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FLORA OF GUATEMALA

PAUL C. STANDLEY

LOUIS 0. WILLIAMS

AND

DOROTHY NASH GIBSON

FIELDIANA: BOTANY

VOLUME 24, PART X, NUMBERS 3 and 4

Published by

FIELD MUSEUM OF NATURAL HISTORY

JUNE 17, 1974

FLORA OF GUATEMALA

PART X

FLORA OF GUATEMALA

PAUL C. STANDLEY

Late Curator of the Herbarium

Field Museum of Natural History

LOUIS 0. WILLIAMS

Chairman Emeritus

Field Museum of Natural History

DOROTHY NASH GIBSON

Supervisor, Herbaria

Field Museum of Natural History

FIELDIANA: BOTANY

VOLUME 24, PART X, NUMBERS 3 and 4

Published by

FIELD MUSEUM OF NATURAL HISTORY

JUNE 17, 1974

Library of Congress Catalog Card Number: J^8-3076

PRINTED IN THE UNITED STATES OF AMERICA

BY FIELD MUSEUM PRESS

CONTENTS

Families Included in Part X, Numbers 3 and 4

PAGE

Bignoniaceae 153

Pedaliaceae 232

Martyniaceae 233

Orobanchaceae 238

Gesneriaceae 240

Lentibulariaceae 315

Acanthaceae 328

Plantaginaceae 462

LIST OF ILLUSTRATIONS

TEXT FIGURES

PAGE

21. Adenocalyma fissum 160

22. Amphilophium paniculatum 163

23. Amphitecna macrophylla 165

24. Amphitecna obcvata 168

25. Anemopaegma puberulum 171

26. Arrabidaea sieberi 177

27. Astianthus viminalis 179

28. Callichlamys latifolia 181

29. Clystostoma binatum 184

30. Crescentia alata 186

31. Cydista diversifolia 191

32. Godmania aesculifolia 194

33. Lundia dicheilocalyx 198

34. Macfadyena uncata 200

35. Mussatia hyacinthina 205

36. Parmentiera aculeata 210

37. Pithecolobium echinatum 213

38. Roseodendron donnell-smithii 219

39. Scobinaria japurensis 221

40. Martynia annua 235

41. Proboscidea trilobia 237

42. Conopholis alpina 239

43. Achimenes pedunculata 249

44. Alloplectus calochlamys 252

45. Alloplectus cucullatus 254

46. Besleria laxiflora 258

47. Besleria pansamalana 260

48. Campanea picturata 263

49. Codonanthe macradenia 265

50. Columnea guatemalensis 269

51. Columna sulfurea 271

52. Diastema rupestre 273

53. Drymonia oinchrophylla 276

54. Drymonia psila 278

55. Drymonia serrulata 280

56. Kohleria deppeana 283

57. Kohleria skutchii 289

58. Monopyle puberula 292

59. Napeanthus bracteatus 295

PAGE

60. Niphaea oblonga 296

61. Phinaea repens 298

62. Rechsteineria warscewiczii 300

63. Rhychoglossum azureum 302

64. Sclenophora maculata 306

65. Solenophora obliqua 307

66. Solenophora toucana 311

67. Solenophora wilsonii 312

68. Genlisea filiformis 317

69. Pinguicula lilacina 319

70. Utricularia foliosa 323

71. Aphelandra aurantiaca 334

72. Averia longipes 338

73. Barleria micans 341

74. Blechum brownei 343

75. Bravaisia grandiflora 346

76. Buceragenia glandulosa 348

77. Carlowrightia costaricana 350

78. Chaetothylax rothschuhii 351

79. Dicliptera assurgens 355

80. Dyschoriste ovata 361

81. Elytraria imbricata 365

82. Hdbracanthus ruberrimus 369

83. Hansteinia monolopha 371

84. Hemigraphis alternata 373

85. Henrya imbricans 375

86. Hygrophila costata 378

87. Justicia fulvicoma 392

88. Justicia spicigera 401

89. Lophostachys guatemalensis 405

90. Louteridium donnell-smithii 407

91. Megalostoma viridescens 409

92. Mendocia lindavii 412

93. Neohallia borrerae 414

94. Odontonema callistachyum 417

95. Poikilacanthus macranthus 421

96. P sender anthemum cuspidatum 425

97. Razisea spicata 428

98. Ruellia donnell-smithii 432

99. Sanchezia parvibracteata 441

100. Siphonoglossa sessilis 443

101. Spathacanthus parviflorus 446

102. Stenandrium dulce 448

103. Streblacanthus parviflorus 451

104. Teliostachya alopecuroidea 453

105. Tetramcrium nervosum 455

106. Thunbergia alata 457

107. Trybliocalyx pyramidatus 460

108. Plantago linearis 465

Flora of Guatemala ' Part X, Number 3

BIGNONIACEAE. Bignonia Family

PAUL C. STANDLEY AND Louis 0. WILLIAMS

Reference: Russell J. Seibert, The Bignoniaceae of the Maya area,

(including Yucatan, Campeche, Quintana Roo, Chiapas, Tabasco,

British Honduras, and Guatemala) Carnegie Inst. Wash. Publ. 522:

377-434, it. 5-7. 1940; Use of glands in a taxonomic consideration

of the family Bignoniaceae, Ann. Mo. Bot. Card. 35: 123-136,

tt. 1948.

Trees, shrubs, or woody vines, rarely herbs, the branchlets terete or angulate;

nodes with interpetiolar glandular fields of small impressed plate-like glands or

interpetiolar ridges, sometimes with both, rarely without either; axillary buds with

the outer scales pseudostipular or frequently foliaceous; leaves generally opposite,

without stipules, simple, unifoliolate, or variously digitately or pinnately com-

pound, in most of the scandent plants often trifoliolate, with the terminal leaflet

often replaced by a tendril; leaflets usually entire; inflorescence a terminal or

axillary raceme, thyrse, dichasium, or reduced to a single flower; bracts and

bractlets present, caducous; calyx gamosepalous, usually campanulate or tubular,

5-lobate, in Crescentia bilabiate, spathaceous, or truncate; corolla gamopetalous,

the tube usually dilated above, the limb somewhat bilabiate, the 5 lobes usually

imbricate; fertile stamens 4, rarely 2, didynamous, generally inserted below the

middle of the tube; anthers glabrous or villous, the cells divergent or parallel;

staminode usually very short and inconspicuous; disc pulvinate to cupuliform or

almost patelliform; ovary 2-celled, rarely 1-celled; ovules numerous, attached in

each cell to axial placentae or to 2 parietal placentae when 1-celled; style simple,

the 2 stigmas usually lamelliform; fruit capsular and bivalvate, or baccate and

indehiscent; seeds usually compressed, often broadly winged, or thick and wingless

and imbedded in pulp.

Genera 100 or more, widely dispersed in tropical regions. Ap-

parently only one other genus, Tanaecium, is represented in Central

America and almost all the Central American species of the family

are covered by the following treatment.

Vines of this family constitute a large part of the vegetation of

the thickets on the Pacific plains, as well as in other lowland areas.

There is scarcely a tree or large shrub on these plains that is not

covered with bignoniaceous vines. Because of their great abundance,

153

154 FIELDIANA: BOTANY, VOLUME 24

one would expect ampler material in herbaria but, as a matter of

fact, it is not easy to find plants in good condition for collecting, or

accessible. Fruits can be found with little difficulty, but the flowers

apparently are open for only a very few days each year, and often

they are produced so high on the trees that their collection is not

worth the labor involved, unless there is reason to believe that some

especially rare species is represented.

The Bignoniaceae of Guatemala, along with those of Mexico, the

rest of Central America and Panama, may be divided into four

easily distinguished tribes, as is done in the key that follows. The

Tourrettieae, with a single genus, the Crescentieae with three genera,

the Tecomeae with ten genera, are not complicated, at least not

unduly so. The tribe Bignonieae, on the other hand, with about 28

genera but relatively few species, is difficult and much in need of

a friend.

A look at the characters by which the genera of the tribe Big-

nonieae are said to be separated leaves one with the impression that

often flimsy or even nebulous characters have been used. This

impression is strengthened by two other sources — the number of

genera into which a single species may have been placed by perfectly

competent botanists in the course of time and the large number of

genera that are monotypic. These considerations may indicate that

the lines that separate these genera are all too vague and consequently

that the "solid morphological characters" are capable of being com-

bined in about as many ways as there are characters.

In the preparation of floras it is not possible to take the time to

do monographic work, if one wishes to finish. The work presented

here is certainly not the final word on Guatemalan Bignoniaceae but

we do hope that it is as complete as our specimens would permit.

We have followed Dr. Seibert's account of the Bignoniaceae of

the Maya area, making those modifications and emendations that

seem to be justified by more adequate material and studies by Mr.

Sandwith and Mr. Gentry. Our herbarium now has perhaps more

than twice as many specimens from Guatemala as Dr. Seibert saw

from all herbaria. This increased representation has made it possible

to add to the "Flora" and in some cases to simplify the generic

concepts. The junior author believes that the true number of

genera is fewer than those presented here.

A few days before this portion of the "Flora" was to go to press,

and well more than a year after the last of this manuscript had been

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 155

prepared, Dr. Alwyn Gentry, the friend that the family needs,

gave me some still unpublished notes on Guatemalan Bignoniaceae.

These notes have been incorporated where possible since they reflect

a healthy and much needed look at generic limits. In addition,

Dr. Gentry has prepared the following generic key, based on his

recent studies. I have changed the key in two or three places to

reflect my opinions where they differ from his. — L. Wms., May 1973.

KEY TO GENERA OF GUATEMALAN BIGNONIACEAE

Key contributed by Dr. Alwyn Gentry*

Herbaceous vines; inflorescence subspicate, the upper flowers mostly sterile; ovary

4-locular; capsule densely uncate-spiny, burr like, the valves not splitting to

base (Tribe: TOURRETTIEAE) Tourrettia.

Woody lianas, shrubs, or trees; inflorescence never spicate, without sterile flowers;

ovary 1-2-locular, capsule not spiny with uncate barbs or if spiny not burr-

like and the two valves completely dehiscent.

Trees or shrubs (Podranea, Pandorea, and Tecomaria, all cultivated, are often

vines) ; leaves various but not 2-foliolate and never tendrillate; stems with-

out phloem arms in cross-section; fruits dehiscing perpendicular to the

septum (TECOMEAE) or indehiscent (CRESCENTIEAE).

Leaves pinnately compound (Tribe: TECOMEAE, in part)

Leaves bipinnate; flowers purplish-blue; staminode longer than fertile

stamens Jacaranda.

Leaves once compound; flowers not bluish; staminode much shorter than

fertile stamens.

Calyx spathaceous; exotic trees; corolla large (more than 7 cm. long and

4 cm. wide), broadly campanulate Spathodea.

Calyx not spathaceous; shrubs, vines, or small trees; corolla relatively

small (less than 6 cm. long and 2 cm. wide), tubular-infundibuliform

to tubular-campanulate.

Usually vines; flowers pink to lavender or white; cultivated.

Leaflets entire, acute or obtuse; calyx 6-7 mm. long, shallowly

lobate with broad, obtuse lobes Pandorea.

Leaflets usually serrate; narrowly long-acuminate; calyx 1-2 cm.

long, deeply lobate, the lobes subulate-mucronate . . . .Podranea.

Usually trees or shrubs; flowers yellow to orange-red; cultivated

or native.

Stamens included; flowers yellow; native shrub or small tree. Tecoma.

Stamens exserted; flowers usually orange-red; exotic shrub or

vine Tecomaria.

Leaves simple, 3-foliolate, or palmately compound.

Fruits indehiscent, pulpy with a hard shell or fibrous-fleshy; leaves alternate,

opposite, or fasciculate; corolla thick, usually with a transverse fold

in throat, whitish or greenish, the flowers borne singly or in fascicles.

(Tribe: CRESCENTIEAE).

Calyx spathaceously split; fruits cylindric with a fleshy pericarp; branch-

lets often armed with short spines; leaves 3-foliolate. . .Parmentiera.

Calyx irregularly cleft or bilabiate; fruits globose or ovoid with a hard

shell; branchlets unarmed; leaves simple except in Crescentia alata.

* Minor changes in the key have been made by the junior author to conform

to names which he has followed in this flora. Notably Distictis has been changed

to Pithecoctenium and Dendrosicus to Amphitecna.

156 FIELDIANA: BOTANY, VOLUME 24

Leaves alternate; placentation axile at extreme base of ovary, parietal

on two placentae at top; seeds large (greater than 12 by 14 mm.)

Amphitecna.

Leaves fasciculate; placentation parietal on four placentae; seeds

small (less than 8 by 6 mm.) Crescentia.

