mixco, Guatemala, Hartweg 596). Cirsium maximum
Benth. op. cit. 289. Cnicus heterolepis Gray, Proc. Amer. Acad. 10:
40. 1875. Cnicus subcoriaceus Hemsl. Biol. Centr. Am. Bot 2: 252.
1881. Cardosanto; espina (Guatemala); aman, tun k'is, sa qi paxl
(Quecchi, Alta Verapaz).
Damp or wet, open fields, thickets, or open forest, sometimes in
pine, oak, Cupressus, or Alnus forest, 1,200-3,500 m., commonly at
rather high elevations; Alta Verapaz; Chimaltenango; Guatemala;
Huehuetenango; Jalapa; Quezaltenango; El Quiche; Sacatepequez;
San Marcos; Santa Rosa; Solola; Totonicapan; Zacapa. Mexico;
Honduras; El Salvador; Costa Rica.
Plants probably perennial, 1-3 m. tall, often much branched above, the stout
stems arachnoid-tomentose; basal and lower cauline leaves often 50 cm. long or more,
the basal ones petiolate, the cauline ones sessile and more or less clasping, not
decurrent on the stem, oblong-lanceolate to elliptic, pinnatifid, the lobes oblong,
coarsely dentate, the lobes or teeth tipped with long, stout spines, arachnoid-
tomentose above when young, in age scabrous or somewhat villous- hispid, whitish or
grayish beneath with dense arachnoid tomentum; heads usually solitary, occasionally
2-4 at the ends of the branches, more or less nutant, 4-7 cm. broad; involucres
orange-red or reddish yellow; phyllaries arachnoid-tomentose with a lax, spreading
tomentum, the outer ones subreflexed, lanceolate, spine-tipped, spinose-pectinate, the
inner phyllaries much longer, erect, much longer than the flowers, linear, entire;
corollas rose-purple, the limb 5-parted almost to the base, about twice as long as the
tube; achenes dark brown, lustrous, 6-7 mm. long; pappus dirty white.
Common, showy, rather handsome plants in many regions of
Guatemala, particularly plentiful in the upper pine zones on the
high volcanoes, or at the upper edge of the dense, wet, mixed forest.
NASH: FLORA OF GUATEMALA 429
TRIBE X. MUTISIEAE
By DOROTHY L. NASH
Perennial herbs or shrubs, monoecious or dioecious, various in habit, erect to
sometimes scandent, sometimes acaulescent; cauline leaves alternate, sessile or
petiolate, the blades entire, lobate or dentate; inflorescences of solitary heads or
cymose and sometimes corymbose, thyrsoid, or paniculate; heads homogamous or
heterogamous; involucres cylindrical or turbinate to hemispheric; phyllaries 2-several-
seriate; ray flowers (when present) ligulate, bilabiate; disc flowers tubular but
bilabiate; anthers caudate at the base; style branches usually short, sometimes
elongated, truncate or obtuse; achenes various, mostly oblong, linear, or fusiform,
usually costate; pappus bristles numerous in 1 -several series, simple or plumose.
More than 50 genera, more common in the New World.
Plants dioecious; pistillate heads larger than the staminate ones Lycoseris.
Plants not dioecious.
Heads heterogamous; achenes rostrate Chaptalia.
Heads homogamous (in ours); achenes not rostrate.
Leaves on petioles mostly 2-10 cm. long.
Plants scandent (in ours); pappus bristles uniseriate Jungia.
Plants erect (in ours); pappus bristles biseriate to multiseriate Onoseris.
Leaves sessile and often auriculate-clasping, or on short petioles less than 1 cm.
long.
Leaves on very short petioles or subsessile; phyllaries 1-2-seriate; corollas
bright yellow Trixis.
Leaves mostly sessile, often auriculate-clasping; phyllaries 2- multiseriate;
corollas white, pink, or purple (in ours).
Flowering stems scapiform, leafless or the few leaves reduced and bractlike;
basal leaves commonly runcinate-pinnatifid, at least near the base;
indument of basal portion of stem long and white, or lacking Perezia.
Flowering stems (in ours) very leafy; basal leaves not runcinate-pinnatifid;
indument of lower stems composed of tufts of brown, wooly tomentum.
Acourtia.
ACOURTIA D. Don
References: Rimo Bacigalupi, A monograph of the genus
Perezia, section Acourtia, with a provisional key to the section
Euperezia, Contr. Gray Herb. 97: 1-81, t. 1-7. 1931. James L. Reveal
and Robert M. King, Re-establishment of Acourtia D. Don
(Asteraceae), Phytologia 27: 228-232. 1973.
Perennial herbs or rarely suffruticose plants with a tuft of dense, wooly or
feltlike, rusty tomentum on the lower stem and sometimes in the leaf axils; leaves
alternate, coriaceous, mostly sessile, sometimes pinnatifid or variously lacerate or
dentate; heads homogamous, few-many-flowered, usually disposed in cymes, the
inflorescences often becoming paniculate, rarely the heads borne individually;
phyllaries 2-several-seriate, imbricate, often mucronulate; flowers hermaphrodite and
430 FIELDIANA: BOTANY, VOLUME 24
fertile; corollas white, pink, or purplish, bilabiate (3 of the segments fused to form the
outer ligulate lip); anthers long-caudate at the base, tipped with a rigid, colored
appendage; style branches flattened, papillose at the apex; achenes linear-cylindric to
fusiform, inconspicuously costate, glabrous or with glandular trichomes; pappus
bristles scabrous, mostly 1-2-seriate, rarely 3-seriate.
About 40 species, all American, with three in Guatemala.
Plants weak, usually scrambling or reclining; leaves very thin and flexible;
inflorescences small, scattered in upper leaf axils, the cymes consisting of few
heads on short, slender, leafy peduncles A. carpholepis.
Plants erect, rigid; leaves thick and coriaceous or if thin, somewhat rigid;
inflorescences large, terminal, paniculate.
Stems essentially glabrous; phyllaries ovate to oblong, obtuse; heads 4-5-flowered.
A. reticulata.
Stems glandular-puberulent; phyllaries linear- lanceolate, acute; heads 10-20-
flowered A. coulteri.
Acourtia carpholepis (A. Gray) Reveal & King, Phytologia
27: 229. 1973. Perezia carpholepis A. Gray, Proc. Amer. Acad. 19:
60. 1883. Figure 136.
Damp or dry, brushy hillsides, in mixed or oak forest, 1,400-
2,200 m.; Huehuetenango; Jalapa; Sacatepequez; Solola. Southern
Mexico.
Perennials, the stems weak, usually scrambling or reclining, sometimes more
elongated and somewhat scandent, slender, flexuous above, slightly scaberulous or
smooth; leaves numerous, chartaceous, sessile, lance-oblong to lance-ovate or broadly
elliptic-obovate, mostly 5-14 cm. long, 2-7.5 cm. wide, acuminate, deeply cordate at
the base with mostly rounded auricles, the margins rather finely spinulose-dentate,
along glabrous, minutely puncticulate beneath, conspicuously reticulate-veined;
cymes of few heads, on short, slender, leafy peduncles arising in the leaf axils and
shorter than the leaves, the peduncles puberulent; heads sessile or short-pedicellate,
10-11-flowered, 13-15 mm. high; phyllaries glabrous, broad, obtuse or rounded at the
apex, ciliate, gradually imbricate; corollas lilac; achenes about 5 mm. long, glandular,
glabrate or puberulent; pappus yellowish.
Acourtia coulteri (A. Gray) Reveal & King, Phytologia 27:
229. 1973. Perezia coulteri A. Grav. Proc. Amer. Acad. 15: 40. 1880.
Reported by Bacigalupi from Huehuetenango, on sparsely
wooded, calcareous slopes, oak forest, Chacula, 1,600 m., C. & E.
Seler 3069. Southern Mexico.
Stout, stiff, erect perennials, about 75 cm. tall, the base of the stems densely
covered with brown, feltlike tomentum, the stems striate, glandular-puberulent, leafy
throughout; leaves sessile or nearly so, chartaceous, oblong-lanceolate or oblong-
oblanceolate, mostly 5-10 cm. long, 1-2.5 cm. wide, acute or obtuse, somewhat
attenuate to rounded or subtruncate at the base, repand-denticulate or subentire,
glandular-puberulent, especially beneath; inflorescences large, corymbiform, 6-15 cm.
broad; heads numerous, 10-20-flowered; involucres 6-10 mm. high; phyllaries linear-
NASH: FLORA OF GUATEMALA 431
lanceolate, acute, usually ciliate, somewhat puberulent or glabrate; corollas lavender-
pink; achenes glandular-puberulent, striate, 5-6 mm. long.
Acourtia reticulata (Lag. ex D. Don) Reveal & King,
Phytologia 27: 231. 1973. Proustia reticulata Lag. ex D. Don, Trans.
Linn. Soc. 16: 200. 1830. Perezia reticulata A. Gray, PI. Wright. 1:
128. 1852. Te de monte (Guatemala, fide Aguilar).
Brushy slopes or in rather dry, oak forest, 1,100-1,800 m.;
Chimaltenango; Guatemala; Huehuetenango; El Quiche. Southern
Mexico.
Stout, erect perennials to 1 m. tall, glabrous or nearly so, the stems densely
leafy; leaves sessile, coriaceous, cuneate-obovate to elliptic- oblanceolate, mostly 6-12
cm. long and 1.5-5 cm. wide, rounded to acute at the apex, gradually narrowed to the
base or becoming somewhat auriculate and clasping at the base, closely reticulate-
veined, usually glandular-puncticulate and puberulent on the veins beneath, the
margins spinulose-dentate; inflorescences terminal, large, usually dense, rarely lax,
thyrisiform; heads sessile or short-pedicellate; involucres 4-6-flowered, 8-10 mm. high;
phyllaries ovate or oblong, obtuse or rounded at the apex, glabrate, ciliate; corollas
pale lavender-pink or white; achenes costulate, 4-5 mm. long, stipitate-glandular;
pappus yellowish white, abundant.
CHAPTALIA Ventenat
Reference: Arturo Burkart, Estudio del genero de Compuestas
Chaptalia, Darwiniana 6: 505-594. 1944.
Acaulescent, perennial herbs; leaves basal, sessile or petiolate, the blades entire,
dentate, or lyrate, white- to mentose beneath; scapes bearing a single head, naked or
with subulate bracts above; heads heterogamous, narrow or broad, erect or often
nutant; involucres campanulate or turbinate; phyllaries multiseriate, narrow,
acuminate, imbricate, graduated; outer flowers bi-multiseriate, pistillate, fertile, their
corollas ligulate, the lamina spreading, subdentate, tridentate or trilobate, white,
violaceous, or reddish; disc flowers hermaphrodite, fertile, or the inner ones or all of
them sterile, the corollas tubular, the limb bilabiate; anthers sagittate at the base,
the auricles long-caudate; style branches of the hermaphrodite flowers linear, obtuse;
achenes oblong or fusiform, 4-6-costate, more or less attenuate into a beak; pappus
bristles numerous, scabrous or barbellate.
Burkart divided the genus into seven sections, recognizing 30
species in the first six sections; of his seventh, Microchaptalia, he
states, "No he dispuesto de material suficiente para estudiar bien
los representantes de este grupo. El numero total de especies
descriptas que a mi juicio deben incluirse en el, es de 23, pero tengo
la impresion que esta cifra debe reducirse." Chaptalia is a genus of
tropical and subtropical America, with only two species known in
Guatemala.
432 FIELDIANA: BOTANY, VOLUME 24
Leaves sessile or short-petiolate, the blades rather thick, the margins entire to
somewhat crenate or inconspicuously undulate-lobate below; flower heads
always erect; beaks of achenes about as long as or sometimes 1-1.5 times longer
than the body C. dentata.
Leaves usually long-petiolate, rarely sessile, the blades very thin, the margins
commonly conspicuously and deeply lyrate-lobate below; flower heads nutant
when young and when in fruit, erect only at anthesis; beaks of achenes 2-3 times
longer than the body C. nutans.
Chaptalia dentata (L.) Cass. Diet. Sci. Nat. 26: 104. 1823.
Tussilago dentata L. Sp. PL ed. 2: 1213. 1763. T. albicans Swartz,
Fl. Ind. Occ. 3: 1348. 1806. Leria albicans A. DC. in DC. Prodr. 7:
42. 1838. L. leiocarpa A. DC., loc. cit. Chaptalia albicans Vent, ex
Steud. Norn. Bot. ed. 2, 1: 344. 1840. C. leiocarpa Urban, Symb.
Ant. 8: 747. 1921.
Grassy or rocky, open places, 1,200-2,100 m.; Chiquimula;
Huehuetenango; Jalapa; Santa Rosa; Zacapa. Southern United
States; Mexico; Honduras; West Indies.
Acaulescent perennials, prostrate or ascending; leaves sessile or short-petiolate,
the blades rather thick, oblanceolate or oblong-oblanceolate, mostly 3-10 (-20) cm.
long, obtuse or acute, attenuate to the base, the margins entire, repand-dentate,
crenate, or sometimes shallowly undulate-lobate, glabrous above or very sparsely
floccose-tomentose, densely white-tomentose beneath, the tomentum closely ap-
pressed; scapes slender, 8-35 cm. long, floccose-tomentose, the bracts small and
subulate or none; heads erect; involucres 1.5-2 cm. high at maturity; phyllaries linear,
attenuate, floccose-tomentose, tinged with purple above; ligulate corollas white or
pinkish, the lamina tridentate; achenes glabrous or pubescent, 4-6-costate, the body
4-6 mm. long, the filiform beak about as long as or 1-1.5 times longer than the body;
pappus white, about 1 cm. long.
Chaptalia nutans (L.) Polak, Linnaea 41: 582. 1877. Tussilago
nutans L. Syst. ed. 2: 1214. 1759. Leria lyrata Cass. Diet. Sci. Nat.
26: 102. 1823. L. nutans A. DC. in DC. Prodr. 7: 42. 1838. Valeriana
(Izabal). Figure 137.
Damp or wet thickets or open forest, often in pine forest,
sometimes in open fields; near sea level to 1,950 m.; Alta Verapaz;
Chimaltenango; Chiquimula; Escuintla; Huehuetenango; Izabal;
Quezaltenango; Retalhuleu; Santa Rosa. Mexico; British Honduras
to El Salvador and Panama; West Indies; South America.
Acaulescent perennials; leaves usually long-petiolate, rarely sessile, the blades
oblanceolate-oblong to oblong-ovate, mostly 8-30 cm. long, very thin, acute or obtuse,
commonly deeply lyrate-lobate below (rarely only crenate-lobate), the terminal lobe
almost entire or repand-denticulate, the lower lobes few and small, glabrous or
glabrate above, densely floccose-tomentose beneath, the tomentum loose, white or
grayish; scapes slender, mostly 15-50 cm. high, floccose-tomentose, ebracteate or
rarely with one or two inconspicuous, subulate bracts above; heads commonly
NASH: FLORA OF GUATEMALA 433
nodding when young and again when in fruit, erect only at anthesis, 2-2.5 cm. high;
phyllaries linear or lance-linear, acuminate, floccose-tomentose; ligulate corollas red-
purple or white tinged with red or purple; achenes about 5 mm. long, pubescent or
almost glabrous, 5-6-costate, the filiform beak 2-3 times longer than the body; pappus
white or yellowish, 1-1.5 cm. long.
JUNGIA Linnaeus, f.
Herbs or shrubs, erect or scandent, variously pubescent; leaves alternate,
petiolate, the blades mostly orbicular-cordate, angulate and dentate, or lobate, often
tomentose beneath; inflorescences paniculate at the ends of the branches; heads
homogamous; involucres cylindrical or campanulate; phyllaries 1-2-seriate, subequal
or a few of the outer ones shorter; receptacle paleaceous; pales rather rigid,
embracing all or most of the flowers; flowers all hermaphrodite and fertile, the
corollas white, yellow, or pink, bilabiate, the outer lip tridentate, the inner one bifid;
anthers sagittate at the base, the auricles narrow, caudiculate; style branches dilated
above, truncate and penicillate at the apex; achenes glabrous or pubescent, oblong or
linear, subterete, 5-costate, contracted above (in ours), or (in some South American
species) the achenes attenuate into a beak; pappus bristles uniseriate, plumose,
barbellate, or denticulate.
[About 25 species, all American and all except two or three in
South America. One is in Guatemala, one in Costa Rica and
Panama, and one has been described from western Mexico.
Jungia guatemalensis Standl. & Steyerm. Field Mus. Bot. 23:
142. 1944. Figure 138.
Dense, wet, mixed forest, 1,400-1,800 m.; Alta Verapaz (type
from Tactic, Tuerckheim 8410); Quezaltenango.
