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; hads 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