Fruit a dehiscent capsule; leaves opposite; corolla thin, more or less tubular

without a transverse fold in throat, color various; inflorescence a ra-

ceme or panicle or reduced to a few flowers (Tribe: TECOMEAE, in part)

Leaves simple; seeds minute (less than 2 by 4 mm); fruit less than

7 cm. long Astianthus.

Leaves palmately compound; seeds larger; fruit more than 11 cm. long.

Flowers less than 1.8 cm. long; anthers pilose; fruits spirally twisted;

leaflets often attenuate Godmania.

Flowers more than 2 cm. long; anthers glabrous; fruits more or less

straight; leaflets conspicuously petiolulate.

Capsule cpstate; calyx thin and membranaceous,

bilabiate Roseodendron.

Capsule not costate; calyx thicker, bilabiate, truncate, or dentate

Tabebuia.

Lianas (some species shrubs when young or open-grown); leaves simple to

3-foliolate, often with tendrils and commonly 2-foliolate in part; stems with

4-16 phloem arms in cross-section; fruits dehiscing parallel to the septum

(Tribe: BIGNONIEAE).

Corolla very small (less than 0.7 cm. long), conspicuously bilabiate with open

mouth; capsule with longitudinal wings Tynanthus.

Corolla usually much larger (always more than 0.8 cm. long), if conspicuously

bilabiate, with closed mouth; capsule not winged.

Corolla orange, the lobes valvate in bud; cultivated exotic Pyrostegia.

Corolla not orange, the lobes imbricate in bud; native plants (though

sometimes also cultivated).

Tendrils uncate-trifid, the branches claw-like; stem irregularly many-

armed in cross-section; corolla yellow; calyx thinly membranaceous.

Capsule oblong, the valves very thick, splitting in half at maturity;

calyx with a short recurved apiculation Melloa.

Capsule linear, the valves thin, not splitting in two; calyx with an

incurved apiculus or not apiculate Macfadyena.

Tendrils not uncate; flowers lavender, white, or yellow; stem evenly

4-16-armed in cross section.

Branchlets six-sided (sometimes rather inconspicuously so in Pithe-

coctenium) with ribbed angles; tendrils 3-15-fid; capsules elliptic-

oblong; corolla thick, white to magenta or reddish; interpetiolar

glandular fields lacking.

Calyx double; corolla bilabiate, the lobes not reflexed, upper 2 and

lower 3 lobes fused; tendrils 3-fprked at tip; leaves usually with

dendroid trichomes; capsule neither echinate nor reticulated.

Amphilophium.

Calyx simple; corolla lobes separate, reflexed; tendrils 3-15-forked

at tip; trichomes all simple; capsule echinate or reticulately

ridged Pithecoctenium.

Branchlets terete to tetragonal, not 6-ribbed; tendrils simple or trifid;

capsules various, often linear; corolla usually thinner, its color

various; interpetiolar glandular fields present or absent.

Leaves conspicuously punctate; fruit very narrow (less than 7 mm.

wide), almost terete, pencil-like; stem hollow Stizophyllum.

Leaves not punctate; fruit more than 8 mm. wide, usually strongly

compressed; stem not hollow.

Leaves dendroid-pubescent, at least in axils beneath (if restricted

to axils, the main veins drying conspicuously reddish below);

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 157

capsule oblong, the valves woody, compressed, smooth, the

seeds with thin bodies and broad wings; tendrils simple.

Interpetiolar glandular fields present; calyx small (less than

9 mm. by 5 mm.); corolla lavender; seed wings hyaline.

Xylophragma.

Interpetiolar glandular fields absent; calyx large (more than

22 mm. by 22 mm.); corolla yellow; seed wings brown.

Callichlamys.

Leaf pubescence, if any, of simple trichomes or lepidote scales;

capsule and seeds various but without above combination of

characters; tendrils simple or trifid.

Flowers yellow or cream, sometimes with reddish streaks;

capsule valves more or less woody, greater than 3 cm. wide,

not echinate or tuberculate; pseudostipules not vertically

3-seriate.

Twigs sharply tetragonal, the angles ribbed; corolla small

(less than 2 cm. long); calyx shallowly campanulate, less

than 2 mm. long Mussatia.

Twigs subterete; corolla greater than 2.5 cm. long; calyx

cupular, greater than 5 mm. long.

Capsule oblong or linear-oblong; ovary stipitate: corolla

glabrous or lepidote without Anemopaegma.

Capsule oblong or linear-oblong; ovary not stipitate;

corolla pubescent without.

Interpetiolar glandular fields present; tendrils trifid.

Ceratophytum.

Interpetiolar glandular fields absent; tendrils simple.

Adenocalymma.

Flowers magenta to lavender or white; capsule valves not

woody unless echinate or tuberculate (except in one variety

of Pachyptera kerere with vertically 3-seriate pseudostipules),

mostly narrower.

Capsule suborbicular, echinate; pseudostipules resembling

miniature bromeliads Clytostoma.

Capsule linear or oblong, not echinate (sometimes tuberculate

or verrucose) ; pseudostipules not a bromeliad-like cluster

of subulate scales.

Tendrils conspicuously trifid; capsule uniformly very flat

or thick and oblong; branchlets terete.

Capsule verrucose-tuberculate; calyx with narrow, elon-

gate (to 5 mm.) twisted teeth; without interpetiolar

glandular fields Adenocalymma.

Capsule smooth; calyx with marginal teeth lacking or

less than 1 mm. long; with or without interpetiolar

glandular fields.

Petiolar and interpetiolar glandular fields absent;

corolla glabrous or very inconspicuously lepidote;

capsule compressed, less than 1.8 cm. wide.

Marline lla.

Petiolar and interpetiolar glandular fields present

(sometimes inconspicious) ; corolla puberulent

without, at least on lobes; capsule thick and ob-

long or compressed but more than 2.5 cm. wide.

Thecae of anthers glabrous Pseudocalymma.

Thecae of anthers pubescent Pachyptera.

Tendrils simple or minutely bifid; capsule linear, compressed

but with edges or midline usually thicker; stems often

angulate.

158 FIELDIANA: BOTANY, VOLUME 24

Anthers pubescent; capsule pubescent with raised midrib.

Lundia.

Anthers glabrous; capsule glabrous or lepidote (pubescent

in Arrabidaea mollissima with median nerve not

raised).

Capsule verrucose; calyx large (more than 18 mm.

long) and evenly bilabiate Scobinaria.

Capsule not verrucose; calyx smaller, subspathaceous

or less than 9 mm. and usually more or less trun-

cate.

Calyx subspathaceous, 12-18 mm. long; capsule

lepidote with margins but not midline some-

what raised; leaves simple or 2-foliolate; stems

terete Neomacfadya.

Calyx more or less truncate or slightly bilabiate,

less than 9 mm. long; capsule as above only in

Cydista aequinoctialis with 2-foliolate leaves

and usually tetragonal stems.

Fertile stamens 2; corolla glandular lepidote;

capsule extremely compressed, glabrous,

without raised midline; some leaves 3-f olio-

late Mussatia.

Fertile stamens 4; corolla pubescent or glandular

lepidote; capsule with raised median nerve or

lepidote or pubescent or the leaves simple or

2-foliolate.

Disc lacking; interpetiolar glands never present;

stems with 8-16 phloem arms in cross-sec-

tion; branchlets often tetragonal; corolla

somewhat lepidote without Cydista.

Disc present; interpetiolar glands often present;

stems with 4 phloem arms in cross-section;

branchlets never tetragonal; corolla puber-

ulous at least on lobes but not lepidote.

Interpetiolar glandular fields almost always

absent; petiolar glandular fields present;

tendril usually minutely bifid; capsule

valves convex, sandpaper-surfaced.

Paragonia.

Interpetiolar glandular fields usually present;

petiolar glandular fields absent; tendril

simple; capsule valves neither convex nor

rough-surfaced Arrabidaea.

ADENOCALYMMA Martius

Woody vines with tendrils, the branches terete, with interpetiolar ridges at

the nodes; leaves 3-2-foliolate, the tendrils simple or rarely trifid; pseudostipules

subulate, not foliaceous; inflorescence a narrow, axillary or terminal raceme or

panicle or thyrse; bracts usually large and conspicuous, caducous; calyx cupular-

campanulate, truncate or denticulate, irregularly lobate or somewhat spathaceously

split, usually with conspicuous plate-shaped glands; corolla funnelform or cam-

panulate-funnelform, densely pubescent outside; stamens included, the anthers

glabrous; disc pulvinate or cupular, large; ovary glandular-lepidote; capsule oblong

or oblong-linear, thick and somewhat flattened, ligneous, smooth, tuberculate or

rugose, septicidally dehiscent; seeds with hyaline wings, the body always very thick.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 159

About 50 species in tropical America, mostly in South America.

Probably only the following are known from Central America.

Corolla yellow; capsule smooth; calyx irregularly and shallowly lobate; leaves

without plate-shaped glands in the axils of the nerves A. calderonii.

Corolla pinkish; capsule tuberculate; calyx 5-costate, with 5 tortuous, subulate

teeth; leaflets with plate-shaped glands in the axils of the 2 basal lateral

nerves A. fissum.

Adenocalymma calderonii (Standl.) Seibert, Carnegie Inst.

Wash. Publ. 522: 428. 1940. Tabebuia calderonii Standl. Journ.

Wash. Acad. Sci. 14: 244. 1924 (type from El Salvador). A. hintonii

Sandw. Kew Bull. 10. 1936.

Dry thickets, 500 m. or less; Peten; Alta Verapaz; Zacapa;

Retalhuleu; Santa Rosa; Escuintla. Southern and central Mexico;

British Honduras; El Salvador; Honduras; Costa Rica.

A large woody vine, drying fuscous, the branchlets with numerous pale

elevated lenticels, almost glabrous; leaves 2-3-foliolate, the leaflets ovate to

elliptic or oblong, 3-13 cm. long, 2-7 cm. broad, obtuse to short-acuminate, obtuse

to shallowly cordate at the base, lustrous, with a distinct cartilaginous margin,

sparsely glandular-lepidote when young, later minutely punctate, otherwise

glabrous; inflorescence of axillary and terminal, narrow racemes or panicles,

somewhat furfuraceous-pubescent; calyx 4-8 mm. long, puberulent-tomentulose,

irregularly lobate or cleft, with a few plate-shaped black glands; corolla yellow,

4-6 cm. long, glabrate near the base, the limb and lobes densely furfuraceous-

pubescent; capsule 10-18 cm. long, 2-3 cm. broad, short-attenuate, rounded at the

base, ligneous, smooth or slightly rugose; seeds transversely oblong, 16-21 mm.

long, 5-8 cm. broad, with membranaceous wings.

Called "bejuco bianco" and "canilla de zopilote" in Tabasco.

The vine apparently is a common one in thickets along the Pacific

coast of Guatemala.

Adenocalymma fissum Loes. Verh. Bot. Ver. Brandenb. 65:

102. 1923. A. seleri Loes. I.e. 101. 1923. Onohualcoa fissa Sandwith,

Kew Bull. 1946: 88. 1947. 0. seleri Lundell, Contr. Univ. Mich.

Herb. 7: 52. 1942.

Moist or dry thickets, 1,200 m. or less; Pete"n; Guatemala.

Southern Mexico.

Branchlets subterete, striate, pale-lenticellate; leaflets 2-3, ovate or ovate-

lanceolate, acute or short-acuminate, obtuse or rounded at the base, 6-14 cm.

long, 2.5-8 cm. broad, coriaceous, not darkening when dried, densely punctate

beneath and sometimes puberulent; axils of the basal lateral nerves with fields of

conspicuous plate-shaped glands; tendrils trifid; inflorescence usually a long,

narrow, terminal or axillary thyrse; calyx 5-10 mm. long, with 5 distinct longitu-

dinal costae, these terminating in subulate teeth 1-2 mm. long, glandular-lepidote

FIG. 21. Adenocalyma fissum. A, pair of leaves about one-half natural size;

B, inflorescence about one-half natural size; G, corolla dissected, X 1; D»

calyx and pistil, X 1.

160

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 161

and puberulent; corolla pinkish lavender, 4-9 cm. long, minutely puberulent and

somewhat lepidote; ovary densely glandular-lepidote; capsule about 19 cm. long

and 2.5-3 cm. in diameter, subcylindric, densely tuberculate-verrucose with

tubercles 2-3 mm. long; seeds 1.5 cm. long and 6 cm. broad.

Maya names are reported from Yucatan as "chacanicab" and

"oppol-che."

This species was segregated from Adenocalymma by Dr. Lundell

based largely upon the verrucose capsule and calyx, as Onohualcoa.