Small vines, herbaceous or suffrutescent, sometimes merely reclining on the
ground, the stems rather stout, subterete, striate, when young densely pilose or
villosulous with brownish, spreading hairs; leaves on petioles 2-9 cm. long, the blades
membranaceous, suborbicular in outline, shallowly cordate at the base, 3.5-11 cm.
long, 5-14 cm. wide, very shallowly 7-lobate or angulate-lobate, villosulous above with
multicellular hairs, somewhat paler beneath, sordid-tomentose and glandular-
punctate, the lobes broadly rounded or very broadly triangular, entire or undulate;
cymes disposed in open and interrupted, leafy panicles at the ends of the branches;
heads 9-10 mm. high, crowded in small, dense cymes, sessile or short -pedicellate; inner
phyllaries about 8, oblong, 5 mm. long, broadly triangular, obtuse and subapiculate at
the apex, very densely and minutely puberulent, the outermost phyllaries few, about
2 mm. long, elliptic-oblong or lance-ovate, acuminate; immature achenes linear,
glabrous, 1.6 mm. long; pappus bristles 5-6 mm. long, white, soft, short- plumose.
This species has been reported from Guatemala as J. ferruginea
L. f. (occurring in Costa Rica and Colombia). That species is very
similar in appearance but has inner phyllaries about 4 mm. long
and heads only 5-7 mm. high. The South American J. spectabilis D.
434 FIELDIANA: BOTANY, VOLUME 24
Don has narrowly acute to acuminate phyllaries; both J. spectabilis
and the Mexican J. pringlei Greenm. have pubescent achenes.
Perhaps conspecific with J. guatemalensis are sterile collections
from Chiquimula and El Progreso, which differ slightly in
pubescence from the Alta Verapaz collections.
LYCOSERIS Cassini
Erect or somewhat scandent shrubs; leaves alternate, sessile or short-petiolate,
the blades entire or nearly so, often subcoriaceous, glabrous above, often white-
tomentose beneath, generally 3-5-plinerved; plants dioecious; pistillate heads larger
than the staminate ones, solitary or few at the ends of the branches, short-
pedunculate, commonly nutant; involucres campanulate or subglobose; phyllaries
subcoriaceous, multiseriate, appressed, narrow, the apices erect or reflexed; receptacle
flat, bearing short fimbrillae; ray flowers uniseriate, pistillate and fertile in the
pistillate heads with the disc flowers fertile or sterile, and all flowers sterile in the
staminate heads; ray corollas yellow or orange, the ligule spreading, 2-3-dentate at
the apex, with staminodia present in those of the staminate heads; disc corollas
tubular, the limb 5-parted; anthers sagittate at the base, the auricles long-caudate;
styles in the disc flowers of the staminate heads undivided, in the pistillate flowers
filiform, obscurely bifid or bilobate, the short lobes not dilated; achenes oblong or
narrowly cylindrical, 5-costate or multistriate, glabrous; pappus bristles numerous,
multiseriate, smooth or scaberulous.
About 10 species, all in tropical America, with only one in
Guatemala. Two others are recorded from Costa Rica and Panama.
Lycoseris crocata (Bertol.) Blake, Bull. Torrey Bot. Club 53:
218. 1926. Carduus cernuus Bertol. Fl. Guat. 431. 1840 (type from
Volcan de Agua, Sacatepequez, Velasquez s.n.), not C. cernuus
Steud. 1821. Aster crocatus Bertol. op. cit 434 (type from Volcan de
Agua, Sacatepequez, Velasquez s. n.). L. squarrosa Benth. Bot. Voy.
Sulph. 121. 1844. Santo Domingo (Retalhuleu and Zacapa). Figure
139.
Dry thickets, brushy plains, sometimes in pine forest, sea level
to 500 m.; Chiquimula; Guatemala; Retalhuleu; Sacatepequez;
Zacapa. British Honduras (Belize); El Salvador to Panama.
Usually erect or arching shrubs, 1-1.5 m. tall, the branches conspicuously striate,
glabrous or when young floccose-tomentose; leaves short-petiolate, the blades
lanceolate or lance-oblong, mostly 7-12 cm. long, narrowly acuminate, acute or
attenuate to the base, entire or obscurely and remotely serrulate near the apex,
triplinerved, shining and glabrous above, arachnoid-tomentose beneath when young
but soon glabrate; heads solitary on leafy, short, terminal or axillary peduncles, the
staminate ones about 1.8 cm. high, the pistillate 3-3.5 cm. high; phyllaries lanceolate
or oblong-lanceolate, attenuate, arachnoid-tomentose or glabrate; corollas orange or
yellow; achenes dark, glabrous, about 7 mm. long; pappus yellowish white.
NASH: FLORA OF GUATEMALA 435
• >
These shrubs are rather showy, blooming through the driest
months of the summer season. Carduus cernuus was based by
Bertoloni on the pistillate plant, Aster crocatus on the staminate
one.
ONOSERIS De Candolle
Reference: Ramon Ferreyra, Revision del genero Onoseris,
Journ. Am. Arb. 25: 349-395. 1944.
Herbs or shrubs, various in habit; leaves radical or cauline and alternate, the
blades white- to men tose beneath; heads large, usually heterogamous, solitary or
disposed in panicles, the peduncles or pedicels Usually elongated, bracteate; involucres
hemispheric or turbinate; phyllaries multiseriate, imbricate, graduate, very acute or
acuminate; receptacle naked or pilose-fimbrillate; ray flowers (when present)
pistillate, uniseriate (wanting, in ours), their corollas bilabiate, the outer lip
liguliform, spreading, 2-3-dentate at the apex, the inner lobe with 2 slender segments;
disc flowers hermaphrodite, fertile, the corollas tubular, the limb scarcely ampliate, 5-
parted; anthers sagittate at the base, the auricles caudate; style of the hermaphrodite
flowers filiform, slightly thickened at the apex, obtuse, simple or very shortly
bilobate; achenes subterete, 5-costate, glabrous or pubescent; pappus bristles
numerous, biseriate to multiseriate, scaberulous, rather rigid.
About 25 species, in tropical America, with two in Guatemala
and two known from southern Central America.
Pedicels and phyllaries bearing short, gland-tipped hairs O. donnell-smithii.
Pedicels and phyllaries without gland-tipped hairs O. onoseroides.
Onoseris donnell-smithii (Coult.) Ferreyra, Journ. Arn. Arb.
25: 368. 1944. Pereziopsis donnell-smithii Coult. Bot. Gaz. 20: 53, t
6. 1895.
Known in Guatemala only from the type, Santa Rosa, Rio de
los Esclavos, 750 m., Heyde & Lux 4527. El Salvador.
Plants coarse and stout, to 3 m. tall, the stems densely white-tomentose; leaves
large, several clustered at the top of the stem below the inflorescence, the petioles
elongated, winged, the blades mostly 6-25 cm. long, 6.5-30 cm. wide, cordiform,
palmately veined, simple or pinnatisect, the margins unequally sinuate-dentate, the
upper surfaces somewhat lanate at first, becoming glabrate, the lower surfaces
covered with dense, white tomentum; heads numerous, disposed in a very large, open
panicle, the peduncles and pedicels densely lanate and rather inconspicuously
glandular-pubescent; involucres turbinate, 17-30 mm. high; phyllaries 7-8-seriate,
acuminate, more or less glandular-pubescent dorsally, the margins scarious; corollas
about 2 cm. long, the tube inconspicuously pubescent; anthers about 1 cm. long;
achenes 3.5-5 mm. long, 1.2-1.4 mm. broad, pubescent with ascending hairs; pappus
bristles numerous, yellowish, as much as 16 mm. long.
Called "papelillo" in El Salvador.
436 FIELDIANA: BOTANY, VOLUME 24
Onoseris onoseroides (HBK.) Robins. Proc. Amer. Acad. 49:
514. 1913. Isotypus onoseroides HBK. Nov. Gen. & Sp. 4: 12, t 307.
1820. Caloseris rupestris Benth. PI. Hartweg. 88. 1841 (type from
Cerro Chorro, 12 miles from Guatemala, Hartweg 599). O. isotypus
Benth. & Hook. Gen. PL 2: 487. 1873. Figure 140.
Dry thickets, usually on very steep hillsides or cliffs, 600-1,800
m.; Chiquimula; Guatemala; Huehuetenango; El Quiche; Sacatepe-
quez; Santa Rosa; Solola. Southern Mexico; Honduras and El
Salvador; Costa Rica; northwestern South America.
Coarse herbs or shrubs, 1-2.5 m. tall, usually simple, sometimes sparsely
branched, the stems densely white- tcmentose; leaves both basal and cauline, rather
few, or several clustered near the top of the stem, the long petioles narrowly winged,
the blades large, simple or pinnatisect, when pinnatisect the terminal segments much
larger than the others and similar to the simple leaves, broadly triangular-ovate,
about 21 cm. long and wide, acute or acuminate, truncate or cordate at the base and
subhastate, with acute or obtuse basal lobes, the margins coarsely dentate, densely
white-arachnoid-tomentose beneath; heads numerous, 2.5-3 cm. high, long-pedicellate,
disposed in large, open panicles; involucres turbinate; phyllaries numerous, linear-
lanceolate, long-attenuate, dark red or purplish, glabrate or sparsely pubescent;
corollas red, glabrous or nearly so, the limb bilabiate; achenes linear, densely
pubescent; pappus bristles stramineous, about 2 cm. long, deciduous.
Called "papelillo" in Honduras. It is said that before the
introduction of matches, the white wool covering most parts of the
plant was much used as tinder, and it may still be so utilized in
remote regions. The plants are large and showy.
PEREZIA Lagasca
Reference: Beryl S. Vuilleumier, The systematics and evolution
of Perezia sect. Perezia (Compositae), Contr. Gray Herb. 199: 1-163.
1970.
Perennial herbs with more or less scapiform flowering stems, producing no
cauline leaves or if cauline leaves present, these reduced and bractlike; basal leaves
sessile or petiolate, commonly runcinate-pinnatifid, at least near the base, rarely the
leaves linear and entire; heads homogamous, solitary or disposed in cymes; phyllaries
several-seriate; flowers hermaphrodite and fertile, fragrant, the corollas blue, white,
pink, or purple (in ours), bilabiate (3 segments fused to form the tridentate ligule);
anthers caudate, appendaged, usually brightly colored; style branches recurved,
truncate or rounded at the apex; achenes ellipsoidal, inconspicuously costate,
generally pubescent; pappus bristles 1-3-seriate
Perhaps 35 species, all American, with only two in Central
America.
Basal leaves mostly 5-25 cm. long; peduncles and pedicels glabrous P. nudicaulis.
NASH: FLORA OF GUATEMALA 437
Basal leaves mostly 30-40 cm. long; peduncles and pedicels densely glandular-
pubescent P. glandulifera.
Perezia glandulifera D. Nash, Phytologia 31: 362. 1975.
Known only from the type, canyon of Rio Seligua, Huehue-
tenango, 1,000-1,200 m., Williams, Molina, & Williams 41167.
Erect perennials, the scapes slender, terete, striate, densely glandular-pubescent
the uppermost leaves much reduced, bractlike, glabrous, the lower stem leaves few,
also reduced, oblong-ovate to oblanceolate, mostly 1-4 cm. long, auriculate-
amplexicaul, glabrous, the margins serrate; basal leaves mostly 30-40 cm. long and 15-
18 cm. wide, runcinate-pinnatifid, the margins dentate or dentate-serrate, glabrous
above and below except the lower part of the costae below densely clad with long,
white indument; inflorescence laxly paniculate; heads numerous, pedicellate, each
with 10-12 flowers; peduncles and pedicels densely glandular-pubescent, the pedicels
mostly 1-2 cm. long; phyllaries about 4-seriate, ovate- lanceolate to lance-oblong,
obtuse or acute, mucronulate, glandular-puberulent, greenish; corollas white; mature
achenes about 5 mm. long, appearing linear (actually very narrowly ellipsoidal),
puberulent; pappus bristles about 5 mm. long.
Perezia nudicaulis Gray, PL Wright. 1: 127. 1852 (type
collected in Guatemala by Skinner, the locality not indicated).
Valeriana (Zacapa); hierba de arriero (Jalapa); falsa contrayerba
(Guatemala, fide Aguilar). Figure 141.
Damp or dry, open or brushy slopes or in forest, often in pine-
oak forest, 900-2,500 m.; Baja Verapaz; Chimaltenango; Chiquim-
ula; Guatemala; Huehuetenango; Jalapa; Jutiapa; Quezaltenango;
El Quiche; Santa Rosa; Solola; Zacapa. El Salvador; Honudras.
Acaulescent perennials from stout, thick, somewhat woody rhizomes, these
densely felted at the apex and emitting numerous, slender, fleshy roots, the scapes
mostly 10-60 cm. high, slender, simple below, glabrous, bearing a few minute bracts or
sometimes 1-2 rather large leaves near the base or at the base of the branches; basal
leaves petiolate or subsessile, the blades chartaceous or thinner, lance-elliptic in
outline, mostly 5-20 (-25) cm. long, 2.5-10 cm. wide, long-pilose beneath along the
costa, elsewhere glabrous or nearly so, or sometimes scabrous above or more or less
glandular-puberulent, reticulate-veined, the margins cartilaginous, triangular-dentate
with sharp-tipped teeth, usually runcinate-pinnatifid below but sometimes not at all
lobate, the base then cordate to truncate; heads few or numerous, long-pedunculate,
about 10 mm. high, 11-13-flowered; inflorescence lax and almost naked; phyllaries
about 5-seriate, oblong or the inner ones linear-oblong, glabrate, very obtuse or
rounded at the apex, ciliate, scarious-marginate; corollas white or pinkish, glabrous;
achenes about 5 mm. long, appressed-hispidulous; pappus brownish, uniseriate.
Widely distributed plants in Guatemala, but seldom abundant
in a given locality, the plants scattered and inconspicuous.
438 FIELDIANA: BOTANY, VOLUME 24
TRIXIS P. Browne
References: B. L. Robinson & J. M. Greenman, Revision of the
Mexican and Central American species of Trixis, Proc. Amer. Acad.
40: 6-14. 1904. C. Anderson, A monograph of the Mexican and
Central American species of Trixis (Compositae), Mem. N. Y. Bot.
Card. 22(3): 1-68. 1972.
Shrubby perennials, usually erect, but sometimes scrambling or scandent, usually
more or less pubescent; leaves alternate, the blades ovate-oblong to linear, the
margins almost entire, serrate, or denticulate; inflorescences terminal, densely
corymbose-paniculate or rarely thyrsoid or the heads solitary; heads homogamous,
cylindrical; involucres bracteate, the bracts usually 3-5, unequal, linear to ovate,
foliaceous; phyllaries 1-2-seriate, erect, linear-oblong; receptacle flat; flowers
hermaphrodite, fertile, the corollas conspicuously bilabiate, the upper lobe erect,
bifid, the lower one spreading or revolute, tridentate or subentire, yellow, becoming
white in age; anthers sagittate and caudate; achenes cylindrical or subfusiform,
slender, finely puberulent, often 5-costate, dark, short-rostrate or apically constricted,
the apex dilated into an orbicular disc; pappus bristles white or fulvous, borne on the
disc of the achene.
Less than 40 species, all in tropical and warm-temperate
regions of America, ranging from the southwestern United States to
South America. Only three have been found in Central America.
Stems with wings up to 11 mm. wide T. chiapensis.
Stems with wings less than 2 mm. wide or not winged.
Leaves thick, densely tomentose or lanate beneath with matted hairs, usually
conspicuously rugose above T. nelsonii.
Leaves thin, glabrous or sparsely sericeous beneath, not conspicuously rugose
above T. inula.
Trixis chiapensis C. Anderson, Brittonia 23: 347. 1971.
Clearings and mixed forest, 900-1,000 m.; Huehuetenango.
Mexico (Chiapas).
Plants to 3 m. tall, usually ascending or trailing on other vegetation, the stems
with wings up to 11 mm. wide, glandular and often more or less setose, pilose, or
hirsute, at least when young; leaves mostly sessile or subsessile, sometimes petiolate,
the blades 2.6-21.3 cm. long, 1.4-6.8 cm. wide, linear-lanceolate to lanceolate or
elliptical, acuminate, attenuate to the base or decurrent on the petiole, densely
glandular and more or less strigcse or pilose on both surfaces or sometimes villous or
lanate beneath; margins denticulate or entire; inflorescence a dense, corymbose
panicle or a thyrse, the peduncles densely glandular and sometimes setose or pilose;
bracts subtending the involucre 4-5, mostly 6-20 mm. long and 1.5-11 mm. wide,
linear-lanceolate to elliptical or oblanceolate, densely glandular and sometimes
setose, pilose, or villous, the margins entire; true phyllaries about 8, linear, 9-16 mm.
long, 1.1-2.7 mm. wide, strigose on the distal portion of the adaxial surface, densely
glandular on the abaxial surface; corollas glandular, 9-15 mm. long; achenes 7-9.5
NASH: FLORA OF GUATEMALA 439
mm. long, puberulent but rarely glandular and then only sparsely so; pappus 7-11
mm. long, white or tawny.