Mr. Sandwith, specialist in the family, seems to agree that the plant

is different and that the species here accounted for may be the same

as the South American Adenocalymma heliocalyx 0. Kuntze or, if

you wish, Onohualcoa heliocalyx Sandwith. The characters used to

separate Onohualcoa seem not very impressive and we prefer to keep

it in Adenocalymma but call attention to the other generic name.

AMPHILOPHIUM Kunth

Woody vines with tendrils; branches hexagonal, the costae distinct, easily

separable, the nodes with interpetiolar ridges; leaves opposite, 2-3-foliolate, the

terminal leaflet sometimes replaced by a trifid tendril; pubescence glandular-

lepidote and often also of stellate or simple hairs; pseud ostipules small, foliaceous;

inflorescence a terminal thyrse (or axillary 2-flowered peduncle) ; calyx campanulate

or subglobose, the orifice somewhat constricted, 2-3-lobate, the limb broad,

undulating and spreading; corolla subcoriaceous, glabrous or finely pubescent, at

least on the lobes, the tube cylindric, the limb bilabiate, with valvate lobes;

stamens included, the anthers glabrous, oblong-oval; disc large, annular and

somewhat pulvinate; ovary ovate, finely pubescent; capsule oblong-elliptic,

somewhat compressed, ligneous, slightly rugose, septicidally dehiscent; seeds

transverse-oblong, with large membranaceous wings.

Perhaps 10 species, in tropical America. Two others have been

described from Central America, one of which, A. pilosum Standl. of

Honduras, might well be expected in the North Coast of Guatemala.

Amphilophium paniculatum (L.) HBK. Nov. Gen. & Sp. 3:

116. 1819. Bignonia paniculata L. Sp. PL 623. 1753. A. purpureum

Brandegee, Zoe 5: 220. 1905. Cucharillo.

Dry or moist thickets, 1,200 m. or less; Pete"n; Alta Verapaz;

Izabal; Chiquimula; Santa Rosa; Escuintla; Jalapa; Sacatepe'quez ;

El Quiche; Suchitepe"quez; Huehuetenango. Southern Mexico;

British Honduras to Panama; West Indies; South America.

A large or small vine, the branchlets glandular-lepidote, the angles short-ciliate,

the nodes with a pubescent interpetiolar ridge; leaves bifoliolate, the leaflets

pellucid-punctate, ovate to broadly ovate or almost rounded, obtusely short-

162 FIELDIANA: BOTANY, VOLUME 24

acuminate, obtuse to cordate at the base, entire, glandular-lepidote on both

surfaces, slightly paler beneath, with a few cup-shaped glands in the axils of the

nerves; pseudostipules foliaceous, 5 mm. long or less, sickle-shaped; thyrses long

and narrow, with 1-2 flowers aborted in many of the cymes; calyx 13 mm. long

or less, densely glandular-lepidote, the interior appendages 3-5 mm. long, obtuse;

corolla 3-4 cm. long, pinkish or purplish, turning white in age, the limb very short,

puberulent to glabrous; capsule 8 cm. long and 3.5 cm. broad, minutely pubescent

or furfuraceous, obtuse at each end.

Known in Tabasco as "pico de pato," in allusion to the fruit,

which suggests a duck's bill; "sac-ac" (Yucatan, Maya). The flowers

are not at all showy nor are they handsome.

Amphilophium paniculatum var. rnolle (Schlecht. & Cham.)

Standl. Field Mus. Bot. 18: 1114. 1938. A. molle Schlecht. & Cham.

Linnaea 5: 120. 1830. Bejuco de cuchamper (probably an erroneous

name); sohbach (Pete"n, Maya).

Dry to wet thickets, 2,500 m. or lower; Peten; Alta Verapaz;

Zacapa; Chiquimula; Jalapa; Jutiapa; Guatemala; Sacatepe"quez ;

El Quiche* ; Huehuetenango; San Marcos. Southern Mexico; British

Honduras to El Salvador and Costa Rica.

Like the typical form of the species except in pubescence; branches, petioles,

and inflorescences with evident stellate pubescence; leaflets glandular-lepidote,

minutely stellate-puberulent above or almost glabrous, densely stellate-pubescent

beneath with short hairs.

This has generally been considered a distinct species, but the

typical form intergrades with typical A. paniculatum, and leaves with

both types of pubescence are found upon the same plant.

Called "pico de pato" in Honduras and El Salvador; "cuchara

de pico," "cuchara de pato," "cucharillo" (El Salvador).

AMPHITECNA Miers

Reference: Louis 0. Williams, Amphitecna and Enallagma,

Fieldiana, Bot. 36 (4) : 21-29. 1973.

Trees or sometimes shrubs, the trunks simple or few-branched, the branchlets

angulate at first, becoming subterete, the bark gray or grayish-white, glabrous,

the nodes often enlarged, pulviniform; inflorescences terminal and 1-few-flowered

or lateral, the lateral sometimes on short-shoots on the branches and 1-few-flowered,

or single flowers in the leaf axils of new wood, the flowers conspicuously pedicellate;

calyx closed in bud, bilaterally or rarely trilaterally cleft at anthesis, rarely (?)

circumscissile at the base; corolla usually white or greenish and somewhat fleshy,

funnelform or campanulate-funnelform, 5-lobate or crenate-lobate; stamens 4 in

two subequal pairs, with a small staminode, all inserted near base of corolla,

FIG. 22. Amphilophium paniculatum. A, habit, X1A\ B, corolla dissected, X

; C, calyx, X 2; D, longitudinal section of flower after Bureau, X %.

163

164 FIELDIANA: BOTANY, VOLUME 24

anthers reaching to throat of corolla, the disc large and persistent on the fruit;

ovary 1-celled or perhaps falsely 2-celled; fruit baccate, ovoid to ellipsoidal, the

pericarp thin (less than 2 mm.) and coriaceous or thick (± 5 mm.) and bony.

The center of distribution of this small genus is Guatemala. It

includes eight species of which six are found in Guatemala. They

are all small trees or shrubs found in the forest, often along streams.

Flowers fasciculate on short bracteplate shoots on old wood; veins on lower

surface of leaves appearing marginate.

Leaves 40-80 cm. long; flowers borne near base of thick stem. .A. macrophylla.

Leaves 6-12 cm. long; flowers borne on defoliated branches A. silvicola.

Flowers in terminal bracteolate fascicles or in (usually singly) the axils of leaves;

veins on lower leaf surfaces prominent but not appearing marginate.

Corolla less than 3 cm. long; calyx to 1.7 cm. long, bilabiate with the lobes

obscurely lobulate; species of the eastern lowlands A. donnell-smithii.

Corolla more than 3 cm. long, calyx more than 2 cm. long.

Leaves coriaceous, oblong-elliptic to usually obovate or sometimes oblanceo-

late; plants of Caribbean coastal swamps or rivers; fruits subglobose.

A. obovata.

Leaves chartaceous or subcpriaceous, oblanceolate to oboyate-elliptic; plants

of the highlands or interior Atlantic forests; fruits ovate-cylindric,

oblongoid or elliptic-ovate.

Petioles 2-4 cm. long; inflorescence a single flower in axil of leaves; calyx

circumscissile at base; fruit oblongoid or elliptic-ovate; montane

species in western highlands A. montana.

Petioles very short, 0.2-0.5 cm. long; inflorescence terminal, usually a

single flower; calyx not circumscissile; fruit ovate-cylindric; forest

species of the lowland interior A. oblanceolata.

Amphitecna donnell-smithii (Sprague) L. Wms. Fieldiana:

Bot. 36: 22. 1973. Crescentia donnell-smithii Sprague, Bull. Herb.

Boiss. ser. 2. 6: 376. 1906. Enallagma donnell-smithii Standl. Field

Mus. Bot. 12:361. 1936.

Wet mixed forests, 300 m. or less; Alta Verapaz (type, Tuerckheim

7P53);Pet<§n. British Honduras.

Shrubs or small trees with slender branches, the nodes not enlarged, about

9 m. tall; leaves papyraceous, oblanceolate, attenuate-acuminate, attenuate to the

base, 7-16 cm. long, 1.5-4 cm. broad, sparsely glandular-lepidote; flowers solitary,

usually axillary; calyx cleft almost to the base, narrow, bilabiate, one lobe minutely

tridentate, the other bidentate, 12-17 mm. long, glabrous; corolla 2.5-3 cm. long,

the tube very narrow, the limb spreading, sparsely glandular-lepidote; ovary

densely glandular-lepidote.

The smallest-flowered of all the species of this group to be found

in our area. It is uncommon and several of the specimens seen and

presumed to belong here are sterile.

Amphitecna macrophylla (Seem.) Miers ex Baill. Rev. Hort.

465. 1882. Crescentia macrophylla Seem, in Hook. Journ. Bot. Kew

FIG. 23. Amphitecna macrophylla. A, leaf, X Yz\ B, corolla dissected, X 1; C,

calyx, X 1; D, capsule, X 1A'> E» section of stem showing cauliflorous inflores-

cence, X±M-

165

166 FIELDIANA: BOTANY, VOLUME 24

Misc. 6: 274. 1854. Neotuerckheimia megalophylla Donn.-Sm. Bot.

Gaz. 47: 258, /. 1. 1909 (type from Coban, Alta Verapaz, Tuerckheim

II 2278. Enallagma macrophylla Lundell, Wrightia 4: 170. 1971.

Moist or wet forest, usually growing on limestone at the edges of

streams, 1,300 meters or less; Alta Verapaz; Izabal. Mexico (Vera-

cruz and Tabasco; type from Teapa, Tabasco).

Shrubs or trees 4.5-12 m. high, simple or with few branches; leaves very large,

clustered near the top of the trunk, almost sessile, oblanceolate or oblong-oblance-

olate, 40-80 cm. long and 6-16 cm. broad or probably even larger, obtuse to

acuminate, long-attenuate to the base, papyraceous, blackening when dried,

glabrous; flowers borne on the thick trunk usually near its base; calyx 1.5-2.5 cm.

long, pale green, splitting bilabiately halfway to the base, somewhat glandular-

lepidote; corolla 2-5 cm. long, pale green, the lobes pale lilac-pinkish, glabrous;

ovary oblong, punctate-glandular; anthers dull purplish brown; fruit pendent,

ellipsoid, 26 cm. long or probably even larger, short-acuminate at the apex, nar-

rowed at the base, with 8 unevenly spaced longitudinal ridges, filled with white,

spongy or finally soft pulp.

Called "guiro de montafia" in Tabasco and "morro cimarron"

in Oaxaca. This plant is striking in appearance. It is plentiful along

the Rio Carcha between Coban and Carcha, probably the type

locality of Neotuerckheimia, where it forms thick stands on the

jagged limestone rocks, the trunks often leaning out across the water.

Pittier compares the mature fruits to those of cacao criollo (Theobroma

cacao), but the younger ones suggest a small wild papaya (Carica).

When cut, they show a solid spongy white mass in which the seeds,

although large, are scarcely distinguishable from the pulp. In the

locality mentioned, at the end of March the trees bore young flower

buds and almost ripe fruits. The huge leaves form a single cluster

at the end of the trunk or its branches or are arranged in several

whorls at the top of the plant.

Amphitecna montana L. Wms. Fieldiana, Bot. 36: 22. 1973.

Small tree in the montane forests on the slopes of Tajamulco

and Tacana volcanoes, mostly at 2,000 m. or higher; San Marcos

(type, Steyermark 36704.) . Mexico (Chiapas).

Small forest trees 10-15 m. tall, the branches angulate at first, becoming terete

with age, glabrous; leaves alternate and scattered along the branches or sometimes

clustered toward the apex, oblanceolate to narrowly obovate-elliptic, acuminate,

cuneate to the base and rather long petiolate (for genus), with 15-20 prominent

lateral nerves anastomosing toward the margins, blades subcoriaceous, 12-30 cm.

long and 3-10 cm. broad, the petioles mostly 2-4 cm. long; inflorescences of single

flowers in axils of leaves; flowers pendent, the pedicel slender, to about 10 cm.

long; calyx fleshy, coriaceous, bilobate or possibly also trilobate to the middle,

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 167

about 3 cm. long, circumscissile at the base and falling away leaving the pedicel

and young ovary; corolla light green, narrowly campanulate, shallowly crenate-

lobate, slightly zygomorphic, about 4.5 cm. long and about 2.5 cm. broad at throat;

fruits oblongoid or elliptic-ovoid, to 15 cm. long and 8 cm. in diameter, shell thin

and fragile, the flesh hard and slightly bitter; seeds thick and the cotyledons

fleshy, notched at both top and bottom, about 18 mm. broad, 12 mm. high and

8-10 mm. thick.

A distinctive and perhaps not uncommon tree in the rather

restricted area where known. The pendent fruits are outstanding,

the petiolate leaves and circumscissile calyx distinguish it from the

several other species of the alliance.

Amphitecna oblanceolata L. Wms. Fieldiana, Bot. 36: 23.

1973.

Wet lowland forests of the Atlantic coastal plain; Pete"n (type,

Tun 14.59) ; Alta Verapaz; Izabal. Endemic.

Shrubs or small trees to 10 m. tall; branches slender, terete or angulate,

whitish, glabrous; leaves oblanceolate or elliptic-oblanceolate, acuminate, cuneate

to the base, glabrous, secondary nerves 10-15 pairs at nearly a right angle to

mid-nerves, the tips anastomosing, the blades often with small plate-shaped

glands, 9-25 cm. long and 2-7 cm. broad, petiole very short, 2-8 mm. long; in-

florescence terminal on young branches, mostly a single pedicellate flower, the

pedicel 2-3 cm. long; calyx bilabiate at anthesis, 1.5-2 cm. long and 0.5-0.8 cm.

broad at the base, the lobes broadly oblong-ovate to narrowly obovate, obtuse;

corolla campanulate-funnelform, zygomorphic, 5-lobate, 2.5-4.5 cm. long; fruit

narrowly ovate-cylindric, probably more than 15 cm. long and about 3.5-5 cm. in

diameter; seeds (nearly mature) immersed in the whitish pulp, subreniform in

outline with the embryo at the center and with lateral wings thin and obtuse.

The long narrowly ovate-cylindric fruit is distinctive.

Amphitecna obovata (Benth.) L. Wms. Fieldiana, Bot. 36: 25.

1973. Crescentia obovata Benth. Bot. Voy. Sulph. 130. t. 46. 1845.

Enallagma obovata Schumann in Engler & Prantl, Pflanzenf. IV. 3b:

247. 1895. Morrito del rio; wild calabash.

Usually in forests along rivers or in mangrove swamps near sea

level or but little above. Southern Mexico; British Honduras to

Panama and the West Indies to northern South America; southern

Florida.

Large shrubs or trees, sometimes 15 m. high but usually lower, the trunk

often crooked, the crown widely spreading, the bark thin; branchlets angulate,

becoming subterete and shallowly fissured, the nodes enlarged; leaves sessile or

nearly so, coriaceous, oblong-elliptic to obovate or oblanceolate, 7-25 cm. long,

2-9 cm. broad, abruptly acute, cuneate to rounded at the base; flowers long-

pedunculate, the calyx 2-4 cm. long, bilabiate, cleft almost to the base, with a

FIG. 24. Amphitecna obovata. A, habit, X 3^; B, corolla dissected to show

stamens and staminode, X 1; C, calyx, X 1; D, base of pistil and gland, X 3.

168

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 169

few scattered impressed plate-shaped glands on the upper half; corolla green or

tinged with purple, 4-6 cm. long, glandular-lepidote; ovary glandular-lepidote;

fruit subglobose, 6-8 cm. in diameter.

Known in British Honduras as "calabash," "morrito del rio,"

"river calabash," or "wild calabash"; "huiro de montana" (Tabasco);

"jicarillo" (Veracruz).

This plant has gone under the name of Enallagma latifolia

(Miller) Small in most of the recent floras. However, Miller's

description of Crescentia latifolia does not fit the plant to which the

name commonly has been applied. Since no authentic material

exists, the name may be considered to be an ambiguous one.

Amphitecna silvicola L. Wms. Fieldiana, Bot. 36: 25. 1973.

Moist dense forest, at about 1,400 m., probably in Pete"n. Mexico

(type from Lago Tsiskaw, Chiapas, on the Guatemalan border,

Breedlove 10064). Known only from the type.

Shrubs about 2 m. tall, the branches angulate, becoming terete, glabrous;

leaves alternate on the new growths, elliptic or oblanceolate, acuminate, short

petiolate with 7-12 lateral nerves, these lighter-marginate on the lower surfaces,

the blades 6-12 cm. long and 2.2-4 cm. broad, petiole to 1 cm. long; inflorescence

fascicles of 1-3 flowers on older wood from an enlarged multibracteolate base,

pedicels about 2 cm. long, slender; flowers white; calyx closed in bud, bilaterally

split in anthesis nearly to the base, each lobe obovate, somewhat cochleate, obtuse,

glabrous, 15-20 mm. long and 10-12 mm. broad; corolla campanulate-funnelf orm ;

crenulate-lobate, muriculate toward the apex outside, smooth and glabrous within,

4-5 cm. long when mature; stamens 4, two unequal pairs, reaching to the throat

of the corolla; staminode cylindric, 3-4 mm. long, arcuate; anthers 2-celled, the

cells explanate and joined through apex of anther, each cell about 5 mm. long; disc

entire, obscurely lobate; young ovary elongate, smooth; style about 30 mm. long,

narrowly truncate; fruits unknown.

The venation on the underside of the leaves has a lighter margin

and the veins look, at first, like the work of leaf-miners. This is

characteristic of only two species in the genus.

ANEMOPAEGMA Martius

Woody vines with simple or tripartite tendrils; branchlets terete, or sub-

quadrate, striate, the older ones with inconspicuous lenticels, the nodes with

interpetiolar ridges; pseudostipules foliaceous and conspicuous or reduced or none;

inflorescence a short axillary raceme or reduced to a single flower; calyx campanulate,

truncate or somewhat lobate, often glandular-lepidote; corolla funnelform, the

tube narrow, yellow; stamens included, anthers glabrous; capsule subterete and

compressed with subligneous valves, or ellipsoidal and somewhat terete with

relatively thin valves; seeds with corky or hyaline wings.

170 FIELDIANA: BOTANY, VOLUME 24

A genus, if such it is, with perhaps some 40 species distributed

from Mexico to Paraguay. It seems unlikely that the two species in

Guatemala belong in the same genus, as genera in the family are

divided. The generic description is written to include our species

but the key that follows will show their main differences.

Pseudostipules large and foliaceous; tendrils simple; capsule ellipsoidal with the

valves relatively thin; the seeds with broad corky opaque wings; lowland

species A. chrysoleucum.

Pseudostipules small (2-3 mm. long) or none; tendrils tripartite at the tip; capsule

compressed with valves sub ligneous; the seeds with broad hyaline wings;

highland species A. puberulum.

Anemopaegma chrysoleucum (HBK.) Sandwith, Lilloa 3:

459. 1938. Bignonia chrysoleuca HBK. Gen. & Sp. Nov. PI. 3: 134.

1819. Anemopaegma belizeanum Blake, Contr. Gray Herb. 52: 91.

1917 (type from British Honduras, Peck 957.

In ciliar forests or in swampy thickets at or near sea level;

Izabal. British Honduras; Honduras; Nicaragua. Northern South

America.

Small or large, woody vines, the branchlets subterete, striate, puberulent or

almost glabrous, the nodes ampliate and complanate, with faint interpetiolar

ridges; leaves bifoliolate, the leaflets oval or ovate-oblong, acute to short-acumin-

ate, mucronate, rounded to cuneate at the base, entire, subcoriaceous, densely im-

pressed-glandular-punctate above and beneath, otherwise almost glabrous;

Pseudostipules orbicular or elliptic, 8-20 mm. long; flowers axillary, solitary or in

3-flowered racemes; calyx 8-10 mm. long, truncate or obscurely lobate, coriaceous,

sparsely ciliate, obscurely glandular-lepidote with numerous pale plate-shaped

glands on the upper half; corolla 5-7 cm. long, yellow, minutely glandular-lepidote;

ovary glandular-lepidote; capsule 8-15 cm. long and 3-4 cm. broad, glabrous; seeds

transversely oblong, with broad corky opaque wings.

This species, like many others of the family, is known from widely

separated localities. It is expected that this erratic distribution, in

this species, may be due to insufficient collections along lowland

rivers and swamps. The seeds of this species, in contrast to most of

those of the tribe Bignonieae, would seem to be adapted for distribu-

tion by water rather than by wind.

Anemopaegma puberulum (Seibert) Miranda, Ann. Inst. Biol.

Mex. 24: 93. 1953. Chodanthus puberulus Seibert, Carnegie Inst.

Wash. Publ. 522: 425, fig. 7. 1940. Chorcha; flor canaria.

In forests or in thickets, 1,200-1,600 m. perhaps both cultivated

and native in Guatemala; El Quiche"; Guatemala. Mexico (Veracruz,

type from Chiapas) ; Honduras.

FIG. 25. Anemopaegma puberulum. A, habit, X M; B, corolla dissected, X 1;

C, calyx and style, X 1; D, pistil with subtending callus, X 2; E, capsule, X K;

F, seed, X M-

171

172 FIELDIANA: BOTANY, VOLUME 24

Branchlets pilose and sparsely glandular-lepidote; leaves opposite or sub-

opposite, bifoliolate; leaflets subcoriaceous, ovate or oblong-ovate, 6.5-11 cm.

long, 4-7 cm. broad, obtuse-acuminate, obtuse or rounded at the base, puberulent

above, densely punctate on both surfaces, impressed-glandular beneath in the

nerve axils, densely velutinous-pilose beneath with short hairs; inflorescence

axillary, the bracts linear-spathulate or subulate, 8-9 mm. long; calyx campanu-

late, truncate, denticulate, 7-8 mm. long, tomentulose and glandular-lepidote;

corolla 5.5-6.5 cm. long, sparsely fulvous-furfuraceous, the lobes 10-13 mm. long;

capsule about 12.5 cm. long and 6.5 cm. broad, strongly compressed, very obtuse

at the apex, abruptly contracted at the acute base into a stout stipe almost 3 cm.

long; seeds transversely oblong with very broad hyaline wing.

This species was originally described as a Chodanthus, a monotypic

genus known only from southern Brazil and Paraguay. Whether or

not it is an Anemopaegma is open to some question.

ARRABIDAEA DC.

Woody vines with tendrils, branchlets terete or subterete, not prominently

angled, with conspicuous interpetiolar gland fields at the nodes usually present or

with interpetiolar ridges; leaves usually ternately divided, the terminal leaflet

often replaced by a simple tendril, pseudostipules small and inconspicuous;

inflorescence often a terminal pyramidal thyrse but may be short and axillary and

rarely on old wood; flowers comparatively small, pale pink to purple; calyx

campanulate, truncate or denticulate, margin rarely thin and patelliform (in one

species), pubescent; corolla hypocrateriform or usually campanulate-funnelform,

densely pubescent outside, the lobes subequal; stamens included, glabrous; disc

annular, pulvinate; capsule linear, compressed, septicidally dehiscent, the valves

coriaceous and with longitudinal median nerves, smooth, rarely tuberculate; seeds

with membranaceous (rarely corky) wings.

The genus is perhaps the largest of the Bignoniaceae in America

and there may be as many as 100 species; they are often attractive

lianas, usually of lowland regions. Ranges of some of the species

are very great and the distribution often with great geographical

breaks. The genus needs to be revised.

Nodes with interpetiolar ridges; corolla lobes in bud prominently white-pubescent

on the limb only A. patellifera.

Nodes with interpetiolar glandular fields; corolla lobes in bud not prominently

white-pubescent on the limb only.

Leaflets glabrous or glabrate on both surfaces.

Leaves simply ternate, with 3 or 2 leaflets.

Leaflets white-barbate beneath in the nerve axils; corolla 1.5-2 cm. long.

A. floribunda.

Leaflets not barbate beneath in the nerve axils; corolla 2.5-3 cm. long.

A. chica.

Leaves 2-3 times ternate, with more numerous leaflets A. inaequalis.

Leaflets with a persistent pubescence beneath.

Flowers in a long narrow thyrse; branchlets with 2 types of pubescence,

short-tomentulose and with longer capitate-tipped hairs. .A. mollissima.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 173

Flowers in a large dichasium; branchlets with only one type of pubescence.

Leaflets ochraceous-tomentulose beneath; calyx glandular-lepidote.

A. litoralis.

Leaflets with a fine dense whitish tomentum beneath; calyx tomentulose.

Costa and nerves of the lower leaf surface densely tomentulose like the

rest of the surface; petioles about as long as the petiolules.

A. sieberi.

Costa and lateral nerves of the lower leaf surface appearing glabrous,

conspicuous in contrast with the pale surface; petioles twice as long

as the petiolules A. candicans.

Arrabidaea candicans (L. Rich.) DC. Prodr. 9: 185. 1845.

Bignonia candicans L. Rich. Act. Soc. Hist. Nat. Paris 110. 1792. A.

pachycalyx Sprague, Bull. Herb. Boiss. II. 6: 373. 1906.

British Honduras (El Cayo District, Mountain pine ridge, San

Agustin, in high alluvial forest, Lundell 6771). Mexico; Costa Rica;

Panama; Colombia to Amazonian Brazil.