Trixis inula Crantz. Inst. Rei Herb. 1: 329. 1766. Inula trixis
L. Amoen. Acad. 5: 406. 1759. Perdicium radiale L. Sp. PL ed. 2.
1248. 1763. Trixis frutescens P. Br. ex Spreng., Syst. Veg. 3: 501.
1826. T. glabra D. Don, Trans. Linn. Soc. London 16: 297. 1830. T.
frutescens var. angustifolia A. DC. in DC. Prodr. 7: 69. 1838. T.
radialis (L.) Kuntze, Rev. Gen. 370. 1891. T. deamii Robins. Proc.
Amer. Acad. 45: 411. 1910 (type from Zacapa, Beam 6359). T.
adenolepis Blake, Contr. U. S. Nat. Herb. 22: 654. 1924 (type from
Zacapa, Beam 324). T. chiantlensis Blake, Brittonia 2: 359. 1937
(type from Huehuetenango, Skutch 1957). Hierba de Santo
Domingo (Guatemala). Figure 142.
Often in damp or dry thickets, brushy slopes, rocky hills, sandy
open areas, sometimes in pine-oak or oak forest, often bordering
grassy plains or pastures, near sea level to 2,500 m.; Alta Verapaz;
Baja Verapaz; Chimaltenango; Chiquimula; Escuintla; Guatemala;
Huehuetenango; Jutiapa; Peten; El Progreso; Quezaltenango; El
Quiche; Retalhuleu; Sacatepequez; Santa Rosa; Solola; Zacapa.
Southeastern Texas; Mexico and Honduras to Panama; West
Indies; northern South America.
Erect, much branched shrubs to 3 m. tall, the stems not winged or rarely with
wings less than 2 mm. wide, glandular and pubescent or strigose when young but
glabrous or only sparsely pubescent in age; leaves subsessile or on very short (1-6
mm.) petioles, the blades elliptical or linear-lanceolate to lanceolate or rarely ovate,
mostly 3-10 cm. long, 1-5 cm. wide, acute or acuminate and apiculate, obtuse or
attenuate to the base, essentially glabrous above or sometimes sparsely strigose,
usually glabrous below but sometimes pilose or rarely villous, especially along the
costae, the margins obscurely denticulate or entire; inflorescences corymbose, often
forming leafy panicles, the peduncles usually sparsely glandular and strigose, pilose,
or sericeous; bracts foliaceous, mostly 3-20 mm. long, equalling or shorter than the
phyllaries; true phyllaries mostly 8-14 mm. long, 1-3 mm. wide, linear-oblong,
commonly acute, sometimes subulate, usually finely appressed-pilosulous to
strigillose, sparsely or densely glandular, rarely glabrous; corollas yellow, the outer
ones 10-16 mm. long, more or less glandular, the inner ones similar but smaller;
achenes 4.5-7 (-9) mm. long, densely and minutely glandular and hispidulous; pappus
sordid or yellowish, 7-11 mm. long.
A weedy, variable, widely distributed species.
Trixis nelsonii Greenm. Proc. Amer. Acad. 41: 270. 1905. T.
amphimalaca Standl. & Steyerm. Field Mus. Bot. 23: 147. 1944
(type from Huehuetenango, Standley 82538).
440 FIELDIANA: BOTANY, VOLUME 24
Dry, rocky hills, 2,000-2,250 m., Huehuetenango. Mexico
(Chiapas).
Erect shrubs to 1.5 m. tall, the stems and branches not winged, densely villous to
lanate, sparsely glandular, at least when young; leaves on petioles 2-13 mm. long, the
blades mostly 2-10 cm. long, 1-25 cm. wide, oblong-lanceolate or narrowly elliptic-
oblong, apiculate and short-acuminate or acute, obtuse or attenuate to the base,
tomentose above when young, in age glandular-pilosulous or glabrate, densely villous
to lanate below and sometimes sparsely glandular, the margins entire or denticulate;
inflorescences commonly corymbose, sometimes thyrsoid; heads short-pedicellate,
crowded; bracts foliaceous, mostly 5-12 mm. long; true phyllaries oblong-linear, 8-15
mm. long, acute or subacute, densely pilose and glandular; corollas yellow, fading to
white, more or less glandular; achenes about 7 mm. long, densely hispidulous; pappus
8.5-12 mm. long, yellowish.
TRIBE XI. CICHORIEAE
By DOROTHY L. NASH
Perennial, beinnial, or rarely annual herbs, commonly with milky sap, the stems
simple or branched; leaves basal and/or cauline, often incised, with dentate or
crenate margins, the cauline leaves alternate; inflorescences of solitary heads on
elongated peduncles or scapes or sometimes sessile, or the heads disposed in cymes,
the inflorescences then sometimes thyrsiform or paniculate; heads homogamous, the
flowers fertile, all ligulate, often yellow; phyllaries imbricate in several series or equal
and uniseriate; receptacle naked or paleaceous; anthers sagittate at the base; style
branches slender, papillose, sometimes somewhat flattened; achenes narrow or flat,
sometimes with a beak bearing the pappus; pappus plumose or not, sometimes of thin
awns or rarely absent.
About 50 genera in both the Old and the New World, with
seven in Guatemala.
Corollas bright blue (in ours); pappus of short, obtuse scales Cichorium.
Corollas variously colored but never bright blue (sometimes very pale blue in
Lactuca); pappus of capillary bristles.
Achenes more or less compressed.
Leaf margins with spine- tipped teeth; achenes not rostrate Sonchus.
Leaf margins not with spine-tipped teeth; achenes more or less rostrate.. .Lactuca.
Achenes not compressed.
Pappus bristles plumose Hypochoeris.
Pappus bristles not plumose.
Corollas lavender or pink Pinaropappus.
Corollas yellow, white, orange, or red.
Scapes hollow; achenes rostrate Taraxacum.
Scapes not hollow; achenes not rostrate Hieracium.
CICHORIUM Linneaus Chicory.
Erect, chiefly perennial, branched herbs with stout stems, the sap milky; leaves
basal and cauline, the cauline ones alternate, mostly small and bractlike; heads large,
NASH: FLORA OF GUATEMALA 441
pedunculate or in sessile clusters along the branches; phyllaries biseriate, herbaceous,
the outer ones short and spreading, the inner ones erect and more or less subtending
the outer achenes; receptacle flat, naked or somewhat fimbrillate; heads
homogamous, the corollas all ligulate, mostly blue, the ligules truncate and 5-dentate
at the apex; anthers sagittate at the base; style branches slender, subobtuse; achenes
5-angulate or 5-costate, truncate, not rostrate; pappus of 2-3 series of short, obtuse
scales.
About eight species, all natives of the Old World.
Cichorium intybus L. Sp. PL 813. 1753. Chicoria; achicoria;
escarola.
Native of Europe; often cultivated in other regions for its roots,
extensively naturalized as a weed in many parts of the United
States; in Guatemala sometimes cultivated for its roots, or in
gardens for ornament, but probably not naturalized.
Perennials from long, deep taproots; stems stout, green, slightly hispid, much
branched, usually less than 1 m. high; basal leaves spathulate, to 15 cm. long,
runcinate-pinnatifid, dentate or lobate, narrowed below into long petioles, the upper
leaves much smaller, lanceolate or oblong, lobate or entire, amplexicaul and
auriculate at the base; heads numerous, 2.5-3.5 cm. broad, the corollas bright blue;
inner phyllaries about 8; pappus of short, obtuse scales.
Chicory has been grown in Guatemala for its fleshy roots
which, when dried and pulverized, are employed for flavoring coffee,
or as a substitute for it. This use of the plant is widespread,
especially in France and other European countries, and even in
some parts of the United States. Bunches of flowering branches are
sometimes sold in Guatemalan markets, especially those of
Quezaltenango, presumably for decoration, although they are not
well suited for this purpose, as the flowers close about noon. In
many parts of the United States chicory has become a rather
troublesome but handsome weed.
HIERACIUM Linneaus Hawkweed.
References: K. H. Zahn, Hieracium, Pflanzenreich IV. 280. Abt.
1, 2, pp. 1-1705. 1921-23; John H. Beaman, Phytologia 32: 45. 1975.
Herbaceous perennials with milky sap, rarely glabrous, usually with two kinds of
indument — simple, often glandular hairs, and smaller, branching or stellate hairs—
often appearing floccose or tomentose; caudex short, usually thick, with fleshy-
fibrous roots; leaves all basal or partly cauline and alternate, sessile or petiolate,
rarely pinnatifid, the margins entire or dentate; flowering stems solitary or several,
the heads solitary, or if numerous, disposed in cymes, the inflorescences sometimes
appearing thyrsiform or paniculate; heads homogamous with only ligulate corollas;
442 FIELDIANA: BOTANY, VOLUME 24
involucres often dark-pilose, cylindric-campanulate or broadly open-campanulate;
phyllaries narrow, herbaceous, the outer ones short, the inner ones subequal;
receptacle flat; corollas commonly yellow, rarely white, orange, or red, truncate and
5-dentate at the apex; anthers sagittate at the base, the auricles shortly setaceous-
acuminate; style branches slender; achenes more or less columnar but mostly
narrowed toward the base, sometimes tapering toward the summit, usually 10-
costate; pappus bristles numerous, 1-2-seriate, simple, rather stiff, persistent, usually
fulvous but sometimes white,
A vast and complex genus in which Zahn recognized more than
750 species, while Bentham and Hooker (1886) guessed the number
of valid species to be scarcely 150, widely distributed in temperate
and cold regions of both hemispheres. There are probably less than
20 in Mexico and Central America, with 10 in Guatemala.
Cauline leaves absent or much reduced and bractlike.
Corollas white H. clivorum.
Corollas yellow.
Involucres mostly 7-9 mm. long H. mexicanum.
Involucres mostly 10-13 mm. long.
Stems glabrate or only sparsely villous below H. skutchii.
Stems more or less long-pilose below.
Leaves linear, linear-lanceolate, or linear-oblanceolate, less than 1 cm. wide;
achenes 2-3 mm. long H. guatemalense.
Leaves oblong-obovate to oblong-spathulate, 2-4.5 cm. wide; achenes 3-4
mm. long H. fendleri ssp. ostreophyllum.
Cauline leaves (at least the lower ones) commonly well-developed, the upper ones,
even when conspicuously reduced, not bractlike.
Heads relatively large, the involucres 10-12 mm. high, commonly long- hirsute.
H. schultzu.
Heads relatively small, the involucres 6-10 mm. high, variously pubescent and more
or less glandular but not commonly long-hirsute.
Inflorescences mostly thyrsiform to narrowly paniculate; achenes tapering at the
base H. abscissum.
Inflorescences laxly cymose to open-paniculate; achenes columnar or tapering
above the middle.
Basal leaves relatively narrow, 1-2 cm. wide, acute or subobtuse and
mucronulate at the apex, the margins remotely but not conspicuously
dentate H. irazuense.
Basal leaves relatively broad, 1.5-4 cm. wide, obtuse or rounded at the apex,
the margins subentire or inconspicuously dentate.
Achenes tapering above the middle; pappus fulvous H. gronovii.
Achenes columnar; pappus white H. stuposum.
Hieracium abscissum Less, in Schlecht. & Cham. Linnaea 5:
132. 1830. H. strigosum D. Don, Trans. Linn. Soc. 16: 175. 1830. H.
thyrsoideum Fries, Symb. Hierac. 141. 1848 (not H. thyrsoideum
Fisch. ex Steud. Nom. ed. 1: 409. 1824, nom. nud.}. H. thyrsoideum
spectabile Fries, loc. cit. H. thyrsoideum plebejum Fries, op. cit. 142.
NASH: FLORA OF GUATEMALA 443
H. anthurum Fries, op. cit. 153. H. comatum Fries, op. cit. 148. H.
comatum var. irregularis Fries, loc. cit. H. multicaule Schaffner ex
Sch. Bip. Bonplandia 9: 173, 326. 1861. Pilosella abscissa (Less.) Sz.
Sz. Flora 45: 436. 1862. P. strigosa (D. Don) Sz. Sz. loc. cit P.
thyrsoidea (Fries) Sz. Sz. torn. cit. 437. H. orizabaeum Arv.-Touv.
Spicileg. Hierac. 17. 1881. H. hirsutum Sesse & Moc. Fl. Mex. ed. 2:
178. 1894. H. abscissum f. hypoglossum Zahn, Pflanzenr. 4 (280):
1103. 1922. H. comatum var. felipense Zahn, torn. cit. 1105. H.
abscissum subsp. morelosanum Blake, Contr. U. S. Nat. Herb. 22:
658. 1924. Figure 143.
Usually in pine or pine-oak forest, sometimes on open, rocky
banks, 1,350-3,350 m.; Alta Verapaz; Baja Verapaz; Chimaltenango;
Huehuetenango; El Progreso; Quezaltenango; Sacatepequez; San
Marcos; Santa Rosa; Zacapa. Mexico; Honduras; Nicaragua; Costa
Rica; Panama.
Erect perennials, the caudex usually short and thick, with numerous fleshy-
fibrous roots, the stems solitary or few, mostly 30-90 cm. tall, more or less long-pilose
with spreading hairs 2-5 (-8) mm. long, commonly densely lanate at the base, usually
somewhat tomentulose above and more or less glandular; basal leaves and those of
the lower stem oblanceolate, mostly 4-25 cm. long, 1-3 cm. wide, narrowing to a
winged petiole, acute, the margins inconspicuously dentate, usually densely long-
pilose, the upper leaves similar or clasping at the base, most of them well developed,
but the uppermost ones often linear and bractlike; inflorescences commonly
thyrsiform or appearing narrowly paniculate; pedicels to 20 mm. long, tomentulose
and/or glandular with dark hairs; heads small, numerous; involucres mostly 6-9 mm.
high; inner phyllaries linear, usually with dark tomentum near the base and more or
less glandular; corollas yellow; achenes dark reddish brown to almost black, tapering
toward the base, 2-3 mm. long, 10-costate; pappus pale fulvous, 4-5 mm. long.
A few atypical Guatemalan specimens have been included here
although they may be hybrids, possibly with H. guatemalense
Standl. & Styerem. or H. irazuense Benth. In all of these the
cauline leaves are fairly well developed as in H. abscissum, but the
inflorescences are more open and with fewer heads, and the
indument of the pedicels and involucres is often a little longer,
darker, and less glandular. These are: Steyermark 48336 and 50578
from Huehuetenango; Steyermark 34154 from Quezaltenango;
Standley 65257 from Sacatepequez; Williams, Molina & Williams
25938 from San Marcos; Steyermark 42513 from Zacapa.
Hieracium clivorum Standl. & Steyerm. Field Mus. Bot. 23:
259. 1947.
Known only from the type, Huehuetenango, Sierra de los
Cuchumatanes, on high bluffs in upper reaches of barranco, above
San Juan Ixcoy, 2,400 m., Steyermark 50065.
444 FIELDIANA: BOTANY, VOLUME 24
Erect perennials from short, thick, horizontal rhizomes, the stems solitary,
slender, monocephalous, 14-27 cm. tall, sparsely long-pilose below, largely glabrate
above to about the middle, then becoming pilosulous upward to densely tomentulose
below the heads with both simple and stellate hairs; basal leaves numerous, 3.5-6.5
cm. long, including the petioles, 0.8-1 cm. wide, the blades oblanceolate, acute,
narrowing to the short or elongating petioles, the leaf margins with 3-4 teeth on each
side, the cauline leaves none or bractlike and linear; peduncles tomentulose and with
some glandular hairs just below the involucre; heads large; involucres about 1 cm.
high, 1-1.5 cm. broad; phyllaries densely glandular; corollas white; achenes juvenile,
glabrous; pappus fulvescent, about 5 mm. long.
Hieracium fendleri Sch.-Bip. Bonplandia 9: 173. 1861. H.
arsenei Blake, Contr. U. S. Nat. Herb. 22: 656. 1924.
Erect, herbaceous perennials, 30-60 (-100) cm. tall, the caudex short, often
producing many thick, fibrous roots, the apex densely long-pilose; basal leaves
present at anthesis and often forming a conspicuous rosette, commonly purple
beneath, oblong-obovate or oblong-spathulate, mostly 4-8 cm. long and 2-4.5 cm.
wide, obtuse or rounded at the apex, sometimes apiculate, narrowed to the subsessile
base, obscurely 4-5-dentate on each side, densely pilose on both surfaces with white
or yellowish hairs 3-7 mm. long, the cauline leaves absent or reduced to 1 or 2 bracts,
these linear to elliptic-oblanceolate or lanceolate, mostly 2-4 cm. long; scapelike
stems long-setose or long-pilose below with hairs 3-5 (-8) mm. long, sparsely short-
pilose and glandular-pilose or glabrate above, more densely glandular-pilose on the
inflorescence; inflorescence lax, corymbiform, the pedicels densely glandular-pilose;
heads usually 4-6 and about 35-flowered; involucres mostly 10-13 mm. high, sparsely
to densely long-pilose; inner phyllaries linear-lanceolate, abruptly acute, long-hirsute,
the longer hairs commonly underlain by floccose hairs, these sparse or dense, the
margins subscarious; corollas yellow or rarely yellowish-orange; achenes dark reddish
brown to almost black, 4-5 mm. long; costate, minutely tuberculate-hispidulous on
the angles; pappus white to yellowish, 5-7 (-9) mm. long.
Only the following subspecies is known in Guatemala.
Hieracium fendleri subsp. ostreophyllum (Standl. & Ste-
yerm.) Beaman, Phytologia 32: 45. 1975. H. ostreophyllum Standl. &
Steyerm. Field Mus. Bot. 23: 104. 1944.
Open, grassy, pine-oak forest, 1,000-2,700 m.; Huehuetenango;
San Marcos (type from northwestern slopes of Volcan de
Tajumulco, Steyermark 36690).
Differs from the typical plants in its more or less pubescent
involucres with gland-tipped hairs, and in the shorter achenes,
mostly 3-4 mm. long.
Hieracium gronovii L. Sp. PL 2: 802. 1753. H. hondurense
Blake, Contr. U. S. Nat. Herb. 24: 31. 1922. H. panamense Blake,
op. cit. 22: 658. 1924. H. minarum Standl. & Steyerm. Field Mus.
NASH: FLORA OF GUATEMALA 445
Bot. 23: 103. 1944 (type from Sierra de las Minas, Zacapa,
Steyermark 29722). Yerba de culebra (Huehuetenango).
In pine forest, 500-2,000 m.; Alta Verapaz; Baja Verapaz;
Chiquimula; Huehuetenango; Solola; Zacapa. Eastern United
States; British Honduras; Honduras; Nicaragua; Panama.
Erect perennials from a short caudex, the stems solitary or few, mostly 30-80 cm.
tall, leafy toward the base, usually densely pilose below, sparsely pilose or glabrate
above or sometimes appressed-pilosulous above and sparsely stipitate-glandular;
basal leaves few, often withering before anthesis, short-petiolate, the blades mostly 3-
10 cm. long, 1.5-3 cm. wide, elliptic-oblong to oblanceolate, obtuse or rounded at the
apex, subentire to inconspicuously dentate, more or less pilose, especially beneath, the
cauline leaves commonly 3-9, rarely only 1 or 2, oblanceolate, obtuse or acute, the
upper ones rounded at the base and sessile to clasping, the lower ones narrowing to a
petiolar base, 3-10 cm. long, 1-3 cm. wide, obscurely dentate, more or less pilose;
inflorescences cymose, sometimes becoming paniculate; heads as few as 3 or 4 or as
many as 45-50; pedicels more or less glandular -puberulent and with very fine stellate
hairs; involucres 7-10 mm. high; inner phyllaries narrowly linear- lanceolate, glabrous
above, usually tomentulose below and more or less glandular; corollas yellow;
achenes dark reddish brown, 3-3.5 mm. long, 8-10-costate, tapering above the middle;
pappus fulvous, 3-6 mm. long.
Hieracivun guatemalense Standl. & Steyerm. Field Mus. Bot.
23: 101. 1944. H. culminicola Standl. & Steyerm. torn, cit.: 100 (type
from Volcan de Santo Tomas, Steyermark 34815).
Open ridges, cliffs, dry rocky slopes of barrancos, grassy slopes,
or in forest, frequently in pine forest, 2,000-3,800 m.; El Progreso;
Quezaltenango (type collected near the summit of Volcan de Zunil,
Steyermark 34860); San Marcos; Zacapa.
Perennials from thick, woody, sometimes branched rootstocks, these densely
fulvous-lanate between the persistent leaf bases and with numerous fleshy-fibrous
roots, the plants acaulescent or nearly so; basal leaves numerous or few, rarely
absent at anthesis, mostly 5-25 cm. long, 0.4-0.8 cm. wide, linear-lanceolate or linear-
oblanceolate, obtuse, callous-apiculate, the margins remotely callous-denticulate and
somewhat runcinate, glabrous or glabrate above, minutely glandular or sparsely
villous beneath, the cauline leaves absent or when present, much reduced, linear, 1-2
cm. long, 1-2 mm. wide; scape monocephalous, 4-15 cm. long, long-villous, some of the
hairs gland-tipped and 1-5 mm. long; heads large, 1.5-2.5 cm. broad; involucre
turbinate-hemispheric; inner phyllaries acute or acuminate, 10-13 mm. long, densely
long-villous; corollas bright yellow; achenes reddish brown, 2-3 mm. long, truncate,
about 10-costate, minutely tuberculate on the angles; pappus bristles pale fulvous or
whitish, 5-6 mm. long.
Hieracium irazuense Benth. in Oerst. Vidensk. Medd. Kjoeb.
1852: 113. 1852. Pilosella irazuensis (Benth.) Sz. Sz. Flora 45: 440.
1862. H. selerianum Zahn, Pflanzenr. IV. 280. 1088. 1922 (type from
446 FIELDIANA: BOTANY, VOLUME 24
Totonicapan or Solola, collected between Totonicapan and Los
Encuentros, C. & E. Seler 2350). H. hypocomum Zahn, torn. cit.
1107. H. maxonii Blake, Contr. U. S. Nat. Herb. 22: 660. 1924. H.
jalapense Standl. & Steyerm. Field Mus. Bot. 23: 102. 1944 (type
from Jalapa, Steyermark 32767). H. tacanense Standl. & Steyerm.
torn. cit. 104 (type from San Marcos, Steyermark 36128). Con-
tragolpe (El Quiche); hierba de culebra (Huehuetenango).
Wet forest and thickets, brushy slopes, rarely in meadows,
1,800-3,800 m.; Chimaltenango; Guatemala; Huehuetenango; Ja-
lapa; El Progreso; Quezaltenango; El Quiche Sacatepequez; San
Marcos; Solola; Totonicapan. Mexico (Chiapas) to Panama.
Perennials from a short caudex with numerous fleshy-fibrous roots, the crown
tufted, the stems 1 -several, to 80 cm. tall, rarely branching, usually densely long-
pilose at the base, more or less short-pilose to glandular-pilose above; basal leaves
commonly short-petiolate, mostly 3-18 cm. long, 1-2 cm. wide (rarely only 0.5 cm.
wide), oblanceolate or elliptic-lanceolate, acute or subobtuse and mucronate, the
margins remotely but conspicuously dentate, pilose on both surfaces, usually long-
lanate with brownish hairs near the base, and the lower surface with at least some
very fine stellate hairs, the cauline leaves usually similar, mostly 5-15 cm. long,
sometimes much reduced and linear, commonly sessile; inflorescences laxly cymose,
the peduncles appressed-pubescent and more or less dark-glandular; pedicels
tomentulose and densely glandular-pubescent; heads usually few; involucres
campanulate, 6-10 mm. long; inner phyllaries narrowly lanceolate, more or less
glandular-pubescent, especially near the base, and commonly sparsely tomentulose,
the margins subscarious; corollas yellow (rarely white); achenes dark reddish brown,
10-costate, 3-3.8 mm. long; pappus fulvous, 4.5-6 mm. long.
Hieracium mexicanum Less, in Schlecht. & Cham. Linnaea 5:
133. 1830. H. niveopappum Fries, Symb. Hist. Hierac. 139. 1848. H.
praemorsiforme Sch. Bip. Bonplandia 9: 327. 1861. Pilosella
niveopappa (Fries) Sz. Sz. Flora 45: 435. 1862. P. praemorsiformis
Sz. Sz. loc. cit. P. mexicana (Less.) Sz. Sz. torn. cit. 436. H.
mexicanum var. niveopappum A. Gray ex S. Wats. Proc. Amer.
Acad. 18: 110. 1883. H. mexicanum subsp. praemorsiforme Zahn,
Pflanzenr. IV. 280. 1105. 1922. H. mexicanum subsp. niveopappum
Zahn, torn. cit. 1106. H. mexicanum subsp. praemorsiforme f.
glabrescens Zahn, loc. cit. H. comaticeps Blake, Contr. U. S. Nat.
Herb. 22: 659. 1924.
Known in Guatemala from a single collection, San Marcos,
Volcan Tajumulco, E. side, ca. 4,100 m., Beaman 3160. Mexico.
Perennials commonly with 1-3 stems from a short, thick caudex, the stems 10-40
cm. tall, more or less pilose and with some stellate pubescence near the base and in
the inflorescence, sometimes with setae and /or glandular hairs in the inflorescence;
basal leaves 3-10 cm. long (including the petioles), 1.5-3 cm. wide, ovate, obovate,
broadly lanceolate, or oblanceolate, mostly acute, narrowing at the base into the
NASH: FLORA OF GUATEMALA 447
short petiole, the margins subentire or inconspicuously dentate, long-pilose to
glabrate above, more or less long-pilose below and minutely tomentulose, the cauline
leaves absent or 1-3, reduced, linear to narrowly lanceolate or oblanceolate;
inflorescences cymose, often appearing thyrsiform or paniculate, the pedicels to 2 cm.
long, tomentose to more or less pilose and sometimes glandular; heads 3-10;
involucres 7-9 (-10) mm. high, 10-12 mm. wide; phyllaries numerous, more or less
long-pilose, glandular and somewhat stellate-pubescent near the base; corollas
yellow; achenes dark reddish brown, tapering at the base, 10-costate, mostly 2-2.5
mm. long; pappus white to somewhat fulvous, 5-6 mm. long.
Hieracium stuposum Fries, Vet. Akad. Forh. 146. 1856. not H.
stupposum Reichb. 1831. Pilosella stuposa (Fries) Sz. Sz. Flora 45:
435. 1862. H. pringlei A. Gray, Proc. Amer. Acad. 19: 69. 1883. H.
jaliscense Robins. & Greenm. op. cit. 40: 23. 1904. H. jaliscense var.
ghiesbreghtii Robins. & Greenm. torn. cit. 24. H. stuppatum Zahn,
Pflanzenr. IV. 280. 1114. 1922. H. jaliscense var. eriobium Zahn, foe.
cit. H. jaliscense subvar. guadalajarense Zahn, loc. cit. H. jaliscense
var. guatemalense Sleumer in Fedde, Repert. Sp. Nov. 41: 118. 1937
(type from Chichicastenango, El Quiche, L. Schultze 947).
In oak, pine, or pine-oak forest, 1,780-2,500 m.; Chimaltenango;
El Quiche; Huehuetenango. Arizona; Mexico.
Erect perennials, the stems solitary or few, mostly to 60 cm. tall, lanate at least
on the lower portion; caudex short and thick with numerous fleshy-fibrous roots;
basal leaves few, commonly short-petiolate but sometimes long-petiolate, the blades
commonly broadly oblong-spathulate or obovate, rarely oblanceolate, mostly 5-15
cm. long, rarely to 30 cm. long, 2-4 cm. wide, usually rounded at the apex, rarely
acute, the margins subentire to repand-dentate, more or less lanate on both surfaces
but more densely so beneath, the cauline leaves 1-5, lanceolate to oblanceolate or the
upper ones linear, narrowing to a sessile base; inflorescences cymose, usually
branching, the peduncles more or less subfloccose and glandular, often densely so;
involucres 7-10 mm. high, narrowly campanulate, glandular, at least near the base;
inner phyllaries linear-lanceolate; corollas yellow; achenes columnar, 10-costate, very
dark reddish brown, 2.5-3.8 mm. long; pappus white, about 5 mm. long.
Hieracium schultzii Fries, Epicris. Hierac. 150. 1862. H. friesii
Sch. Bip. Bonplandia 9: 326. 1861 (not H. friesii Hartm. 1843).
Pilosella friesii (Sch. Bip.) Sz. Sz. Flora 45: 436. 1862. H. bulbisetum
Arv.-Touv. Ann. Cons. Jard. Geneve 3: 27. 1899. H. oaxacanum
Robins. & Greenm. Proc. Amer. Acad. 40: 21. 1904. H. liebmannii
Zahn, Pflanzenr. IV. 280. 1104. 1922. H. wrightii var. palmeri Zahn,
torn. cit. 1108. H. nicolasii Blake, Contr. U. S. Nat. Herb. 22: 657.
1924. H. jaliscopolum Blake, torn. cit. 659.
Grassy slopes and pine forest, 1,800-3,200 m.; Huehuetenango;
Solola. Southwestern United States; Mexico.
448 FIELDIANA: BOTANY, VOLUME 24
Perennials, caudex short, stem usually solitary, sometimes several, mostly 10-70
cm. tall, commonly densely long-pilose below, more or less pilose above, also
inconspicuously stellate- pubescent above and with some yellow, glandular hairs from
black bases, these sometimes quite dense; basal leaves one or several, mostly 3-10 cm.
long (including the petiole when present), 1-3 cm. wide, lance-oblong to la nee- elliptic
or elliptic-oblanceolate, commonly acute, rarely rounded, narrowing into the petiolar
base, the margins subentire or dentate, more or less pilose, the cauline leaves similar,
those near the middle of the stem somewhat reduced and clasping, commonly 3-8
below the middle of the stems, 2-15 cm. long, 0.5-3.5 cm. wide, usually acute or short-
acuminate, rarely obtuse, more or less long-pilose; inflorescences cymose; pedicels
more or less glandular and with stellate tomentum; heads fairly large; involucres 1-
1.2 cm. high, 1.2-1.7 cm. wide; phyllaries numerous, tomentulose below, more or less
glandular above and with some long setae usually present; corollas yellow; achenes
dark reddish brown, 2.3-4.3 mm. long, 10-costate, tapering toward the base; pappus
commonly fulvous, rarely white, to 6 mm. long.
Hieracium skutchii Blake, Brittonia 2: 360. 1937.
On rocky cliffs in coniferous forest, 2,400-3,700 m.; Huehue-
tenango (type from Chancol, Skutch 1258).
Erect perennials from slender rhizomes, the stems solitary, 10-35 cm. tall,
glabrate or sparsely villous below, more densely villous or tomentulose to more or less
setose above; basal leaves few or numerous, thick, mostly 2.5-16 cm. long (including
the petiole), 0.3-1.4 cm. wide, obtuse or acute, narrowing into the short-petiolar base,
the margins essentially entire or inconspicuously dentate, glabrate or sparsely pilose
at least along the costa and/or near the margins, the cauline leaves commonly 3 or 4,
much reduced, linear, sessile; heads 1-3, large; involucres about 12 mm. high, 15-20
mm. wide; phyllaries densely long-pilose with light hairs from a dark base,
eglandular; corollas yellow; achenes dark reddish brown, 10-costate, 2-2.5 mm. long;
pappus fulvous, 4-5 mm. long.
Very similar in general appearance to H. clivorum Standl. &
Steyerm., which differs in its glandular involucres and white
corollas.
HYPOCHOERIS Linneaus
Perennial or rarely annual herbs; leaves mostly basal, rosulate, the margins
entire, dentate, or pinnatifid; stems erect, scapiform or sparsely branched; cauline
leaves usually all bractlike; heads long-pedunculate, homogamous, the flowers all
ligulate; involucres oblong-cylindric or campanulate; phyllaries multiseriate, imbri-
cate, appressed, graduate; receptacle flat or somewhat convex; pales narrow, usually
hyaline; corollas yellow, ligulate, truncate and 5-dentate at the apex; anthers
sagittate at the base, the auricles long-acuminate; style branches slender, subobtuse;
achenes oblong or linear, terete (in ours), 10-costate, little contracted but often
narrowing at the base, the apex of the inner ones terminating in a slender beak, the
outer ones truncate; longer pappus bristles uniseriate, plumose, with numerous short,
simple ones often present.
About 50 species, native in Europe, Asia, and the mountains of
NASH: FLORA OF GUATEMALA 449
South America. Two have become naturalized in the mountains of
Guatemala and Costa Rica.
Leaves glabrous; achenes dimorphic, the outer ones truncate, the inner ones rostrate.
H. glabra.
Leaves hirsute, at least on margins and costae, sometimes with scattered hairs on
both surfaces; achenes uniformly rostrate H. radicata.
Hypochoeris glabra L. Sp. PL 810. 1753. Figure 144.
Roadsides, open fields, sometimes in mixed forest, 1,800-2,800
m.; Guatemala; Quezaltenango; San Marcos; Solola; Totonicapan.
Native of Europe and Asia; naturalized in some parts of the United
States.