Large vines as much as 25 m. long, the branchlets subterete or sub-tetragon-

ous, striate, minutely pubescent at first, becoming glabrate; leaves 2-3-foliolate,

turning somewhat reddish when dried; leaflets oblong-ovate to broadly ovate, 4-14

cm. long, 2.5-8 cm. broad, shortly obtuse-acuminate, cuneate to cordate at the base,

sparsely pubescent above and minutely glandular-lepidote, lustrous, densely and

minutely white-tomentulose beneath, except on the nerves, with some depressed

plate-shaped glands toward the base on each side of the costa, subcoriaceous;

inflorescence a long terminal pyramidal thyrse; calyx campanulate, truncate and

minutely denticulate, 3.5-5 mm. long, finely tomentulose; corolla 2-3 cm. long,

purplish, densely furfuraceous-pubescent outside; ovary densely glandular-lepidote;

capsule linear, 30 cm. long and 1 cm. broad or sometimes much shorter, glabrous.

This species, rare in our area, is similar to A. sieberi and is dis-

tinguished by the minor characters given in the key.

Arrabidaea chica (Humb. & Bonpl.) Verlot, Rev. Hort. 154.

1868. Bignonia chica Humb. & Bonpl. PL Aequin. 1: 107, t. 31. 1808.

Mashaste.

Wet thickets or forest, at or near sea level; Izabal. British

Honduras; El Salvador; Nicaragua; Brazil.

Large vines, the branches terete, glabrate, the nodes somewhat compressed

and dilated, conspicuously enlarged at the attachment of the petiole; leaves

2-3-foliolate, drying reddish, the leaflets ovate to oval, 7-9 cm. long, 3-5 cm.

broad, acutely acuminate, rounded or obtuse at the base, subcoriaceous, glabrous

or nearly so but sometimes sparsely glandular-lepidote, the costa and lateral

nerves sometimes with a sparse white pubescence, the tertiary veins prominulous

below; inflorescence a large, axillary or terminal, lax, pyramidal thyrse; calyx

campanulate, truncate and denticulate, 3-4.5 mm. long, densely tomentulose;

corolla rose or mauve, 2.5-3 cm. long, densely pubescent outside; capsule drying

reddish purple, about 25 cm. long and 1 cm. broad, glabrous except for scattered

depressed glands, lustrous.

174 FIELDIANA: BOTANY, VOLUME 24

It is probably this species that is reported by Wisdom as in use

by the Indians of the Jocotan region, Chiquimula, for dyeing, to

give a reddish brown color. Use of this plant for dyeing is widely

spread and said to be well known in some parts of South America.

Arrabidaea floribunda (HBK.) Loes. Repert. Sp. Nov. 16: 209.

1919. Bignonia floribunda HBK. Nov. Gen. & Sp. PI. 3: 104. 1819.

Pete"n. British Honduras (Honey Camp, Orange Walk District,

Lundell 674) ', southern Mexico.

Branchlets subterete, striate, with short white pubescence when young, soon

glabrate; leaves 2-3-foliolate, the leaflets ovate to ovate-oblong, 3.5-9 cm. long,

1.5-4.5 cm. broad, obtuse-acuminate, rounded or obtuse at the base, papyraceous,

glandular-lepidote above, glabrate and somewhat punctate in age, the axils of the

lateral nerves white-barbate beneath, the veins sparsely white-pubescent; in-

florescence terminal or sometimes axillary, consisting of a large dichasium; calyx

campanulate, truncate and denticulate, 2-3 mm. long, purplish-tomentulose;

corolla 1.5-2 cm. long, lavender to purple, densely reddish-pubescent, the tube

very slender, curved, the lobes deeply cut; capsule linear, usually somewhat

curved, 9-18 cm. long, 7-9 mm. broad, lustrous, glandular-lepidote, with some

impressed glands.

Maya names reported from Yucatan are "zacac," "anicab," and

"sebaque-ac;" also "cafe"-ac," a combination of Spanish and Maya.

Arrabidaea inaequalis (DC.) Schum. in Engler & Prantl,

Pflanzenf. 4, 3b: 214. 1894. Bignonia inaequalis DC. ex Splitg.

Tijdschr. Nat. Geschied. en Phys. 9: 9. 1842.

British Honduras (Machaca, Toledo District, in forest, Schipp

S580). South America.

Woody vines, 18 m. long, the branchlets terete, finely striate, drying blackish,

minutely glandular-lepidote; leaves biternate or triternate, rarely simply ternate,

the leaflets ovate-oblong to obovate-oblong, 5-10 cm. long, 2-5 cm. broad, abruptly

acuminate, cuneate to rounded at the base, papyraceous, drying fuscous, glabrous,

the veins sparsely glandular-lepidote, the axils of the lateral nerves slightly barbate

beneath; inflorescence a long, more or less pyramidal thyrse; calyx campanulate,

truncate and denticulate, 4-6 mm. long, finely brownish-tomentulose; corolla

3-3.5 cm. long, pinkish purple, densely short-pubescent; ovary densely glandular-

lepidote.

Only the collection cited is known from North America.

Arrabidaea litoralis (HBK.) Standl. Contr. U. S. Nat. Herb.

23: 1318. 1926. Seibert, Carnegie Inst. Wash. Publ. 522: 406. 1940.

Bignonia litoralis HBK. Nov. Gen. & Sp. PI. 3: 139. 1819. A. chica

var. viscida Donn.-Sm. Bot. Gaz. 20: 7. 1895 (type from Casillas,

Santa Rosa, Heyde &Lux 4550).

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 175

At 1,200 m. or less; Santa Rosa. Mexico.

Branchlets terete, striate, densely tomentulose, later glabrate, the nodes with

small interpetiolar glandular fields; leaves 2-3-foliolate, the leaflets ovate to almost

rounded, 8-10 cm. long, 3-7 cm. broad, acuminate, obtuse to cordate at the base,

drying brownish, sparsely pubescent above and glandular-lepidote, tomentulose

or glabrate beneath; inflorescence a large, axillary or terminal, somewhat ovoid

dichasium; calyx campanulate, truncate, 4-6 mm. long, densely glandular-lepidote

and with a few short capitate hairs; corolla pink, 3.5-4 cm. long, in bud densely

tomentulose, later sparsely tomentulose and glandular-lepidote; ovary densely

glandular-lepidote; capsule 21 cm. long or more, about 1 cm. broad, glabrous.

We have seen no authentic material of A. litoralis nor do we

know that A. chica var. viscida is the same as that. We follow Seibert

but suggest that monographic study may show the Guatemalan

plant to be another species.

Arrabidaea mollissima (HBK.) Bur. &Schum. in Mart. Fl. Bras.

8, pt. 2: 46. 1896. Bignonia mollissima HBK. Nov. Gen. & Sp. PI.

3: 103. 1819. A. mollicoma Blake, Contr. Gray Herb. 52: 92. 1917

(type from Nicaragua). A. seleriana Loes. Verh. Bot. Ver. Bran-

denb. 65: 99. 1923 (type from Chiquimula, Seler 3348).

Dry or moist thickets, 400 m. or less; Chiquimula; Zacapa; El

Progreso; Jutiapa; Santa Rosa. Southern Mexico; El Salvador;

Nicaragua; Panama; Venezuela and Colombia.

Branchlets terete, striate, with 2 kinds of pubescence, densely and closely

tomentulose, also glandular-pilose, the nodes with large glandular fields; petiole

scars large and conspicuous; leaves 2-3-foliolate, the leaflets ovate to rounded,

4-7 cm. long, 2.5-5 cm. broad, short-acuminate, truncate or shallowly cordate at

the base, densely pubescent on both surfaces, the hairs equal, rarely glandular-

capitate, sparsely glandular-lepidote; pseudostipules acicular-subulate, 6 mm.

long; inflorescence a terminal or axillary, long, narrow thyrse or dichasium; calyx

campanulate, truncate, obscurely denticulate, 4-6 mm. long, puberulent, the hairs

unequal; corolla pink or lavender, 4-5 cm. long, densely short-pubescent; ovary

densely and minutely glandular-lepidote; capsule 22-25 cm. long, 1.3-1.5 cm.

broad, densely brownish-tomentulose.

Called "chupachupa" in El Salvador.

Arrabidaea patellifera (Schlecht.) Sandwith, Kew Bull. 22:

413. 1968. Bignonia patellifera Schlecht. Linnaea 8: 516. 1833.

Petastoma patelliferum Miers, Proc. Roy. Hort. Soc. Lond. 3: 195.

1863. Vainilla.

Moist or dry thickets, in rather open forests or sometimes in pine

forests, 1,200 m. or less, chiefly at low elevations; Jalapa; Jutiapa;

Santa Rosa; Escuintla; Guatemala; Retalhuleu; Quezaltenango;

176 FIELDIANA: BOTANY, VOLUME 24

Huehuetenango. Southern Mexico; British Honduras to El Salvador

and Panama; southward to Amazonian Brazil.

Small or large vines, the branchlets pale-lenticellate, densely tomentulose

pilose, or glabrate, somewhat glandular-lepidote; leaflets ovate-elliptic to elliptic-

oblong or obovate-oblong, 13 cm. long and 9 cm. broad or smaller, obtuse to

acuminate or cuspidate, obtuse or narrowly rounded at the base, glabrate or often

densely pilose or almost tomentose, glandular-lepidote, especially beneath, the

nerve axils barbate beneath; pseudostipules 8 mm. long or less; inflorescence a

large, terminal or axillary, pyramidal thyrse; calyx 3-5 mm. long, saucer-shaped,

truncate, the margin undulate, somewhat glandular-lepidote; corolla deep pink or

purple, 2.5-4 cm. long; ovary sparsely glandular-lepidote; capsule 15-25 cm. long,

1.2-1.5 cm. broad, the margins of the valves somewhat thickened, minutely

glandular-lepidote and with numerous pale lenticels; seeds about 1 cm. long and

3 cm. broad.

Called "coral" or "bejuco de coral" in El Salvador. When in

flower, in herbarium specimens, this plant may be recognized at once

by the contrast between the densely tomentulose corolla lobes and

the glabrous tube, a form of corolla pubescence not found in any

other local member of the family. The plant is extremely variable

in leaf pubescence, more so than is usual in other Central American

members of this family.

Arrabidaea sieberi P. DC. in DC. Prodr. 9: 186. 1845; Sandwith,

Kew Bull. 22: 404. 1968. A. blanchetti P. DC., I. c. A. lundellii Standl.

Field Mus. Bot. 8: 48. 1930 (type from British Honduras, Lundell

831).

In forest edges or thickets, 1,200 m. or usually much less; Pete"n;

Huehuetenango. Mexico ; British Honduras ; northern South America

to Brazil.

Large woody vines, climbing to the tops of tall trees, the branchlets terete or

subangulate, striate, rather densely tomentulose; nodes with a slightly elevated

ridge extending from each side of the glandular field to the petiole; leaves bif olio-

late, drying pale green; leaflets elliptic-ovate, 6-10 cm. long, 3.5-5 cm. broad,

obtuse to shortly obtuse-acuminate, obtuse or rounded at the base, densely and

minutely tomentulose on both surfaces, greener above and glandular-lepidote;

inflorescence a very large, broad thyrse; calyx campanulate, truncate and dentic-

ulate, 3.5-4.5 mm. long, densely tomentulose and sparsely glandular-lepidote;

corolla 2.5-3.5 cm. long, pink, densely short-pubescent; ovary glandular-lepidote;

capsule linear, 18-30 cm. long, 1 cm. broad, minutely glandular-lepidote.

Called "tietie" in British Honduras, an allusion to the fact that

the tough stem, like those of other scandant Bignoniaceae, are used

as a substitute for rope and twine, especially in constructing the

framework of huts.

FIG. 26. Arrabidaea sieberi. A, habit with capsules, X K; B, corolla dissected

showing stamens (one reduced) and staminode, Xl^J G, calyx and style, X IY2;

D, seed, X 1.

177

178 FIELDIANA: BOTANY, VOLUME 24

ASTIANTHUS D. Don

Small or medium-sized trees, the branches terete; leaves simple, linear

ternate; pseudostipules small, subulate; inflorescence terminal, composed of loose

paniculate cymes and thyrses, the flowers bright yellow; calyx campanulate,

dentate, almost glabrous but with plate-shaped glands on the upper half; corolla

funnelform, pubescent on the upper third; anthers glabrous, sagittate; disc cupular;

ovary constricted at the base, oblong; capsule linear, scarcely compressed, septici-

dally dehiscent, the valves coriaceous, not nerved ; seeds small, transverse-oblong,

with broad whitish hyaline wings.

The genus consists of a single species.

Astianthus viminalis (HBK.) Baillon, Hist. PI. 10: 44. 1888.

Bignonia viminalis HBK. Nov. Gen. & Sp. PI. 3: 103. 1819. Tecoma

viminalis Hemsl. Biol. Cent. -Am. Bot. 2: 497. 1882. Chilca.

On river banks and gravel bars along streams, at 500 m. or less;

Zacapa; Jutiapa; Chiquimula; El Progreso; Santa Rosa. Southern

Mexico; El Salvador; Honduras.

Trees, sometimes 15 m. high but usually lower, almost glabrous, with a thick

ridged trunk and stout subterete branches, the bark grayish white; branchlets

terete, glabrous; leaves linear, long-attenuate to each end, 13-28 cm. long, glandu-

lar-lepidote; pseudostipules 3 mm. long or less; panicles large and many-flowered;

calyx 8 mm. long, minutely and very sparsely pubescent, with pale plate-shaped

glands on the upper half; corolla 4-5 cm. long; capsules pendent, 6-9 cm. long,

8-9 mm. thick, glabrous; seeds 2 mm. long, 6 mm. broad, with white hyaline wings.

In Guatemala this handsome tree is almost confined to the lower

Motagua Valley, where it is abundant on stream banks, the trees

often standing deep in water at the flood season. It seems to be

confined to sandy or gravelly stream beds, in this respect recalling

the closely related genus Chilopsis of northern Mexico and south-

western United States. The foliage is decidedly willow-like and of a

vivid green. The trees are a beautiful sight when covered with their

abundant, brilliant yellow blossoms.

GALLICHLAMYS Miquel

Woody vines with tendrils, the branchlets subterete, the nodes with indistinct

interpetiolar ridges; leaves mostly trifoliolate, often terminated by a simple

tendril; inflorescence an axillary raceme; calyx very large, inflated, campanulate,

its tissue thick and spongy, irregularly and deeply lobate; corolla funnelform, thin,

minutely glandular-lepidote, the tube constricted above the disc and ovary;

stamens included, the anthers glabrous; ovary ellipsoid, constricted above the

broad depressed-pulvinate disc; capsule thick, ligneous, somewhat rugose, com-

pressed-ellipsoid, septicidally dehiscent; valves without an elevated longitudinal

FIG. 27. Astianthus viminalis. A, habit, X

X 1; C, capsule, X 1; D, seed, X 4.

; B, flower partially dissected,

179

180 FIELDIANA: BOTANY, VOLUME 24

median nerve, densely glandular-lepidote; seeds transverse-oblong, with very

broad, membranaceous wings.

The genus consists of a single rather variable species.

allichlamys latifolia (L. Rich.) Schum. in Engler & Prantl,

flan enf. 4, 3b: 223, 1894. Bignonia latifolia L. Rich. Act. Soc.

Hist. Nat Par. 110. 1792. Tabebuia speciosa Standl. Field Mus.

Bot. 8: 49. 1930 (type from Middlesex, British Honduras, Schipp

S51. Boxac (Maya); bejuco negro (Pete"n, fide Lundell).

In thickets, 400 m. or less; Pete"n. British Honduras; Costa

Rica; Panama; southward to Bolivia and Brazil.

Large vines, sometimes 15 m. long, the branchlets somewhat striate, drying

purplish black, minutely glandular-lepidote when young, the nodes greatly

ampliate; leaflets ovate to ovate-oblong, cuspidate-acuminate, rounded to abruptly

cuneate at the base, minutely glandular-lepidote and punctate above and beneath,

with some impressed glands on each side of the costa; inflorescence a furfuraceous-

puberulent raceme, becoming glabrate; calyx yellow, 2.8-3.5 cm. long, minutely

furfuraceous at the apex, with scattered immersed blackish plate-shaped glands;

corolla deep golden-yellow, 6-8 cm. long or larger, the limb very broad, minutely

glandular-lepidote; capsule as much as 32 cm. long and 6-8 cm. broad, glandular-

lepidote, appearing granular; seeds about 4 cm. long and 10 cm. broad.

The species is known from only two collections in our area and

then skips Honduras and Nicaragua, where it certainly must occur,

and is found in Costa Rica and Panama.

CERATOPHYTUM Pittier

Woody vines, the branchlets terete, striate, the nodes with distinct inter-

petiolar gland fields; leaves 3-2-foliolate, the tendrils trifid; pseudostipules in-

distinct; inflorescence a terminal corymbiform thyrse; calyx coriaceous, cam-

panulate, truncate, sometimes minutely denticulate, densely glandular-lepidote,

with impressed glands on the upper half; corolla funnelform, tomentulose outside,

white or yellowish; stamens included, the anthers glabrous; disc pulvinate; ovary

oblong- linear, densely lepidote; cpasule oblong-linear, smooth, terete, somewhat

ligneous, septicidally dehiscent, gibbously thickened at the base, tapering to the

apex; seeds transverse-oblong, with broad membranaceous wings having hyaline

margins.

Three or four species, in tropical America. One other is known

from Central America.

Ceratophytum tobagense (Urban) Sprague & Sandw. Kew

Bull. 322. 1933. Anemopaegma tobagense Urban, in Fedde, Repert.

Sp. Nov. 14: 311. 1916. C. brachycarpum Pittier, Journ. Wash. Acad.

Sci. 18: 65. 1928. Adenocalymna heterophyllum Standl. Field Mus.

FIG. 28. Callichlamys latifolia. A, habit, X 1A; B, corolla dissected, X

G, capsule, X ^; D, seed, X 1.

181

182 FIELDIANA: BOTANY, VOLUME 24

Bot. 8: 49. 1930 (type from Yucatan). A. standleyanum Lundell,

Carnegie Inst. Wash. Publ. 478: 221. 1937.

Moist thickets, at or near sea level ; Pet£n ; San Marcos. Yucatan

Peninsula of Mexico; British Honduras; northern South America.

Branchlets glandular-lepidote, the leaf scars greatly enlarged; leaves mostly

trifoliolate, the tendrils recurved-trifid; leaflets ovate or ovate-elliptic, sometimes

obovate, short-acuminate, cuneate to subcordate at the base, 6-15 cm. long,

4-8 cm. broad, subcoriaceous, somewhat glandular-lepidote, otherwise glabrous

but the axils of the lateral nerves beneath somewhat barbellate; calyx 9-12 mm.

long, coriaceous, densely glandular-lepidote, with areas of somewhat impressed

glands on the upper half; corolla white with yellow throat, 6.5-8 cm. long, densely

tomentulose outside; ovary linear-oblong, glandular-lepidote; capsule as much as

28 cm. long and 3.5 cm. broad, tan or pale cinnamon-colored, dotted with immersed

plate-shaped glands; seeds 1.5 cm. long, 4 cm. broad.

Called "aguijon" in Yucatan, and "duppy beans" in British

Honduras.

The name Ceratophytum tetragonalobum (Jacq.) Sprague & Sandw.

(Kew. Bull. 1934: 223. 1934) is thought to be the same plant, and

Alwyn Gentry (in lit.} apparently concurs. Jacquin's name Bignonia

tetragonaloba is based on a detached fruit and will always be open

to a bit of doubt.

GLYTOSTOMA Miers

Woody vines, the branchlets tetragonous to subterete, striate, the young

lateral branches with numerous conspicuous cataphylls near the base, the nodes

with interpetiolar ridges; leaves simple or 2-3-foliolate, the terminal leaflets often

replaced by a simple tendril; pseudostipules small, subulate-lanceolate; inflo-

rescence axillary, usually a few-flowered pedunculate thyrse or cyme, or a fascicle

of 1-4 flowers; calyx broadly campanulate, truncate, usually denticulate, thin,

glandular-lepidote; corolla delicate, lepidote and short-pubescent, funnelform;

stamens included, the anthers glabrous; disc annular or plate-shaped; ovary oblong

to ovoid, densely and minutely tuberculate; capsule short, oblong or ellipsoid,

thick, densely echinate, loculicidally dehiscent; seeds transverse-oblong, somewhat

corky, the wings narrow, rather firm.

Perhaps 10 species, in tropical America. No other species is

known from Central America.

Clytostoma binatum (Thunb.) Sandwith, Rec. Trav. Bot.

Ne"erl. 34: 235. 1937. Bignonia binata Thunb. PL Bras. 13: 35. 1821.

Adenocalymna ocositense Donn.-Sm. Bot. Gaz. 18: 209. 1893 (type

from Quezaltenango, Donnell-Smith 2688). Petastoma ocositense

Kranzlin, in Fedde, Rep. Sp. Nov. 17: 61. 1921. C. elegans Standl.

Carnegie Inst. Wash. Publ. 461: 86. 1935 (type from British Hon-

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 183

duras, Schipp 1127). C. ocositense Seibert, Carnegie Inst. Wash.

Publ. 522:413. 1940.

Moist to wet thickets at little above sea level, sometimes in

mangrove swamps; Pete"n; Alta Verapaz; Izabal; Retalhuleu; Que-

zaltenango; San Marcos. Mexico (Veracruz, Tabasco, Chiapas);

British Honduras; Honduras south to Panama; Venezuela; Colombia;

northern Brazil.

Small or large vines, often with tendrils, the branchlets conspicuously len-

ticellate, becoming verrucose in age, glabrous, the lateral branchlets with numerous

small cataphylls toward the base; leaves bifoliolate, the leaflets elliptic-oblong to

lance-oblong, 14 cm. long and 6 cm. broad or smaller, acuminate, obtuse at the

base, coriaceous, obscurely glandular-lepidote on both surfaces, in age conspic-

uously punctate, petioles and petiolules both short, mostly 0.5-2 cm. long; pseu-

dostipules small, early deciduous, covered with large glands; inflorescences con-

sisting of 1-4-flowered fascicles, lateral, usually on obvious peduncles; calyx with

subulate teeth, somewhat glandular-lepidote, the margin ciliate, 4-7 mm. long;

corolla lavender-purple, 5-7.5 cm. long, lepidote above, rather densely puberulent

or sublepidote on the tube and toward the base; disc small, annular; ovary minutely

tuberculate; capsule ellipsoid, somewhat compressed, 5 cm. long, 4 cm. broad,

densely covered with long, stiff, spinelike bristles or tubercles.

Called "bejuco de tres lomos" in Tabasco and "trastraz" in

Honduras.

There seems to be no doubt that the Mexican and Central

American Clytostoma is the same as that found in northern South

America. The Panamanian C. isthmicum Pittier is the same, and

there are several synonyms from South America not cited here.

CRESCENTIA L.

Small or medium-sized trees, the branchlets angulate at first, becoming sub-

terete, the nodes prominent and much enlarged in age; leaves simple or trifoliolate,

spirally disposed, usually produced in fascicles on short shoots in the axils of fallen

leaves; inflorescence of 1-3 pedicellate flowers from the nodes on old wood, often

produced on the trunk below the branches; calyx large, coriaceous, deeply cleft or

variously lobate; corolla broadly campanulate, with very broad tube, transversely

plicate on one side below the middle, lepidote or glandular lepidote outside, the

lobes broadly deltoid; stamens somewhat exserted, the anthers glabrous; disk

very large, hemispheric-pulvinate; ovary 1-celled, ovoid-ellipsoid, densely glandu-

lar-lepidote; fruit large, often very large, gourdlike, globose to oval, with a hard

corky pericarp, pulpy within; seeds somewhat compressed, not winged, embedded

in the pulp.

The genus probably consists of only the two following species

both of which may have been widely distributed in pre-Columbian

times for their useful fruits.

FIG. 29. Clystostoma binatum. A, habit, X ^ B, corolla dissected, X 1; C,

calyx dissected, with style, X 1; D, capsule, side and edge views, X 1.

184

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 185

Leaves simple C. cujete.

Leaves trifoliolate C. alata.

Crescentia alata HBK. Nov. Gen. & Sp. PI. 3: 158. 1819. Mono;

jicaro (sometimes so called, but this same is usually applied to C.

cujete); simax, rutc (Chord, fide Wisdom).

Common or abundant on essentially dry but at seasons often

very wet plains and hillsides, 1,200 m. or less; Izabal (cultivated);

Alta Verapaz; Zacapa; Chiquimula; El Progreso; Baja Verapaz;

Jalapa; Jutiapa; Santa Rosa; Guatemala (Fiscal). Mexico; El

Salvador; Honduras; Nicaragua; Costa Rica. Introduced into the

Philippines and other Pacific islands.

Low trees, rarely 12 m. high, with a rounded or spreading crown, the trunk

to 50 cm. in diameter, short, the branches thick, and sometimes interlaced; bark

light brown, scaly or deeply fissured, fibrous; leaves trifoliolate, or in juvenile

stages sometimes simple or bifoliolate, the petiole broadly winged and resembling

a leaflet; leaflets sessile, linear to narrowly obovate, entire, obtuse to rounded at the

apex, cuneate at the base, coriaceous, somewhat lepidote beneath; calyx bilabiate,

deeply cleft, 1.5-2 cm. long, glabrous; corolla 6-7 cm. long, greenish and purple-

brown, sometimes with streaks of rose-purple; fruit oval or subglobose, usually

10-15 cm. long.