Erect perennials, 25-50 cm. tall, the stems solitary or several, glabrous, bearing
several heads; leaves usually spreading on the ground, oblanceolate, 5-12 cm. long,
shallowly lobate or the margins coarsely dentate, glabrous; involucres oblong-
cylindric, 12-15 mm. high; phyllaries glabrous; heads 2-2.5 cm. broad; flowers about
equalling the involucres, the corollas bright yellow, somewhat pubescent below the
base of the limb; achenes 10-costate, the costae scabrous, the outer achenes truncate,
the inner ones long-rostrate.
Hypochoeris radicata L. Sp. PL 810. 1753.
In open field, cloud forest area, 2,300 m., San Marcos. Native of
Europe, naturalized in the eastern United States and in Costa Rica.
Differs from H. glabra primarily in its hirsute leaves and
uniformly rostrate achenes.
LACTUCA Linneaus
Glabrous or hispid herbs with milky sap, usually perennial, sometimes annual;
leaves basal or cauline and alternate, entire, dentate-lobate, or pinnatifid, the
margins often setulose-ciliate, the cauline ones often auriculate-amplexicaul;
inflorescences mostly paniculate; heads sessile or pedicellate, homogamous, the
flowers all with ligulate corollas; involucres cylindrical, usually narrow, sometimes
enlarged below in fruit; phyllaries few-seriate, imbricate, scarious-marginate, the
inner ones elongated and subequal, the outer ones gradually shorter; receptacle flat,
naked; corollas yellow, white, or bluish, the ligules 5-dentate at the apex; anthers
sagittate at the base, the auricles acute or more or less caudate; style branches thin,
short, acute; achenes oval to oblong or linear-fusiform, more or less compressed,
contracted at each end and often rostrate at the apex, the sides 3-5-costate, often
transverse-rugose; pappus bristles copious, multiseriate, persistent or caducous.
About 90 species have been reported, in both hemispheres; a
few are native in temperate North America and Mexico, but only
two reach Central America. A third, the widely cultivated Lactuca
sativa L., is also treated here.
450 FIELDIANA: BOTANY, VOLUME 24
Flowers yellow; cultivated plants L. sativa.
Flowers pale blue, rose-purple, or white; native species.
Inflorescences racemose-paniculate; flowers white; achenes narrow, linear to linear-
oblong, conspicuously muricate, somewhat contracted at the apex but not
conspicuously rostrate L. intybacea.
Inflorescences cymose-paniculate; flowers blue or rose-purple; achenes broad,
flattened, smooth or nearly so, abruptly contracted at the apex into a
conspicuous beak L. graminifolia.
Lactuca graminifolia Michx. Fl. Bor. Amer. 2: 85. 1803. L.
graminifolia var. mexicana McVaugh, Contr. Univ. Mich. Herb. 9:
370. 1972. Figure 145.
Damp or wet thickets, open banks, or open pine forest, 1,300-
1,400 m.; Alta Verapaz. Southeastern United States; Mexico.
Erect perennials, the roots often fleshy, the stems to 1 m. tall, mostly simple
below, naked or bearing a few reduced leaves, terete, glabrous, glaucescent; basal
leaves few or numerous, mostly 5-20 (-30) cm. long, oblanceolate or linear-
oblanceolate in outline, acute, usually pinnatifid with few or numerous shallow,
broadly triangular, obtuse or acute, denticulate or entire lobes, glabrous except for a
few scattered hairs on the lower surface along the costae, most of the leaves
narrowing at the base into long, winged petioles, the cauline leaves smaller, acute or
subauriculate at the base, entire or shallowly lobate or pinnatifid, long-attenuate
(those above mid-stem frequently narrow and entire); inflorescences cymose-
paniculate; heads on slender, elongated, bracteate pedicels; involucres often purplish,
narrow, 12-15 mm. long; inner phyllaries linear-lanceolate, long-attenuate, the outer
ones gradually smaller, lanceolate or lance-ovate; corollas pale blue, rose-purple,
pinkish, or white; achenes dark brown, flattened, 4-5 mm. long, abruptly contracted
into a beak 1-2 mm. long; pappus white, 8-9 mm. long.
Lactuca intybacea Jacq. Icon. PI. Rar. 1: 16. 1786. Phoeni-
xopus intybaceus Less. Syn. Gen. Compos. 137. 1832. Brachy-
ramphus intybaceus DC. Prodr. 7: 177. 1838.
Open banks or dry thickets, sometimes along sandy stream
beds, 400-1,250 m.; Chiquimula; El Quiche. Southern Mexico; West
Indies; northern South America.
Erect, glabrous, annual or perhaps sometimes perennial herbs, usually less than 1
m. tall, the stems solitary or several, terete; basal and lowest cauline leaves obovate
or oblanceolate, acute or obtuse, mostly 8-20 cm. long, runcinate-pinnatifid or
irregularly dentate, setulose-serrulate, the cauline leaves few and small, often
bractlike; inflorescences racemose, branching and forming large, almost naked
panicles; heads short-pedicellate or sometimes subsessile; involucres 10-12 mm. high;
outer phyllaries small, ovate, acute, scarious marginate, the inner ones linear-
lanceolate; corollas white; achenes about 5 mm. long, linear or linear-oblong,
conspicuously muricate, somewhat contracted at the apex but not rostrate; pappus
white, soft, a little longer than the achenes.
Lactuca sativa L. Sp. PI. 795. 1753. Lechuga. Lettuce.
NASH: FLORA OF GUATEMALA 451
Native of the Old World, perhaps of Asia, but probably
unknown in a wild state; grown for food in most inhabited parts of
the earth.
Basal and cauline leaves broad, clasping; flowers yellow.
Lettuce is grown abundantly in Guatemala at almost all
elevations, but probably is little cultivated by the Indians except for
sale. Here, as in North America and Europe, it is the most common
salad plant, and large quantities are on sale at all seasons of the
year. Most of that grown in Guatemala is of the "leaf type but
some of the lettuce of Coban forms good-sized, cabbagelike heads of
the type popular in the United States. Most of the lettuce seed
planted in Central America is imported from the United States.
PINAROPAPPUS Lessing
Glabrous, perennial herbs from thick, fleshy roots, the stems scapiform and
leafless or branched and sparsely leafy; basal leaves narrow or rather broad,
pinnatifid or entire, the cauline leaves when present narrower and mostly entire;
heads usually solitary on long peduncles, the flowers homogamous, the corollas all
ligulate; involucres campanulate; phyllaries multiseriate, membranaceous, dark and
often speckled or spotted at the apex, the inner ones mostly acute, the outer ones
broader and gradually shorter; receptacle flat; pales when present hyaline; flowers
white, pink, or lavender, the ligules truncate and minutely 5-dentate at the apex;
anthers sagittate at the base, the auricles acute or attenuate; style branches slender;
achenes glabrous, linear, terete or subangulate, multicostate, the costae rugulose, the
apex acuminate and rostrate; pappus bristles numerous, simple or minutely
barbellate, the inner ones uniseriate, connate into a basal ring, the outer ones bi- or
triseriate, distinct, shorter.
Several species have been described, their status uncertain; the
group has not been studied critically in recent years, with the
exception of McVaugh's synopsis of the genus in his Compositarum
Mexicanarum Pugillus, Contr. Univ. Mich. Herb. 9: 359-484. 1972.
Only the following species is known in Central America.
Pinaropappus spathulatus Brandg. Zoe 5: 241. 1906. P.
caespitosus Brandg. Univ. Calif. Publ. Bot. 4: 388. 1913. P.
spathulatus var. chiapensis McVaugh, Contr. Univ. Mich. Herb. 9:
377. 1972. Margarita de raonte (Huehuetenango). Figure 146.
Dry, rocky banks, often in the crevices of cliffs, frequently on
limestone and in Quercus or Juniperus forest, 1,350-3,700 m.; El
Quiche; Huehuetenango. Southern Mexico.
Slender, glabrous perennials from thick, perpendicular taproots, or sometimes
with somewhat elongated rootstocks, the stems usually numerous, to 30 cm. long,
452 FIELDIANA: BOTANY, VOLUME 24
commonly scapiform and bearing a single head but sometimes bearing 2 heads, naked
or with a few reduced, subulate or linear bracts; basal leaves numerous, mostly
linear, 3-6 (-15) cm. long, very thin and lax, glaucous, rarely oblanceolate or
spathulate, entire or rarely runcinate-lobate; heads about 1.5 cm. high; involucres 8-
10 mm. high and about as wide, broadly campanulate; phyllaries numerous, graduate,
narrowly oblong, more or less elliptic, or the outer ones nearly ovate, acute or obtuse,
the apex usually ciliolate, the upper half dry, purple or blackish, the inner or middle
phyllaries 1.3-3 mm. wide; corollas lavender to pink; achenes brown, slender, about
4.5 mm. long, the beak shorter than the body; pappus slightly longer than the achene,
brownish.
According to McVaugh, only the var. chiapensis (distinguished
from the typical from chiefly by the slightly wider phyllaries, 1.5-3
mm. instead of 1-1.5 mm.) occurs in Chiapas and Guatemala. He
points out that the type of P. spathulatus (Purpus 1165 from
Veracruz, Mexico) represents a local population and has apparently
been little collected, known only from Veracruz and Pueblo, and
states, "The relationships between populations in this genus, and
the inter- and intra-specific variations, merit experimental study
beyond the scope of this paper."
Our plants have been reported from Guatemala as P. roseus
Less., a common species in southern Texas, Arizona, and Mexico.
Until the whole genus has been carefully studied, it is impossible to
determine which of the described species are valid. The leaves of the
Guatemalan material are commonly narrower than those of
Mexican specimens. What is presumed to be typical P. roseus differs
in having larger, broader heads, usually at least the lowermost
leaves are runcinate-pinnatifid, and the upper dark portion of the
phyllaries is usually bordered by a white, scarious margin.
SONCHUS Linnaeus
Annual or perennial, more or less succulent herbs with milky sap; leaves cauline
and alternate or basal, often auriculate and clasping at the base, entire to pinnatifid
or dissected, mostly with spinescent teeth; inflorescences corymbose-paniculate or
subumbellate; heads homogamous, the flowers all ligulate, usually numerous in each
head (80-250); involucres ovoid or campanulate, rarely nearly cylindrical, in age
thickened or dilated at the base; phyllaries multiseriate, imbricate; receptacle flat,
naked; corollas yellow, the ligules truncate and 5-dentate at the apex; anthers
sagittate at the base with acuminate auricles; style branches slender, almost filiform,
acute; achenes oval to oblong, rugose, more or less compressed, 6-20-costate, not
rostrate (in ours); pappus bristles numerous, multiseriate, capillary, setose, or
plumose, or sometimes reduced to a small crown or scales, or absent, when present
subconnate at the base into an inconspicuous annulus.
Perhaps 70 species, all native to the Old World. Several are
NASH: FLORA OF GUATEMALA 453
widely naturalized in America but only the following is known from
Central America.
Sonchus oleraceus L. Sp. PL 794. 1753. Lechuga (Jalapa);
lechuguilla (Quezaltenango and Sacatepequez). Figure 147.
Damp or wet thickets or fields, occasionally in forest, mostly
along trails, common in damp or rather dry waste or cultivated
ground, 200-3,300 m.; native of the Old World but abundantly
naturalized as a weed in many parts of America, in both tropical
and temperate regions; Alta Verapaz; Chimaltenango; Escuintla;
Guatemala; Huehuetenango; Jalapa; Peten; Quezaltenango; Saca-
tepequez; San Marcos; Santa Rosa; Solola; Totonicapan; Zacapa;
probably in all the departments. United States and Mexico to
Panama; West Indies; South America; Australia.
Erect annuals, 0.3-1 m. tall, glabrous throughout except for some scattered
glandular hairs sometimes on the involucres and the peduncles, the stem stout,
hollow, often dark red or purplish; lower leaves petiolate, to about 15 cm. long, the
blades runcinate-pinnatifid or lyrate, sometimes not lobed, the margins spinulose-
dentate, the upper leaves mostly oblong-lanceolate, clasping at the base and with
large auricles; inflorescences corymbose, often shorter than the leaves; heads few,
mostly 1-2 cm. broad in anthesis; involucres somewhat glandular-hispid or glabrate;
phyllaries 4-seriate, linear- lanceolate, acute, the outermost ones very short and
narrow, the inner ones to about 9 mm. long; corollas yellow, very numerous, the
ligules linear-oblong; achenes light brown, 2-3 mm. long, more or less oblong, flat,
rugulose between the costae; pappus bristles white.
TARAXACUM Zinn Dandelion.
Perennial, scapose herbs with milky sap; leaves basal, usually numerous, the
margins pinnatified or sinuate-dentate; heads large, homogamous, solitary at the ends
of hollow, naked scapes; involucres oblong or campanulate; inner phyllaries
uniseriate, subequal, subconnate at the base, the outer ones multiseriate, shorter,
more or less spreading or at maturity often reflexed; receptacle flat, naked; flowers
all ligulate, the corollas bright yellow, the ligules truncate, 5-dentate at the apex;
anthers sagittate at the base, the thecae often long-acuminate; style branches
slender, subobtuse; achenes oblong or linear-fusiform, 4-5-angulate, 5-10-costate,
rugose or spinulose-muricate, at least upward, attenuate into a very slender beak;
pappus of numerous capillary, unequal, simple, persistent bristles.
Primary species probably 25 or less, but hundreds of segregates,
based on minute or inconsequential characters, have been publish-
ed, from the northern hemisphere. None are native in tropical
America, but one European species has become thoroughly
naturalized throughout most of North America.
Taraxacum officinale Weber in Wiggers, Prim. Fl. Hoist. 56.
454 FIELDIANA: BOTANY, VOLUME 24
1780. Leontodon taraxacum L. Sp. PL 798. 1753. T. dens-leonis
Desf. Fl. Atlant. 2: 228. 1800. Boton de oro (Quezaltenango);
Lechuguilla (Alta Verapaz); diente de Leon. Figure 148.
Damp to rather dry fields, meadows, clay banks, often a weed
in cultivated ground, frequent in streets or about dwellings, 1,300-
3,500 m.; Alta Verapaz; Chimaltenango; Guatemala; Huehue-
tenango; Jalapa; Quezaltenango; Sacatepequez; San Marcos;
Totonicapan. Ubiquitous weeds, native of Europe, widely natural-
ized through much of North America, also common in Central
America, West Indies, and parts of South America.
Perennials from deep, thick roots, the sap very bitter; leaves oblong to
spathulate, mostly 6-25 cm. long, glabrous or more or less pubescent, at least when
young, often very numerous, frequently forming rosettes, the apex acute or obtuse,
the blades narrowing at the base into short, broad petioles, the margins pinnatifid,
sinuate-dentate, or rarely subentire; scapes erect, mostly 3-25 cm. long; heads 2-4 cm.
broad, the flowers very numerous (commonly 150-200), the ligules bright yellow;
inner phyllaries linear or linear-lanceolate, the outer ones similar but much shorter,
reflexed, green, acute; achenes greenish brown or reddish brown, fusiform, spinulose
above, the beak 2-3 times as long as the body; pappus white, abundant.
The common dandelion is thoroughly naturalized in the
highlands of Guatemala, but is seldom the serious pest that it has
become in many parts of the United States. In Europe and in the
United States the tender leaves are often eaten, either cooked or
raw in salads, but we have not seen the plants so utilized in Central
America.
FIG. 1. Elephantopus angustifolius. A, habit of plant with larger leaves indicated
in background, X Vr, B, portion of inflorescence, in fruit, X IVz; C, single head
showing phyllaries, X 3; D, mature achenes with pappus, X 3; E, portion of corolla
dissected, x 9'/2.
455
FIG. 2. Elephantopus mollis. A, habit, X '/2; B, inflorescence, X 2; C, flowering
head with detail, X 3; D, corollas, one dissected to show detail, X 13; E, anther, X
25.
456
FIG. 3. Elephantopus spicatus. A, habit, X V2; B, detail of inflorescence, X 3; C,
head with phyllaries removed, X 2'/2; D, E, and F, achene with pappus, corolla, and
partially dissected corolla, X 5; G, stamen, X 10.
457
FIG. 4. Harleya oxylepsis. A, habit, X Vfc; B, detail of inflorescence, x 2Vfe; C,
phyllary, x 5; D, flowers, in bud and at anthesis, x 5; E, flower partially dissected
to show detail, x 5; F, stamen, x 10.
458
FIG. 5. Lepidonia paleata. A, habit, X 4; B, fruiting heads showing detail, X 3;
C, flower at anthesis, x 11; D, corolla partially dissected, X 11; E, anther, X 20.