Sometimes called "morrito," "cuchara," and "cutuco" in El

Salvador. This tree of distinctive aspect is common throughout

most of the lower Motagua Valley, except in the lowest wettest part.

It is even more plentiful in some parts of western Guatemala, es-

pecially near the border of El Salvador, where it forms extensive

and almost pure stands of distinctive appearance, called morrales.

In such places there occurs little other vegetation except grasses and

low herbs, and such areas supply good grazing land. Through the

parts of Guatemala where this species is native, C. cujete seldom is

seen. The fruits of C. alata are never very large, although some of

those produced by C. cujete also are small. These fruits, of distinctive

appearance, often are offered for sale entire in Guatemalan markets

remote from places where the trees grow naturally, to be used in

household medicine. The pulp is used particularly in the treatment

of colds, and for other purposes. The fallen fruits, which contain a

rather sweet pulp, are eaten commonly by stock of all kinds. There

is said to be in Guatemala a form of this tree that bears exceptionally

small fruits, little larger than a hen's egg. The small fruits often

are used to fashion spinning tops for children. They have a very

important use in Guatemala for making drinking cups and various

other kinds of containers. A fruit cut in half makes two small cups,

one of which is carried by almost every Indian pedestrian, often

attached to his waist, as a drinking cup.

FIG. 30. Crescentia alata. Famous Sesse and Mocino plate 336 in the Deles-

sert Herbarium in Geneva, made in Mexico before 1800. Field Museum neg.

30803.

186

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 187

The leaves of Crescentia alata have the form of a cross, and such a

feature could not fail to draw the attention of the early Spaniards,

who quite naturally attached a supernatural significance to it.

Oviedo writing 400 years ago says: "Each leaf is in the form of a

cross, as I have drawn it; and this seems to me a very noteworthy

thing, in which appears a testimony of the cross, a thing of which

these people can not have been ignorant. These higuero trees, which

have the leaves formed like crosses, I have seen in the Province of

Nicaragua, especially in Negrando, where lies the city of Leon, and

in other parts of that country; and marveling upon the leaves, I

gathered some to show in Spain, as I did show them, and I still have

some in my possession."

Crescentia cujete L. Sp. PL 626. 1753. Xigal (Pipil of Salama) ;

jicaro; horn (Quecchi).

Planted in many parts of Guatemala, and apparently native

along the Pacific plains, perhaps also in Alta Verapaz, usually brushy

plains or in open fields, and at 350 m. or less; Pete"n; Alta Verapaz;

Baja Verapaz; Izabal (planted); Santa Rosa; Escuintla; Suchitepe"-

quez; Retalhuleu; San Marcos; El Quiche" (probably only in cultiva-

tion) . Mexico, the West Indies, south through Central America, and

south to Peru and Brazil. Cultivated in the Old World tropics.

Small or medium-sized trees, usually 10 m. high or less, with thick trunks

and dense rounded crowns; leaves short-petiolate, cuneate-oblanceolate or spathu-

late, rounded to short-acuminate at the apex, attenuate to the base, minutely

glandular-lepidote, sometimes pilose beneath on the nerves, subcoriaceous; calyx

1.5-2.5 cm. long, deeply cleft, glandular-lepidote and with small scattered im-

pressed glands; corolla yellowish white or greenish with dark purple veins, 4.5-7.5

cm. long, glandular-lepidote; fruit very variable in size and shape, often globose

and 25 cm. in diameter, frequently much smaller, and often oval.

Called "calabash" and "wild calabash" in British Honduras,

also "giiiro," and Maya names are reported as "luch" and "huaz."

The wood is light brown or yellowish brown, with fine veining of

darker color, without distinctive taste or odor; moderately hard and

heavy, tough and strong, the specific gravity 0.60; coarse-textured,

with the consistence of elm ( Ulmus), fairly easy to work, takes a

smooth finish; probably not durable. The wood is used for ox yokes,

tool handles, and vehicle parts. Thick crooked limbs often are used

in Guatemala for making saddle trees. The trees seem to afford a

good habitat for epiphytes, and they often are covered with orchids,

bromeliads, and other plants. A syrup made from the pulp of the

fruit is a popular remedy for colds, and an infusion of the leaves is

188 FIELDIANA: BOTANY, VOLUME 24

sometimes administered for dysentery. The largest fruits, globose

in form, upon the tree remind one somewhat of green pumpkins, and

a small tree loaded with these huge fruits is an almost unbelievable

sight. The dry fruits are a very important article in Guatemala and

throughout Central America and Mexico, where they have been

used since ancient times as containers for various purposes. Their

most general use is for drinking vessels, but the larger ones serve to

store all sorts of articles. Sections of the oblong forms are much

used in place of spoons. Many of the jicaras, as the cups made from

the shells are called, are handsomely decorated in colors or by

incised designs. Particularly handsome ones are made in the

neighboring republic of El Salvador. Oviedo states that the Indians

of Costa Rica and Panama had jicaras adorned with gold, "with

handles of gold, and so handsome that the most powerful king

might drink from them without reproach. These come from the

great river of San Juan, which empties into the Gulf of Uraba."

Mexico and Guatemala likewise were noted for the handsomely

designed jicaras, from which chocolate was drunk. These cups, of

course, are rounded on the bottom, so that some support must be

given them when set down, either a stand especially made for the

purpose or a mere twist of cloth or cotton. The fruits of this species

also are eaten by cattle during the dry season, but there is a belief

that the fruit often produces abortion. It is difficult to tell just where

in Guatemala C. cujete is truly native, but it probably is in the

Pacific plains, and it and C. alata seem to occupy distinct areas of

the country. The tree is planted abundantly at low elevations or

even somewhat higher because of its local commercial value. The

word jicaro appears in such place names as El Jicaro, a village of

Guatemala, and El Jicaral (a grove of jicaro trees) in Chiquimula.

The place name Jicaltepeque is believed to signify "Jicara mountain."

The wood of Crescentia cujete and probably also of C. alata has

been used from Colonial times to the present to make stirrups —

some of those of the colonial period are beautifully carved and are

real objects of art. The wood is easy to carve when still green but

when thoroughly seasoned is "like iron" and some now in the col-

lection of Mrs. Louis 0. Williams had perhaps been in use for

"hundreds" of years, going back into the colonial period (before

1821) . While the wood is occasionally used today in Central America

for stirrups no beautifully carved modern ones have been seen. Most

stirrups probably originated in Honduras where there has been a

long history of making beautifully carved objects of wood.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 189

We suspect that Crescentia cujete was widely distributed in

tropical America in the pre-Columbian era, because of the useful

"shell" and may have subsequently evolved into several forms

known today. The original area of dispersal is not easily discernible

but erratic distribution in Central America may indicate an intro-

duced tree here.

CYDISTA Miers

Woody vines with tendrils, the branches tetragonal, at least when young, with

4 distinct costae, the nodes with interpetiolar ridges; leaves simple or bifoliolate,

the tendrils simple; pseudostipules inconspicuous or large and foliaceous; inflo-

rescence an axillary or terminal thyrse or raceme, usually few-flowered, the flowers

white to lavender or purple; calyx campanulate, truncate or bilabiate, variously

dentate or lobate, glandular-lepidote, with or without depressed glands on the

upper half; corolla funnelform, thin, glandular-lepidote outside; disc none; stamens

included, the anthers glabrous; ovary densely glandular-lepidote; capsule linear,

compressed, septicidally dehiscent, the valves smooth, with elevated and thickened

margins, the median nerve inconspicuous or slightly impressed; seeds transversely

semi-ellipsoid, brown, compressed, with broad firm opaque wings, these often with

hyaline margins.

Species perhaps 15, all in tropical America. Apparently all the

Central American ones are represented in Guatemala. — The genus

Cydista (1863) differs in no substantial way from Lundia P. DC.

(1838).

Pseudostipules foliaceous; leaflets cordate at the base C. diversifolia.

Pseudostipules inconspicuous; leaflets not cordate.

Lower leaf surface without impressed glands in the nerve axils C. potosina.

Lower leaf surface with impressed glands in the axils of the lateral nerves.

Calyx bilabiate; flowers produced on leafless branches C. heterophylla.

Calyx truncate; flowers produced with the leaves.

Branchlets tetragonous; leaves glandular-lepidote only. . .C. aequinoctialis.

Branchlets almost terete; leaves pilose and glandular-lepidote beneath.

C. sarmentosa.

Cydista aequinoctialis (L.) Miers, Proc. Roy. Hort. Soc. Lond.

3: 191. 1863. Bignonia aequinoctialis L. Sp. PI. 623. 1753. Bejuco

de ajo.

Moist or wet thickets, 1,400 m. or less; Pete"n; Izabal. Mexico;

British Honduras to Panama; West Indies; South America.

Branches tetragonous and usually with 4 distinct costae, glandular-lepidote,

the nodes with distinct interpetiolar ridges; leaves bifoliolate, drying reddish

brown or purplish; leaflets ovate to oblong or narrowly elliptic, acuminate and

somewhat cuspidate, obtuse to subcordate at the base, 15 cm. long and 7 cm.

broad or smaller, subcoriaceous, lustrous, glandular-lepidote on both surfaces,

190 FIELDIANA: BOTANY, VOLUME 24

otherwise glabrous, with impressed glands in the axils of the lateral nerves, es-

pecially toward the base of the blade; pseudostipules inconspicuous; inflorescence

of terminal few-flowered racemes or thyrses in which the main axis is suppressed;

calyx truncate-campanulate, 5-8 mm. long, glandular- lepidote, with impressed

glands on the upper part; corolla 4.5-8 cm. long, lavender to rose with darker

veins, sometimes almost white with lavender streaks; capsule 40 cm. long or less,

2-2.5 cm. broad, densely glandular-lepidote.

Cydista aequinoctialis we have accepted much in the sense of

Seibert. The species is assumed to occur widely in Central America

but the complex is much in need of study. However it may include

C. sarmentosa given below.

Cydista diversifolia (HBK.) Miers, Proc. Roy. Hort. Soc. Lond.

3: 192. 1863. Bignonia diversifolia HBK. Nov. Gen. & Sp. PI. 3:

104. 1819. Pleonotoma diversifolium Bur. & Schum. in Mart. Fl.

Bras. 8, pt. 2: 274. 1897. Bejuco de ajo.

Moist or dry thickets, 700 m. or less; Pete"n; Zacapa; Chiquimula;

Retalhuleu; El Quiche. Mexico; British Honduras; El Salvador;

Honduras; Nicaragua; Costa Rica; West Indies; northern South

America.

Branchlets tetragonous, with 4 distinct, easily separable costae, striate,

glandular-lepidote, the nodes with a narrow pubescent interpetiolar ridge; leaves

bifoliolate or sometimes simple, the leaflets broadly ovate to oblong-ovate, 4-10

cm. long, acuminate, cordate at the base or sometimes rounded, sparsely glandular-

lepidote above, densely so beneath, sometimes pubescent beneath, usually 3-

nerved, with large submerged glands in the axils of the lateral nerves; pseudostipules

large, foliaceous, broadly ovate or rounded, 6-20 mm. long; inflorescence a terminal

or axillary thyrse; calyx bilabiate or irregularly dentate, 4-5 mm. long, somewhat

glandular-lepidote; corolla pinkish lavender or purple, 3-4 cm. long, glandular-

lepidote; ovary glandular-lepidote, pubescent at the base; capsule 30 cm. long

and 1.5 cm. broad, densely glandular-lepidote, the longitudinal median nerve

somewhat impressed.

Maya names reported from Yucatan are "chacnetoloc," "anicab,"

"zolac," "tsolac," and "xcolac."

Cydista heterophylla Seibert, Carnegie Inst. Wash. Publ. 522:

417. 1940.

Pete*n; Izabal. Southern Mexico; British Honduras; Honduras;

Nicaragua; Costa Rica; Panama.

Large vines, the branchlets tetragonous, subterete at maturity, striate,

glandular-lepidote, ampliate at the nodes, usually leafless at flowering time; leaves

simple or rarely bifoliolate, the tendrils simple; leaflets papyraceous, ovate,

8-15 cm. long, 4-9.5 cm. broad, acutely acuminate, truncate to subcordate at the

base, glandular-lepidote, 3-nerved, glandular beneath in the axils of the lateral

nerves; pseudostipules small; inflorescences axillary, laxly racemose, borne on

FIG. 31. Cydista diversifolia. A, habit, X 1A'>

calyx and stype, X 2; D, capsule, about X ^.