459
FIG. 6. Liabum bourgeaui. A, apical part of plant, X M>; B, disc flower with
dissection of staminal column (left), X 10; C, ray flower, X 10; D, cleistogamous (?)
disc flower, X 10; E, mature achene with pappus, X 7; F, achenes, X 2; G, view of
capitulum to show phyllaries, X 2; H, tooth on leaf margin, much enlarged.
460
FlG. 7. Pacourina edulis. A, habit, X Vz; B, flower at anthesis, X 2'/z; C, achene
dissected, and ovule, X 2*4; D, corolla dissected, X 6; E, stamen, X 6.
461
FIG. 8. Piptocarpha chontalensis. A, habit, X Vfe; B, inforescence, X 2; C, detail
of phyllaries and deciduous achene, X 5; D, corolla dissected, with detail of anthers,
achene, and style, X 10; E, achenes with pappus, X 10.
FIG. 9. Spiracantha cornifolia. A, habit, X '/2; B and C, mature heads with
subtending, spinose bract: B (X 4'/2), C (X 2'/2); D, inflorescence at anthesis, X T>i;
E, bract, X VVfe; F, inflorescence, early anthesis, X 7Vy, G, inflorescence partially
dissected, X 7Mz; H, detail of flower, X 15.
463
FIG. 10. Struchium sparganophorum. A, habit, X !/2; B, detail showing axillary
inflorescences, natural size; C, mature head showing achenes and receptacle, X 5; D,
mature head with fallen corollas showing top achenes, X 3Vr, E, flower at anthesis,
X 16; F, corolla showing position of lobes and stamens, X 18, stigma X 25.
464
FIG. 11. Vernonia standleyi. A, habit, X Vz; B, segment or inflorescence, X V/z;
C, flower head to show phyllaries, X 3Vi; D, flower, X 6'/2; E, corolla with style and
stamens in natural position, X 10; F, corolla partially dissected, X 10; G, anther, X
15.
465
FlG. 12. Adenostemma hirtiflorum. A, habit, X Vz; B, capitulum, X 4; C,
capitulum partially dissected, X 3; D, flower with corolla, X 10; E, corolla dissected,
X 10; F, stamen, X 18.
466
FIG. 13. Ageratum cordatum. A, branch from plant, X Vfc; B, capitulum, X 6,
with flowers at anthesis; C, flower at anthesis, X 22; D, flower with corolla removed,
X 22; E, portion of corolla, dissected, X 22.
467
FIG. 14. Ageratum rugosum. A and B, branch and root of the plant, X Vfc; C,
portion of the inflorescence, X 2'/2; D, flower at anthesis, X 15, showing detail of
petal lobe and of achene surface, the pappus is typical of part of this genus; E,
corolla dissected, X 15, with detail of anther and diagram of anther position.
468
FIG. 14a. Brickellia paniculata. A, habit of plant, X Vz; B, flowering head, X
2'/2; C, phyllary, X 7Vz; D, corolla opened to show stamens and pistil, X 7, and (left)
young ovary and a stamen, both greatly enlarged; E, corolla, X 5; F, flower with
achene and pappus, X 5; G, apical portion of achene with some pappus bristles
removed, greatly enlarged.
469
FlG. 15. Carminatia tenuiflora. A, habit of plant, X Vfe; B, capitulum partially
dissected, X 3; C, flower, X 5V&; D, corolla dissected to show stamens, X 6; E,
stamen, X 20.
470
FIG. 16. Eupatorium collinum. A, habit, x Vr, B, flower heads, one partially
dissected, x 6; C, flower at anthesis, x 9; D, flower with corolla removed showing
stylopodium, X 9; E, corolla dissected, X 10; F, stamen, X 15.
471
FIG. 17. Eupatorium lanicaule. A, habit, x Vz\ B, capitulum, x 4; C, series of
phyllaries from outermost to innermost, X 4; D, flower, X 7, with tip of pappus
bristle much enlarged; E, corolla dissected, X 7!/2j F, style with stylopodium, X 7Vz;
G, anther, X 15.
472
FIG. 18. Eupatorium monticola. A, habit, x l/r, B, portion of inflorescence, X 2;
C, flower, x 9, with much enlarged apex of a pappus bristle; D, corolla with styles,
X 9; E, achene, X 9; F, corolla dissected, X 9; G, anther much enlarged. Drawn
from the type.
473
FlG. 19. Eupatorium muelleri. (Mallinoa corymbosa Coulter). Original plate from
Botanical Gazette, used with permission of University of Chicago Press. 1, habit, X
Vr, 2, head, enlarged; 3, flower, enlarged; 4, two stamens, enlarged; 5, pistil, enlarged.
Exact magnifications may be calculated from the description of the species.
474
FIG. 20. Eupatorium nubivagum. L. Wms. A, habit, X Vr, B, inflorescences, x 1;
C, capitulum, partially dissected, X 3; D, flower, x 12, with greatly enlarged detail;
E, corolla dissected, x 12; F, anther, much enlarged.
475
FIG. 21. Eupatorium sodali (Piqueria standleyi). A, habit, X 2Vr, B, segment of
an inflorescence, x 2Vr, C, head of immature flowers, x 10; D, head of epappose
flowers at anthesis, x 10; E, epappose floret, x 17; F, epappose floret dissected, x
17; G, pappose floret, x 17; H, Paul Carpenter Standley (1884-1963), for whom this
plant was originally named, and renamed, when about 50 years old.
476
FIG. 22. Isocarpha oppositifolia. A, habit and portion of inflorescence, X Vr, B,
head at anthesis, X 3, with a phyllary, X 6; C, achenes, a corolla in natural position,
and a subtending pale, all X 19; D, corolla partially dissected, X 25, with an anther
and a stigma much enlarged.
477
FIG. 23. Macvaughiella standleyi. A, branch of the plant, X '/z; B, detail of a
portion of an inflorescence, X 3; C, flowers preanthesis and at anthesis, X 10, and
with enlarged detail; D, flower dissected, X 20; E, phyllaries, X 10; F, Rogers
McVaugh, American botanist (1909- ).
478
FIG. 24. Mikania vitifolia. A and B, portion of a vine, X '/2; C, capitula, X 3; D,
flower at anthesis, X 8; E and F, flower dissected, X 8, with enlarged detail; G,
stamen X ± 15.
479
FIG. 25. Oxylobus glanduliferus. A, habit, X Vfe; B, capitulum, X 3; C, flower, X
10; D, corolla partially dissected, X 10; E, anther, X 14.
480
FIG. 26. Piqueria trinervis. A, habit of a portion of a large plant, X Vr, B, habit
of a smaller plant, X l/2; C, portion of inflorescence, X 2Vr, D, capitulum with one
phyllary pulled out of way, X 6; E, flower dissected, corolla and achene X 15,
stamen and stigmas much enlarged.
481
FIG. 27. Stevia polycephala. A, Central American botanist Antonio Molina
Rossito (1926- ) with plant in Guatemalan highlands; B, detail of branch of plant,
X '/2; C, capitulum at anthesis and one preanthesis, X 4Vr, D, flower at anthesis, X
6, with detail of corolla surface; E, flower dissected, x 6; F, stamen, much enlarged.
482
FIG. 28. Achaetogeron guatemalense. A, habit of two specimens, X Vr, B,
capitula, natural size; C, capitulum partially denuded, X IVi; D, ray flower and disc
flower, both immature, X 12; E, disc flower dissected, X 13; F, anthers front and
back, x 15.
483
FIG. 29. Archibaccharis asperifolia. A, branch, X Vi; B, portion of inflorescence,
X 2V2; C, flowering head, X 5, with phyllary, X 15; D, hermaphrodite flower ready
to open and one at anthesis, X 20; E, corolla dissected to show stamens and style, X
20; F, detail of anthers, X 25; G, stigma of hermaphrodite flower greatly enlarged;
H, pistillate flower, x 15.
484
FIG. 30. Aster spinosus. A, habit of part of plant, X '/z; B, detail of a branch,
natural size; C, head at anthesis and denuded receptacle, X 3; D, ray flower and
ligulate corolla and style from same, X 7; E, disc flower, X 7, and dissected corolla,
X 10; F, achene with pappus, X 7; G, one anther, X 20.
485
FIG. 31. Astranthium purpurascens. A, habit of 2 specimens, both X Vr, B,
flowering head, X 4; C, ray flower with achene, X 9; D, stigmatic branches, greatly
enlarged; E, hermaphrodite flower, X 9; F, hermaprodite flower opened to show
stamens and style, X 15, the single stamen and the stigma (inset) greatly enlarged.
486
FIG. 32. Baccharis vaccinioides. A, habit, X Vz; B, flowering head, X 4; C,
flower, X 8; D, tip of segment of pappus, greatly enlarged; E, flower with corolla
removed, X 8; F, corolla dissected, X 8; G, anthers, X 12.
487
FIG. 33. Conyza bonariensis. A, habit, X Vz; B, flowering head, X 2'/2; C,
stamen, X 14; D, hermaphrodite corolla opened to show stamens and style, X 9'/2; E,
pistillate flower (left) and hermaphrodite flower (right), X 9'/2.
488
FIG. 34. Egletes liebmannii var. yucatana. A, habit, X Vr, B, stem, leaves, and
head, X 2'/2; C, head dissected, X 5; D, ray flower and disc flower with dissections
from the disc flower, all X 25.
FIG. 35. Erigeron karvinskianus. A, habit, X Vz\ B, flowering head and denuded
disc, X 2'/2 (one phyllary greatly enlarged); C, disc flower (left) and ray flower
(right), X 10 (inset of pistillate stigma greatly enlarged); D, disc flower dissected to
show achene with style (left) and stamens (right), X 15 (inset of stigmatic branches
greatly enlarged); E, one stamen, X 32.
490
FIG. 36. Gymnosperma glutinosa. A, habit, X >/2; B, portion of inflorescence, X
4; C, phyllary, X 10; D, ray and disc flowers with dissections, all X 10.
491
FIG. 37. Haplopappus stoloniferus. A, habit of tall plant, X Vfe; B, habit of
average plant, X Vi; C, flowering head partially dissected, X 1%; D, ray flower and
disc flower, X 6; E, disc corolla, partially dissected, X I'/z; F, stigma and anther,
much enlarged.
492
FIG. 38. Heterotheca graminifolia. A, habit, X V2; B, flowering head, X 2'/2; C,
two views of ray flower, X 7; D, disc flower, X 7; E, disc flower corolla dissected, X
18; F, stigma, X 35; G, anther, X 18.
493
FlG. 39. Lagenophora cuchumatanica. A, habit, X '/2; B, flowering head, X 4; C,
disc flower (left), ray flower (right), both X 11; D, style, greatly enlarged; E, disc
flower, the corolla opened to show stamens and style, X 15; F, detail of anther and
style, greatly enlarged.
494
FIG. 40. Solidago stricta. A, habit, X '/v B, portion of inflorescence, X 2Vfc; C,
ray flower (left) and disc flower (right), X 6 (insets of stigma and detail of pappus,
greatly enlarged); D, disc corolla opened to show stamens and style, X 10 (the single
stamen and upper part of style with stigmatic branches, greatly enlarged).
495
FIG. 41. Achyrocline deflexa. A, habit of plant, X Vz; B, detail of inflorescence
(in bud), X 1'4; C, flowering head in anthesis, X 5; D, hermaphrodite and pistillate
florets, X 10; E, hermaphrodite corolla dissected to show style and stamens, X 30
(with detail of anther and stigmatic branches greatly enlarged); F, upper portion of
pistillate corolla with style, X 30.
496
FIG. 42. Adenocaulon lyratum. A, habit of plant, with leaves from a larger plant,
X Vr, B, detail of flowering head with flowers and maturing achenes, X 4'/2; C,
hermaphrodite corolla, X 12!/2; D, hermaphrodite corolla partially dissected, X 14
(detail of anther and stigma greatly enlarged); E, two achenes, one with persistent
pistillate corolla, X 6'/2; F, corolla showing stigmatic branches, X 7l/2.
497
FIG. 43. Epaltes mexicana. A, habit of plant, X Vfe; B, detail of winged stem,
natural size; C, cross-section of stem, X 4; D, inflorescence, X 2; E, flowering head,
the disc partially denuded, with hermaphrodite and pistillate florets, X 6; F, two
views of hermaphrodite corolla and one pistillate corolla, X 24; G, upper portion of
pistillate corolla, showing stigmatic branches, greatly enlarged; H, upper portion of
hermaphrodite style, greatly enlarged.
498
\w
FIG. 44. Gnaphalium americanum. A, habit of plant, X Vr, B, inflorescence, X
34; C, two hermaphrodite florets, one with corolla opened to show stamens, X 25
(inset of stigmatic branches, greatly enlarged); D, pistillate floret, X 25 (inset of
corolla limb with stigmatic branches, greatly enlarged); E, flowering heads, one in
anthesis (top), the other in fruit, X 3; F, two views of pappus showing bristles united
at base into an annulus, greatly enlarged.
499
'-;(/* irvi ?tr/ '1*7 ^ ^lt-: •'-.'-• Wi rl ***Gan\ «•"' vc"V¥ *^
FIG. 45. Gnaphalium brachyphyllum. A, habit of plants, X '/2; B, detail of
inflorescence, X 2; C, hermaphrodite floret (left), pistillate floret (right), X 18; D,
hermaphrodite corolla dissected to show stamens and pistil, X 22'/2.
500
FIG. 46. Pluchea odorata. A, habit, X Vz; B, flowering head (with bud), X 2; C,
hermaphrodite corolla dissected to show stamens and pistil, X 13'/2 (inset of ovary,
greatly enlarged); D, stamens, X 15; E, hermaphrodite floret and achene (left),
pistillate floret (right), X ll'/2 (insets of stigmatic branches greatly enlarged).
501
FIG. 47. Pseudoconyza viscosa. A, habit, X '/2; B, branching inflorescence, X Vz;
C, detail of inflorescence, X 2; D, pistillate flower (left), hermaphrodite flower
(right), X 3; E, two phyllaries, X 7Vz; F, upper portion of hermaphrodite corolla
opened to show stamens, X 7Vz; G, detail of hermaphrodite stigma, X 7Vz.
502
FIG. 48. Acanthospermum hispidum. A, upper part of leafy, flowering branch, X
Vs; B, disc flower complete with achene and pale, X 25; C, disc corolla dissected to
show style and stamens, X 25; D, stamen, greatly enlarged; E, front view of
capitulum, showing the enlarged inner phyllaries enveloping the ray achenes, with 4
sterile disc flowers in center, X 4; F, rear view of capitulum, X 4; G, one ray achene
with the persistent ray corolla, X 7'/2; H, ray flower, greatly enlarged; I, stigma of
disc flower, greatly enlarged.
503
FIG. 49. Aldama dentata. A, part of flowering stem, X Vz; B, capitulum, X 2; C,
disc flower with corolla opened to show stamens and style, X 10; D, detail of stamen
and style, X 20; E, two disc flowers, one with embracing pale (left), one with pale
opened (right), X 7; F, ray corolla, X 7; G, achene and two pales, X 5.
504
f/i
1 1 1 II <p
FIG. 50. Alepidocline annua. A, habit of plant, X Vz; B, mature head, X 4; C,
head dissected to show receptacle, X 4; D, two disc flowers, one more mature with
pappus fallen, X 19; E, disc corolla dissected to show anthers, X 27; F, anthers, X
27; G, ray corolla, X 19.
505
FIG. 51. Ambrosia cumanensis. A, flowering stem, x Vfc; B, pistillate head with
solitary fruit, x 10; C, immature pistil with involucre of fused phyllaries, x 10; D,
anther, x 16; E, staminate corolla dissected to show stamens and style, x 15; F,
stigma, greatly enlarged; G, staminate corolla, closed, x 15; H, two staminate heads,
x 6.
506
FIG. 52. Aphanactis standleyi. A, habit of plant x Vfc; B, base of plant, showing
roots, x IVfe; C, entire plant viewed from above, X 2Vz; D, capitulum, X 6; E, disc
flower opened to show stamens and style, X 20; F, detail of stamen, greatly enlarged;
G, disc flower (left) and ray flower (right) with achenes and pale, X 12, H, achene, x
17; I, stigmatic branches, greatly enlarged.
507
FIG. 53. Baltimore, recta. A, flowering branch, x Vz; B, root system, x Vz; C,
stem section, natural size; D, cyme of inflorescence, terminal head in anthesis, X 2;
E, disc corolla opened to show stamens and style, x 9; F, anther, X 13; G, pale with
sterile disc achene, x 9; H, disc corolla (left) with sterile pistil removed (right), x 9;
I, ray flower complete with achene, X 6 (inset, left, of cyathiform pappus, greatly
enlarged); J, disc flower and pale, x 6; K, three achenes, X 5.
508
FIG. 54. Bidens squarrosa. A, habit of plant, X Vz; B, inflorescence, natural size;
C, achene, X 5; D, (left to right) pale, ray flower, disc flower, X 6; E, stamen,
greatly enlarged; F, disc flower opened to show stamens and pistil, X 12; G,
stigmatic branches, greatly enlarged.
509
FIG. 55. Borrichia arborescens. A, section of leafy stem (glabrous form), X Vz\ B,
section of stem with leaves and inflorescences (sericeous form), X Vr, C, disc corolla
dissected to show stamens and pistil, X 10; D, disc flower (left) and ray flower (right)
with achenes and pale, X 4; E, basal view of capitulum to show phyllaries, X 2; F,
detail of stamen, greatly enlarged.
510
FIG. 56. Calea skutchii. A, flowering branch, X Vr, B, disc corolla dissected to
show style and stamens, X 12; C, two heads, one in bud, one in anthesis, X 3; D, ray
flower with achene, X T/r, E, disc flower with achene and pale, X 7'/2j F, anther,
greatly enlarged.
511
FlG. 57. Calyptocarpus wendlandii. A, habit, X Vz; B, inflorescence with one
head in anthesis, x 4; C, ray flower with immature achene, x Th\ D, disc flower
with immature achene, X 7Vi; E, pale, X 7V6; F, disc corolla dissected to show
stamens and style, x 15; G, mature achene, X 7.
512
FlG. 58. Chrysanthellum americanum. A, upper part of plant, X Vz; B, detail of
basal leaf, X 5; C, phyllary, much enlarged; D, ray flower with pale, X 18; E, disc
flower with pale, x 18; F, achene and pale, x IVr, G, two achenes, one dissected,
showing seeds within, X 7Vz; H, disc corolla opened to show stamen and style, X 34.
513
FlG. 59. Clibadium arboreum. A, flowering branch, X Vz; B, detail of
inflorescence, X 2; C, opened capitulum, X 5; D, pistillate corolla opened with style
removed, X 10; E, pistillate flower, X 10; F, sterile hermaphrodite flower, X 10; G,
hermaphrodite corolla dissected to show stamens and style, X 10.
514
FIG. 60. Coreopsis mutica var. microcephala. A, flowering branch, x W, B,
capitulum, natural size; C, style of disc flower, x 13; D, disc corolla dissected to
show stamens, X 13; E, ray flower, X 5; F, two achenes, X 3; G, disc flower with
pale, X 5.
515
FIG. 61. Cosmos caudatus. A, part of leafy stem, x M>; B, fruiting heads, x Vz;
C, two achenes, x 3Vz; D, stigmatic branches, greatly enlarged; E, disc flower partly
dissected to show stamens and style, X 8 (inset shows enlarged corolla lobe apex); F,
stamen, greatly enlarged; G, disc flower, X 4J/2; H, ray corolla, X 4Vr, I, pale, X 4Vz.
516
FIG. 62. Cuchumatanea steyermarkii. A, habit, X ¥2; B, habit, X 2l/z\ C,
inflorescence (capitulum with subtending spathulate leaves), X 7'/2; D, flower with
pale X 32'/2; E, corolla dissected to show style and stamens, X 50; F, anther and
stigma, greatly enlarged.
517
FIG. 63. Dahlia coccinea. A, part of leafy stem, X Vz; B, inflorescence, X Vz; C,
capitulum, X l!/2 (enlargement of veins and glands of corolla shown in inset); D, disc
flower with pale, X 4; E, ray flower, X 4; F, disc corolla opened to show stamens
and style, with immature ovary removed; G, stigmatic branches, X 5.
518
FIG. 64. Delilea berterii. A, habit, X Vr, B, inflorescence, X 2; C, two capitula,
one opened to show ray achene and stipe of disc flower, X 3'/2; D, capitulum, showing
disc and ray flowers, X 9; E, disc corolla dissected to show stamens and style, X 20;
F, sterile disc flower, X 15; G, fertile ray flower, X 15; H, detail of stamen, X 30.
519
FlG. 65. Desmanthodium guatemalense. A, flowering branch, X Vi; B,
inflorescence, natural size; C, disc corolla (left) opened to show stamens and style,
sterile disc flower (right), detail of style and one stamen (lower left), X 13; D, detail
of inflorescence, X 2; E, pistillate flowers with achenes, one enclosed in the
subtending phyllary, X 11.
520
FIG. 66. Eclipta alba. A, section of plant showing flowering branches, x Vi; B,
detail of inflorescence, X 2; C, disc flower with immature achene and pale, X 20; D,
ray flower with immature achene, x 20; E, two mature achenes, x 17; F, disc
corolla dissected to show stamens and style, x 30; G, stamen and style, greatly
enlarged.
521
FIG. 67. Eleutheranthera ruderalis. A, habit, x W, B, head with two phyllaries
removed to show hermaphrodite flowers, x 3; C, two views of involucres, x 3; D,
three achenes to show variation in surface ornamentation, x 10; E, flower complete
with immature achene and pale, x 13; F, limb of corolla, x 13; G, corolla dissected
to show stamens and style, x 20; H, detail of anther, greatly enlarged.
522
FIG. 68. Galinsoga urticaefolia. A, habit, X Vfc; B, disc corolla opened to show
stamens and style, X 18, with detail of style branches and an anther, greatly
enlarged; C, capitulum, X 2'/2; D, pale, X 13; E, ray flower, X 13; F, disc flower, X
13; G, mature achenes, X 13.
523
FIG. 69. Garcilassa rivularis. A, flowering stem, X Vi; B, inflorescence, X 2'/z; C,
corolla opened to show stamens and style, X 18; D, flowers with immature achenes
and subtending pales, X 9; E, mature achene, X 9; F, detail of style and anther,
greatly enlarged.
524
FIG. 70. Goldmanella sarmentosa. A, flowering stems, X Vz; B, capitulum in
anthesis, X 3; C, disc corolla dissected to show stamens, X 13'/2j D, achene, X 10; E,
detail of anther and style, greatly enlarged; F, disc flower with pale, X 7; G, ray
flower, X 7.
525
r
FIG. 71. Heliopsis buphthalmoides. A and B, flowering stems, X Vr, C, disc
corolla opened to show stamens and style, X 9; D, anther and style, X 11; E, disc
flower with pale, X 4; F, ray flower, X 4; G, section of ray flower showing
staminodia and style, X 7Vz; H, capitulum with some phyllaries and flowers removed
to show receptacle, X 3A.
526
FIG. 72. Heterosperma pinnatum. A, habit, X '/2; B, mature capitulum, X 4; C,
ray flower with pale, X l2Vr, D, disc flower with pale, X 12V4; E, disc corolla
dissected to show stamens and style, X 25; F, detail of stamens and style, greatly
enlarged; G, ray achenes, three views, X 5; H, disc achenes, X 5.
527
FIG. 73. Hidalogoa ternata. A, habit, X Vr, B, dorsal view of capitulum, X l'/4;
C, stamens, X 10; D, disc corolla opened to show stamens and style, X 4; E, base of
style, X 10; F, disc flower with sterile achene and pale, X 4; G, ray flower with
immature achene, X 4; H, mature ray achene, X 7!/2.
528
FIG. 74. Hymenostephium guatemalense. A, habit, X V4; B, disc corolla dissected
to show stamens, X 10; C, disc flower with pale, X 5; D, ray flower, front view, and
(left) ray corolla, dorsal view, X 5; E, detail of anther and style branches, greatly
enlarged; F, capitulum in anthesis, X 2'/2.
529
FIG. 75. lostephane trilobata. A, habit, X Vz; B, ray flower, X 4Vz; C, disc flower
with pale, X 4Vi; D, phyllary, X 4'/2; E, disc corolla dissected to show stamens (style
removed), X 8; F, detail of anther, greatly enlarged; G, achenes, X 6.
530
FIG. 76. Jaegeria hirta. A, habit, X Vz; B, capitulum in anthesis, X 9; C, disc
corolla opened to show stamens and style, X 27; D, disc flower with achene
separated, X 27; E, stamen, greatly enlarged; F, phyllary, X 15; G, pale, X 15; H,
ray flower, X 15; I, disc flower, X 15.
531
FIG. 77. Lagascea helianthifolia. A, flowering stem, X Vfe; B, stamen, greatly
enlarged; C, flower with achene enclosed in calyxlike involucre, X 3'/2; D, corolla
dissected to show style and stamens, X 3Vz; E, flower in bud, X 4; F, portion of
involucre showing top of achene within, X 5.
532
FIG. 78. Lasianthaea fruticosa. A, flowering branch, X Vr, B, two views of ray
corolla, X 4; C, achenes; left, disc achene with pale, right, two ray achenes, X 6; D,
disc corolla, X SVi; E, disc corolla dissected to show stamens and style, X SVz; F,
detail of anther and of style branches, greatly enlarged; G, capitulum in anthesis, X
2.
533
FlG. 79. Melampodium divarication. A, habit, X Vfe; B, capitulum in early
anthesis, X 2J/2; C, capitulum with some flowers removed, showing ray achene at base
of inner phyllary, X 2Vi; D, anther, greatly enlarged; E, disc corolla dissected to show
stamens and style, X 10; F, detail of stigmatic branches, partly divided but fused at
apex, greatly enlarged; G, disc flower with pale, X 10; H, four views of ray flowers,
three with achenes, X 4'/2.
534
FIG. 80. Melanthera nivea. A, flowering stem, X Vz; B, leaf from lower stem, X
'/2; C, dorsal view and partial side view of capitula prior to anthesis, X 3; D, corolla
dissected to show stamens and style, X 15; E, flower complete with immature achene
and pale, X 15; F, mature achenes, X 15.
535
FIG. 81. Milleria quinqueflora. A, flowering stem, X Vfc; B, part of fruiting stem,
X l'/2; C, fruits (achenes enclosed in persistent involucres), X 2'/2; D, achene, X 2Vfe;
E, capitulum in anthesis, X 4; F, disc flower, X 10; G, ray flower, X 8; H, detail of
stigma and anthers, greatly enlarged; I, disc corolla dissected to show stamens and
style, X 10.
536
FIG. 82. Montanoa guatemalensis. A, flowering branch, X Vr, B, inflorescence (in
fruit), X 1/2; C, capitulum in fruit, X IVz; D, capitulum in anthesis, X 2; E, ray
flower, X 4; F, disc flower within pale, X 4; G, two pales enclosing achenes, X 4'/2;
H, disc flower opened to show stamens and part of style, X 8; I, detail of anther,
greatly enlarged.
537
FIG. 83. Neurolaena macrophylla. A, habit, X Vz; B, portion of inflorescence, X
2; C, capitulum X 3; D, dissected capitulum, X IVz; E, flower with pale, X &/z\ F,
dissection of corolla showing stamens and style, X GVfe; G, anther, X 20.
538
FIG. 84. Notoptera scabridula. A, flowering branch, X Vfc; B, detail of
inflorescence, X 3; C, stigmatic branches of style and anther, greatly enlarged; D,
corolla opened to show stamens and style, X 13; E, flower with pale, X 13.
539
FIG. 85. Oteiza ruacophila. A, flowering branch, x !/2; B, ray flower, x 4'/2; C,
capitulum in anthesis, x 3; D, disc corolla opened to show stamens and style, x 12
(style, left, and anther, right, both greatly enlarged); E, disc flower with pale, x 7V&.
540
FIG. 86. Otopappus glabratus. A, flowering branch, X '/z; B, enlargement of leaf
section showing venation and oblique base; C, detail of inflorescence, X 2; D,
stamen, X 13; E, corolla dissected to show stamens (style removed), X 12'/2j F,
flower with pale, X 8; G, style branches, X 13.
541
FIG. 87. Parthenium hysterophorus. A, habit, X Vr, B, two views of capitulum in
anthesis, X 3'/2; C, detail of inflorescence, X I'/a; D, two views of ray flower with
achene enfolded in fused inner phyllaries, X 9; E, dissection showing attachment of
phyllary to ray achene, X 9; F, disc flower in bud, with pale, X 9; G, disc corolla
opened, showing stamens and style, X 16, with enlargements of stigma (left) and
anther (right); H, seed removed from achene, X 9; I, disc flower with pale (side view
of pale, left), X 9; J, part of sectioned capitulum showing numerous disc flowers, X
13.
542
FIG. 88. Perymenium grande. A, flowering branch, X Vi; B, dorsal view of
capitulum in anthesis, X 2; C, disc flower with immature achene, the corolla
dissected to show stamens (style removed), x 12; D, anther and upper part of style,
both greatly enlarged; E, disc corolla, X 5; F, ray flower, X 5; G, mature disc
achene, X 6'/2.
543
FIG. 89. Philactis liebmannii. A, flowering branch, X '/2; B, enlargement of leaf
section showing strigose indument; C, phyllary, X 5; D, disc flower with pale, X 5Vz;
E, ray flower, X 5'/2; F, basal part of ray corolla sessile on the achene, X 5'/2; G, disc
flower dissected to show stamens, style, and apex of achene, X 13; H, upper part of
style and anther, greatly enlarged; I, disc flower, X 13; J, capitulum in anthesis, X 2.
544
FIG. 90. Podachaenium eminens. A, flowering branch, X Vi; B, venation (lower
leaf surface), X 1/6; C, capitulum in anthesis, X 2; D, disc flower, X TVfe; E, ray
flower, X 7'/2; F, disc corolla dissected to show stamens (style removed), X 17Vfc; G,
disc corolla, X 17'/2; H, style branches, greatly enlarged; I, stamen, X 18; J, ray
corolla opened to show staminodes, X 15; K, ray achene, X 15.
545
FlG. 91. Polymnia maculata. A, flowering branch, X '/2; B, capitulum, showing
ray achenes embraced by inner phyllaries, X IVr, C, dorsal view of capitulum, X I'/a;
D, disc flower, X 4'/2; E, ray flower, X 4'/2; F, disc flower with pale, the corolla
opened to show stamens and style X IVi; G, style and two stamens, X 20.
546
FIG. 92. Rensonia salvadorica. A, flowering branch, X '/2; B, detail of
inflorescence, X 3; C, sterile disc flower and two pales, X 6; D, fertile ray flower, X
6; E, stamen, greatly enlarged; F, disc corolla dissected to show stamens and style, X
1.
547
FlG. 93. Rojasianthe superba. A, flowering branch, X Vz; B, capitulum in fruit,
natural size; C, habit of plant, greatly reduced; D, section of stem, X '/z; E, disc
flower with immature achene, X 3'/z; F, ray flower with immature achene, X 3'/z
(right, pappus bristle, greatly enlarged); G, pale of disc achene, opened, X 3'/2; H,
mature achene, X 4; I, disc flower dissected to show stamens (style removed), X 7'/z;
J, stamen and upper part of style, greatly enlarged.
548
FIG. 94. Rumfordia standleyi. A, flowering branch, X Vz; B, disc corolla dissected
to show stamens and style, X T'i (detail of hairs in inset, greatly enlarged); C,
anther, x 10; D, ray flower with paleiform phyllary (left) and disc flower with
opened pale (right), X 5.
549
FIG. 95. Sabazia pinetorum. A, habit, X >/2; B, capitulum, X 2'/2; C, section of
stem, X 5; D, ray flower with pale and part of phyllary (left) and disc flower with
pale (right), X 6; E, disc achene, X 12; F, disc corolla, X 12; G, disc corolla opened
to show stamens and style, X 12'/2; H, two anthers, greatly enlarged; I, upper part of
style, greatly enlarged.
550
FIG. 96. Salmea scandens. A, flowering stem, X Vr, B, detail of inflorescence, X
l'/2; C, disc corolla dissected to show stamens and style, X 20; D, detail of stigmatic
branches and anther, greatly enlarged; E, flower with pale, X 12V2; F, corolla and
achene, X 18.
551
FIG. 97. Sanvitalia procumbens, A, habit, X Vr, B, disc flower with pale, X 9; C,
upper part of disc achene, X 20; D, apex of disc achene, X 20; E, disc achene opened
and seed removed, X 4'/2; F, detail of inflorescence, X 2; G, stamen and pistil of disc
flower, greatly enlarged; H, disc corolla opened to show stamens and pistil, X 15; I,
style of ray flower, greatly enlarged; J, three views of ray achene with persistent
corolla, X 6.
552
FIG. 98. Schistocarpha platyphylla. A, flowering branch, X Vr, B, capitula, X 3;
C (left), pistillate ray flower, (center and right), two slightly different disc flowers, all
X 9; D, corolla partially dissected to show detail, X 9, with style (right) and stamen
(left) much enlarged.
553
FIG. 99. Sclerocarpus divaricatus. A, habit, X '/z; B, ray corolla, X 3Vr, C, disc
corolla, X 3'/2; D, disc achene, X 3'/2; E, upper portion of disc flower, X 7; F, mature
fruits, one with achene removed from the accrescent pale, X 5Vr, G, anther, greatly
enlarged; H, disc corolla dissected to show stamens and style, X 7Vz.
554
FIG. 100. Selloa obtusata. A, habit, X '/2; B, parts of flowering stems, X l'/2; C,
disc corolla partly opened to show stamens and style, X 20; D, detail of anther and
stigmatic branches of style, greatly enlarged; E, disc flower, X 17'/2; F, disc flower
(left) H, receptacle with persistent pales, much enlarged.
FIG. 101. Sigesbeckia agrestis. A, flowering stem, X Vz; B, detail of inflorescence,
X 7; C, ray flower with pale, X 14; D, disc flower with pale, X 14; E, style branches
of ray flower, greatly enlarged; F, stamen, X 30; G, disc corolla dissected to show
style and stamens; H, style branches of disc flower, greatly enlarged.
556
FIG. 102. Simsia amplexicaulis. A, habit, X Vfe; B, neutral ray flower, X 5; C,
fertile disc flower, with pale, X 5; D, anther, greatly enlarged; E, disc corolla partly
dissected to show stamens and style, X 12'/2; F, three achenes, one (right) immature,
X 5; G, phyllaries, greatly enlarged.
557
FIG. 103. Spilanthes ocymifolia. A, habit, X Vi; B, disc flowers, one with pale, X
10; C, ray flower, X 10; D, disc corolla opened to show stamens and style, X 25; E,
detail of anther and style, greatly enlarged; F, disc achene, X 15; G, two capitula, X
3.
558
FIG. 104. Synedrella nodiflora. A, habit, X Vr, B, inflorescence, X l'/2; C,
capitulum, X 3; D, ray flower, X 10; E, disc flower, X 10; F, disc corolla partly
dissected to show stamens and style, X 18; G, part of style and a stamen, greatly
enlarged; H, phyllary, X 6; I, two disc achenes, X 6; J, ray achene, X 6.
559
FlG. 105. Tithonia longiradiata. A, flowering stem, X Vfe; B, detail of portion of
capitulum, X 3; C, stamen, X 14; D, disc corolla opened to show stamens (style
removed), X 12; E, disc flower with pale, X 5; F, neutral ray flower, X 5.
560
FIG. 106. Tragoceras schiedeanum. A, habit, X Vr, B, capitulum X 4'/2; C, three
views of ray achene with persistent corolla, X 5; D, base of ray corolla, with style,
greatly enlarged; E, sterile disc flower with pale, X 8; F, sterile disc achene, X 16; G,
disc corolla dissected to show style and stamens, X 11; H, style of disc flower, greatly
enlarged; I, stamen, greatly enlarged; J, inner surface of corolla lobe, greatly
enlarged; K, lower leaf surface, X 4.
561
FlG. 107. Trichospora verticillata. A, habit, X '/2; B, detail of leaf, X 2Vfc; C,
inflorescence (two capitula), greatly enlarged; D, disc flower, X HVz; E, two views of
phyllary, X HVz; F, corolla dissected to show stamens (style removed), X 22'/2; G,
anther and upper part of style, greatly enlarged; H, outer tip of corolla lobe showing
viscid exudation, greatly enlarged.
562
FIG. 108. Tridax procumbens. A, two flowering stems, x Vr, B, capitulum, x 2;
C, disc flower with pale, X 7; D, pappus bristles, greatly enlarged; E, ray flower
(left) and ray corolla, dorsal view (right), x 7; F, disc corolla opened, showing
stamens and style, X 10; G, anther, greatly enlarged; H, upper part of style, greatly
enlarged.
563
FlG. 109. Trigonospermum annuum. A, habit (basal leaf in background), X Vr, B,
fertile ray flower with pale, X 6; C, three views of ray achene, X 6; D, fruiting head,
X 5; E, section of disc X 3; F, disc flower, X 13; G, disc corolla opened to show
stamens and style, X 13; H, upper part of style of disc flower (left) and anther
(right), both greatly enlarged.
564
FIG. 110. Verbesina fraseri. A, portion of flowering branch, X Vr, B, disc corolla
opened to show stamens and style, X 8; C, disc flower, X 6'/2; D, two achenes, X 4.
565
FlG. 111. Viguiera dentata. A, portion of flowering branch, X Vz; B, detail of
stamen and style branches, both greatly enlarged; C, disc achenes, X 9; D, disc
corolla dissected to show style and stamens, X ll'/2; E, disc flower, X 6; F, disc
flower with pale, X 4; G, ray flower with sterile achene, X 4.
566
FIG. 112. Wedelia acapulcensis. A, flowering branches, X '/2j B, capitulum, X
2'/2; C, pale, X 6; D, disc flower, X 6; E, ray flower, X 6; F, anther, greatly enlarged;
G, disc corolla dissected to show stamens and style, X 11; H, style branches, greatly
enlarged; I, three disc achenes, the one on the left immature, X 5Vz.
567
FIG. 113. Wedelia trilobata. A, B, and C, three flowering stems taken from
different specimens, to show variation in leaf outline, all X Vi; D, ray flower, X 7; E,
disc flower and pale, X 7; F, side view of capitulum, X 2; G, apex of disc achene,
greatly enlarged; H, disc corolla dissected to show stamens and style, X 12!/2.
568
FIG. 114. Zexmenia phyllocephala. A, habit, X Vz\ B, disc flower with pale, X 6;
C, two views of ray corolla, X 4Vfc; D, stamen, greatly enlarged; E, disc corolla
opened to show stamens and style, X 12; F, achenes, X 7; G, detail of style branches
and base of style, greatly enlarged.
569
FIG. 115. Zinnia peruviana. A, habit, typical form, X Vz; B, habit, small form, X
>/2; C, two views of ray flower, X 2'/2; D, seed removed from achene, X 2Vz; E,
capitulum, X 2'/2; F, disc corolla opened (stamens removed, right), X 7'/2; G, apex of
corolla lobe, X 15; H, disc flower with pale (seed removed from achene, right), X 3;
I, stamen, greatly enlarged.
570
FlG. 116. Bahia depauperata. A, habit, X 3 with detail of pubescence (inset,
greatly enlarged); B, flower with nearly mature achene, X 15; C, the pappus as seen
from above, X 15; D, detail of corolla, stamens, and style, X 20; E, anther and
stigma, much enlarged.
571
FIG. 117. Dyssodia montana. A, upper part of a plant, X Vz; B, at left a disc
flower and at right a ray flower, both X 4Vz; C, corollas, at left in natural position
and at right dissected, both X 6'/2; D, anther much enlarged.
572
FIG. 118. Espejoa mexicana. A, plant, X '/•>; B, capitulum at anthesis, X 2'/2; C,
flower past anthesis with nearly mature acene, X 5; D, corolla, X 6; E, dissected
corolla, X 10, with enlarged anther and style.
573
FIG. 119. Florestina latifolia. A, habit, X >/2; B, capitulum, X l'/2; C, flower, X
5; D, pappus, X 10 and apex of achene enlarged; E, corolla in natural position and
one dissected, X 10; F, anther, X ± 22; G, from a collection of the author (16920).
574
FIG. 120. Galeana pratensis. A, habit, X '2; B, portion of inflorescence, X 1; C,
capitulum dissected to show sterile disc flowers and fertile (pistillate) ray flowers, X
7; D, sterile disc flower and two views of fertile (pistillate) ray flowers, X 10; E, two
views of corollas from pistillate ray flowers, X 20: F, corolla from fertile
(hermaphrodite) disc flower and G, dissected corolla with style, both X 25; H, anther
enlarged.
575
FlG. 121. Helenium integrifolium. A, habit of plant past anthesis, X !/z; B, heads
at anthesis from photograph, X ± \'y, C, longitudinal-section through a head, X ''2;
D, disc and ray flower, X 4; E, disc flower, X 8; F, corolla dissected, showing
stylopodium anthers and style, X ± 13; G, anther much enlarged; H, the plant is
said to be poisonous to sheep.
576
FIG. 122. Pectis elongata. A, habit, X Vz; B, portion of inflorescence, X 2Vz; C,
ray flower and disc flower, X 14, with enlarged detail; D, disc corolla with stigma, X
15 with much enlarged stigma and anther; E, disc corolla dissected, X 20.
577
FIG. 123. Porophyllum punctatum. Habit, X 1, from the type of Porophyllum
millspaughii Rob. collected near Progreso, Yucatan, Mexico, Millspaugh 1648.
578
FIG. 124. Schkuhria i-irgata. A, habit of a large plant, X ]/r, B, habit of a small
plant, X '/2; C, capitulum, X 5; D, fertile ray and disc flowers, X 6: E, corollas of
fertile ray and disc flowers, X 12'/2: F, disc flower dissected, X 15; G, anther and
style, much enlarged; H, the pappus and summit of achene, X 74; I, achene, X 7'/2.
579
FlG. 125. Tagetes nelsonii. A, habit of apex of plant, X Vv. B, capitulum, and
involucre showing glands and lobing, X \Vz: C, fertile ligulate flower, X 4, and below
the pappus much enlarged; D, disc flower, X 4; E, disc flower partially dissected, X
7'/2; F, anther much enlarged.
580
FIG. 126. Achillea millefolium. A, habit, X Vz; B, detail of leaves, greatly
enlarged; C, inflorescence, X 2; D, ray flower (left), disc flower (center), and a pale
(right), X 11; E, disc flower opened to show stamens and style, X 22; F, stamen,
greatly enlarged.
581
FIG. 127. Artemisia mexicana. A, basal leaves of young plant, X Vz; B, section of
leafy stem of mature plant, X Vz; C, sections of flowering stems, X Vz; D, detail of
inflorescence, X 2Vz; E, ray flower (left) and disc flower (right), X 12; F, disc corolla
dissected to show stamens and style, X 22; G, one anther, greatly enlarged.
582
FIG. 128. Chrysanthemum parthenium. A, upper half of stem with inflorescence,
X Vz; B, detail of inflorescence and leaf, X l'/2; C, ray flower (left), disc flower
(center), and pale (right), X 10; D, mature achene, X 16; E, disc corolla dissected to
show stamens and style X 25; F, one stamen, greatly enlarged.
583
FIG. 129. Matricaria courrantiana. A, habit, x Vfe; B, detail of leaf, greatly
enlarged; C, capitula, greatly enlarged; D, disc flower and pale (left), ray flower
(center), disc achene (right), X 9; E, disc corolla dissected, showing stamens (style
removed, right), x 22W, F, anther and style branches, greatly enlarged; G, disc
achene, x 25.
584
FIG. 130. Emilia sonchifolia. A, habit, X Vz; B, flower head partially dissected, X
2l/2; C, detail and dissection of flowers, X 7, except anther much enlarged. Erechites
valerianaefolia. D, habit, X Vz; E, flower head, X 2; F, detail and dissection of
flowers, X 4, except stamen and style much enlarged.
585
FIG. 131. Psacalium pinetorum. A, habit, X Vfe; B, capitulum, X 3'/2; C,
capitulum partially dissected, X 3Vr, D, style, stamens, and corolla lobes, X 7; E,
corolla with stamens and style, partially dissected, X 7.
586
FIG. 132. Senecio cobanensis. A, terminal portion of a branch, X Vz; B, portion
of an inflorescence, X ± IVi; C, receptable, X 7; D, disc flower, partially dissected,
X 7; E, ray flower, X 7, with detail of pappus bristle; F, detail of staminal column,
dissected, X 10.
587
FlG. 133. Senecio nubivagus. A, habit, X '/z; B, mature flower, X 8; C, corolla
dissected showing stamens and portion of style, X 8; D, stamens in natural position,
X 25; E, style branches and stigmas, X 25.
588
FIG. 134. Werneria nubigena. A, habit from a photograph, x ± .Vz; B, partial
section through a capitulum to show receptacle, X ± Vr, C, partial dissection of
corolla showing style branches and stamens of a disc floret; D, swollen apex of
filament, much enlarged; E, disc florets, X 2V6; F, ray floret, X 2Vr, G, style branches
from a ray floret, much enlarged; H, barbellate pappus bristle, much enlarged.
589
FlG. 135. Cirsium mexicanum. A, flowering stem, X Vz; B, two detached capitula,
one in bud (left), one in late anthesis (right), X Vz; C, corolla opened to show stamens
and style, X 10; D, stamen, X 13; E, flower complete with achene and pappus, X 5;
F, leaf and section of stem, X Vz.
590
FIG. 136. Acourtia carpholepis. A, habit, X Vfe; B, lower stem node with tuft of
wooly tomentum, X 2; C, flower, X SVz; D, corolla dissected to show stamens (style
removed), X 5Vz; E, style branches, greatly enlarged; F, stamen, greatly enlarged; G,
capitula, X IVfe.
591
\
\
FIG. 137. Chaptalia nutans. A, two plants, both x Vfc, large and small forms; B,
disc corolla dissected to show stamens and style, x 15; C, anther, greatly enlarged;
D, disc flower, X 7!/2; E, ray flower, X 7'/2; F, mature achene, X 15.
592
FIG. 138. Jungia guatemalensis. A, inflorescence, X Vfe; B, part of leafy stem, X
'/2j C, stamen, greatly enlarged; D, corolla opened to show stamens and style, X 8'/2j
E, style branches, greatly enlarged; F, two flowers, one with pale removed, X 7; G,
capitulum X 5.
593
FIG. 139. Lycoseris crocata. A, flowering branch of staminate plant, X Vz; B,
portion of flowering branch of pistillate plant, X Vz; C, two stamens, X 5; D, disc
corolla dissected to show stamens (style removed), X 5; E, disc flower, X 3; F, base
of pappus bristles, greatly enlarged; G, portion of ray corolla from staminate head,
opened to show staminodia, greatly enlarged; H, two views of ray corolla, X 3; I, ray
achene, X 3; J, disc flower of pistillate head, X 3.
594
FIG. 140. Onoseris onoseroides. A, upper part of flowering stem, X Vfe; B, portion
of winged stem, X Vz; C, corolla dissected to show stamens and style, X 4 (detail of
stigma in inset, greatly enlarged); D, stamen, X 5; E, flower, X 4.
595
FIG. 141. Perezia nudicaulis. A, normal habit of plant, X Vz; B, dwarf form, X
Vr, C, immature capitulum X 2Vr, D, corollas dissected to show stamens and part of
style, X 5; E, detail of stigmatic portion of style, greatly enlarged; F, stamen, X 10;
G, flower, X 5.
596
FIG. 142. Trixis inula. A, upper part of flowering stem, x '/z; B, capitulum in
anthesis, complete with bracts, X 2; C, corolla opened to show stamens and style. X
4'/2 (detail of anthers, left, x 4Vz), and tip of corolla lobe in inset, greatly enlarged; D,
stamen, greatly enlarged; E, flower, x 4'/2; F, stigmatic portion of style, greatly
enlarged; G, front view of corolla, X 4'/2.
597
FIG. 143. Hieracium abscissum. A, habit, X Vz; B, detail of inflorescence, X IVz;
C, stamen, greatly enlarged; D, style branches, greatly enlarged; E, flower with
corolla opened to show stamens and style, X 13]/2; F, achene, greatly enlarged.
598
FIG. 144. Hypochoeris glabra. A, habit, X Vi; B, capitulum and denuded
receptacle, X 2; C, achene of outer flower, X 7'/2 (inset showing greatly enlarged
surface ornamentation); D, achene of inner flower, X 7'/2 (inset, above, left, showing
greatly enlarged plumose pappus bristle); E, style branches, greatly enlarged; F,
anther, greatly enlarged; G, two views of ligulate corolla, X 14.
FIG. 145. Lactuca graminifolia. A, habit x Vr, B, detail of basal leaf, x Vfe; C,
capitulum, X 2'/2; D, stamen, greatly enlarged; E, corolla with stamens removed from
style, X 8; F, flower prior to anthesis, with immature achene, X 8; G, style, greatly
enlarged; H, mature achene, X 6.
600
FIG. 146. Pinaropappus spathulatus. A, habit, X Vz; B, outermost phyllary, X 9;
C, achene with pale, X 2Vz; D, corolla with stamens removed from style, X SVfc; E,
flower, X 5'/2; F, stamen, X 12'/2; G, pappus bristle, greatly enlarged.
FlG. 147. Sonchus oleraceus. A, habit, X Vz; B, detail of lower leaf margin,
greatly enlarged; C, achene, X 7!/z; D, flower in early anthesis, X GVi; E, stamen,
greatly enlarged; F, style branches, greatly enlarged.
602
/
FIG. 148. Taraxacum officinale. A, habit, fruiting plant, X Vr, B, habit,
depauperate or dwarf form, flowering plant, X Vt; C, portion of corolla opened to
show stamens, X 16; D, basal portion of anther, greatly enlarged; E, style branches,
greatly enlarged; F, two views of apex of corolla, greatly enlarged; G, phyllary, X
41/2; H, corolla in bud, X 4'/z; I, flower, X 8.
603
Publication 1229