, corolla dissected, X 1; C,

191

192 FIELDIANA: BOTANY, VOLUME 24

leafless branches; calyx subcoriaceous, 5-7 mm. long, campanulate, bilabiate,

densely glandular-lepidote; corolla purplish pink, 5-7 cm. long, glandular-lepidote;

stamens included, the anthers glabrous; ovary linear, densely glandular-lepidote;

capsules unknown.

This is one of the more common of the bignoniaceous vines in

dry lowland regions of Central America and is conspicuous because

it usually is without leaves at flowering time. Although good

collections are at hand we have not a single collection of the plant in

fruit. The junior author suspects that this plant does not belong

in the genus Cydista but until fruits are known its proper generic

designations will not be known.

Cydista potosina (Schum. & Loes.) Loes. in Fedde, Repert. Sp.

Nov. 16: 209. 1919. Arrabidaea potosina Schum. & Loes. Bull. Herb.

Boiss. 3: 618. 1895. Clytostoma mayanum Standl. Carnegie Inst.

Wash. Publ. 461: 86. 1935 (type from Pet&i, Yaxha-Remate road,

Lundell 4008). Bejuco de coral.

Thickets and second-growth forest, 300 m. or less; Pete"n; Alta

Verapaz; Escuintla; Retalhuleu. Southern Mexico; British Hon-

duras; El Salvador.

Small or large vines, the branchlets tetragonous, with 4 easily detachable cos-

tae, striate, glandular-lepidote especially when young, with some very short

pubescence near the nodes, surrounded at the base by more or less persistent, small

bracts or cataphylls; nodes with a narrow, conspicuous elevated interpetiolar

ridge; leaves bifoliolate, the leaflets elliptic to oblong-elliptic or obovate-elliptic,

acuminate, rounded or obtuse at the base, sparsely glandular-lepidote, hirtellous

beneath along the costa and nerves, without glands in the axils of the lateral nerves;

pseudostipules small, broadly sickle-shaped; inflorescence a few-flowered terminal

raceme; calyx campanulate, subtruncate, dentate, 4.5-5.5 mm. long, glandular-

lepidote; corolla white or cream with lavender throat, or lavender throughout, 4.5-

5.5 cm. long, densely glandular-lepidote; capsule somewhat glandular-lepidote, 20-

25 cm. long, 3-4.5 cm. broad; seeds brown, with firm broad wings having a hyaline

margin.

The Maya names "ekixil" and "xekkixi" are reported from

Yucatan. The capsules are relatively shorter and broader than in

most other members of the genus.

Cydista sarmentosa (Bertol.) Miers, Proc. Roy. Hort. Soc.

Lond. 3: 191. 1863. Bignonia sarmentosa Bertol. Fl. Guat. 425. 1840

(type from "Salto de Forola" [Torola], Escuintla, Velasquez). Arra-

bidaea guatemalensis Schum. & Loes. in Engler, Bot. Jahrb. 23: 129.

1896 (type from Retalhuleu, Bernoulli & Carlo 2056). Cydista

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 193

pubescens Blake, Contr. U. S. Nat. Herb. 24: 23. 1922 (type from La

Florida, Dept. Copan, Honduras). Ito; campana.

Moist or dry thickets, 1,000 m. or less; El Progreso; Jutiapa;

Retalhuleu. Southern Mexico; Honduras; Nicaragua; Costa Rica;

Panama.

Branches tetragonous when young, in age subterete, striate, sparsely pubescent

and glandular-lepidote, the nodes dilated, with an indistinct interpetiolar ridge;

leaves bifoliolate, drying brownish or fuscous, lustrous; leaflets rounded-ovate to

ovate, 7-13 cm. long, 4-11 cm. broad, abruptly short-acuminate, obtuse to broadly

rounded at the base, subcoriaceous, sparsely pubescent above, densely pilose

beneath with spreading hairs, with large fields of impressed glands in the axils of

the lateral nerves; pseudostipules small, caducous, densely pubescent; inflo-

rescences terminating the branchlets, usually composed of 3 few-flowered thyrses;

calyx campanulate, truncate, denticulate, glandular-lepidote, with impressed

glands on the upper half; corolla pink, 4.5-6 cm. long, densely papillose-glandular;

capsule about 40 cm. long, with thickened and raised margins, lustrous, glandular-

lepidote; seeds with almost opaque wings having hyaline margins.

GODMANIA Hemsley

Medium-sized or rather large trees; leaves deciduous, opposite, long-petiolate;

digitately 5-9-foliolate; leaflets usually 7 or 9, petiolulate, entire, or in juvenile

leaves often coarsely dentate; flowers small, numerous, in dense terminal corymbs;

calyx small, broadly campanulate, with 5 short obtuse teeth; corolla ventricose-

campanulate, barbate within, the limb somewhat bilabiate, the lobes broad,

rounded, unequal, spreading or recurved; stamens included, the anthers pilose,

their cells oblong, divergent, the filaments short-pilose; staminode capitate; disc

annular; capsule long and slender, cylindric, costate, loculicidally dehiscent, the

valves coriaceous; seeds with a long hyaline wing on each side.

The genus consists of a single species.

Godmania aesculifolia (HBK.) Standl. in Standl. & Calderon,

Lista Prel. El Salvador 200. 1925; Contr. U. S. Nat. Herb. 23: 1319.

1926. Bignonia aesculi/olia HBK. Nov. Gen. & Sp. PI. 3: 109. 1819.

Tecoma fuscata DC. Prodr. 9: 218. 1845. G. macrocarpa Hemsl.

Diag. PI. Mex. 35. 1879. Palo bianco; palo de agua; cort£s (perhaps

an erroneous name) ; chorequillo (Huehuetenango) .

Mostly in rather dry thickets or forest, chiefly on the Pacific

plains, sometimes on open hillsides or in pastures, 1,300 m. or less;

Peten; Izabal; Jutiapa; Santa Rosa; Escuintla; Guatemala; Solola;

Suchitepe"quez; Retalhuleu; Huehuetenango. Southern Mexico; El

Salvador to Panama; British Guiana; Venezuela.

Small to large trees, sometimes 15 m. high, the bark gray, corky, the branch-

lets puberulent, thick; leaflets mostly oblong-obovate, sometimes cuneate-obovate

FIG. 32. Godmania aesculifolia. A, habit, X J-i; B, corolla dissected, X 3;

C, calyx and style, X 3; D, capsule, X^; E, seed, X 1A-

194

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 195

or oblanceolate, 5-17 cm. long, acute or acuminate, attenuate to the base, short-

pilose on both surfaces but often glabrate, especially above, glandular-lepidote

beneath; corymbs small and very dense, shorter than the leaves; calyx minute,

short-pilose; corolla 10-13 mm. long, yellow or yellowish, or buff -brown and tinged

with red, puberulent; capsules 70 cm. long or often shorter, 1-1.5 cm. in diameter,

often curved or contorted, about 16-costate.

Called "corte"z bianco" in El Salvador; "cacho de toro" (Chiapas) ;

"joco" (Campeche); "cuerno de chivo" (Guerrero). The name

"senorita" has been reported from Guatemala for this tree, but

doubtless in error, the informant having confused it with Bombax

ellipticum, to which that name is applied currently in Guatemala.

The tree is fairly common on the Pacific plains, and is rather con-

spicuous when almost leafless and bearing quantities of the very long

slender capsules, which remind one somewhat of those of Catalpa.

JACARANDA Jussieu. Jacaranda.

Large trees, the branchlets sub terete; leaves bipinnate, with numerous large

or small, entire, lobate, or dentate leaflets; pseud ostipules none or obscure; in-

florescence a terminal or axillary thyrse, often many-flowered; calyx tubular-

campanulate, subtruncate, or shallowly cupular and with acute deltoid teeth;

corolla funnelform or campanulate-funnelform, the tube contracted toward the

base, densely pubescent to almost glabrous outside; stamens short, the staminode

elongate and much exceeding the stamens; anther cells very unequal, one of them

greatly reduced, glabrous; disc pulvinate or stipitiform; ovary ovoid or ovoid-

ellipsoid, usually glabrous; capsule short and broad, compressed, smooth, loculi-

cidally dehiscent, the septum very narrow, the valves hard and ligneous; seeds

broadly transverse-elliptic or suborbicular, with broad hyaline wings.

Species about 40, in Central and South America. One or two

others in southern Central America.

Leaflets large, mostly 2.5-7 cm. long, often dentate J. copaia.

Leaflets small, mostly less than 1 cm. long, entire J. mimosifolia.

Jacaranda copaia (Aubl.) D. Don, Edinb. Phil. Journ. 9: 267.

1823. Bignonia copaia Aubl. PI. Guian. 650, t. 265, t. 261 J.I. 1775.

Wet forest, at or little above sea level; Izabal. British Honduras;

Honduras; Nicaragua; Costa Rica; Panama; northern South America.

Large trees, often 25-30 m. high, with thick trunks, the branches smooth,

drying somewhat purplish, the nodes slightly compressed, somewhat dilated;

leaflets numerous, elliptic to oblong or rhombic, obtuse to short-acuminate,

cuneate at the base, often or usually coarsely dentate, 2.5-7 cm. long, conspic-

uously punctate, usually sparsely pubescent beneath on the costa; calyx 4-6 mm.

long, truncate, the lobes very short, pubescent and glandular-lepidote; corolla

light blue, 3-4.5 cm. long, the limb stellate-pubescent, the tube glabrous; capsule

196 FIELDIANA: BOTANY, VOLUME 24

8-14 cm. long, 5-8 cm. broad, rounded at the apex, very hard and woody; seeds

suborbicular, 2.5 cm. long and 4 cm. wide, the wings hyaline.

We have not seen this tree in flower in Guatemala, but it is not

uncommon in the forests of Izabal. It is one of the most beautiful

and showy of Central American trees, and bears a high reputation

for beauty in Costa Rica and Panama, where it is plentiful. The

wood is oatmeal-colored or dingy white, with a somewhat satiny

luster, without distinctive odor or taste; light and soft but firm,

the specific gravity 0.40-0.47; grain straight; texture medium,

uniform; very easy to cut, saws woolly when green, finishes smoothly,

holds nails well, is not resistant to insects or decay. The wood has

some of the properties of white pine (Pinus strobus), and is suitable

for light interior construction, boxes, crates, etc.

Jacaranda mimosifolia D. Don, Edinb. Phil. Journ. 9: 264.

1823. Jacaranda; gigante.

Native of South America, from Colombia to Argentina. Planted

in vast numbers for ornament in Guatemala, chiefly at middle

elevations but also in the lowlands, most plentiful in the central

departments, and especially about Guatemala; abundantly natural-

ized in many regions, and spreading rapidly by its winged seeds;

noted in Alta Verapaz, Baja Verapaz, El Progreso, Jalapa, Jutiapa,

Escuintla, Guatemala, Sacatepe"quez, Solola, Totonicapan, Huehue-

tenango, Quezaltenango, Suchitepe"quez, and Retalhuleu, and prob-

ably planted in most of the other departments.

Small to large trees with pale bark, the branchlets glabrous or nearly so;

leaflets very numerous, oblong or oblong-lanceolate, mostly 6-8 mm. long, acute

and mucronulate, sessile, glabrous or nearly so; panicles large and many-flowered,

15-25 cm. long; calyx 2 mm. long, broadly campanulate, denticulate, almost

glabrous; corolla blue, 3.5-4.5 cm. long, subtomentose; capsule suborbicular,

glabrous, about 6 cm. long and 5 cm. broad, truncate or apiculate at the apex;

seeds 1.5-2.5 cm. broad.

Whoever loves Jacaranda trees will be delighted with Guatemala,

for in few regions is one very far from at least a small number of

them. One Jacaranda tree is a lovely sight, never to be forgotten, but

hundreds of thousands rather pall upon one. For most of the year

the trees are flowerless, and then not at all impressive. The blooming

season is a short one, usually beginning in late February or early

March, and continuing for two or three weeks. The date of flowering

is partly dependent upon temperature, and on the very hot lake

shores at Panajachel, trees were observed in full bloom January 15.

Near Pastores a tree partially in flower was noted December 26, but

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 197

other trees nearby showed no signs of flowers. The peculiar hard

flattened pods remain upon the tree through the flowering season.

The jacaranda trees are most attractive because of the vast abundance

of their blossoms and their rare coloring, blue being even less com-

mon in the tropics, perhaps, than in the north temperate zone. If

the ground beneath the trees is fairly clean, it often is covered with

a carpet of blue formed by fallen corollas, that gives a rather dazzling

effect of a reflection of the crown above.

LUNDIA DC.

Woody vines with tendrils, the branchlets subterete, with somewhat ampliate

and compressed nodes having large interpetiolar glandular fields; leaves bifoliolate

or sometimes trifoliolate, the tendrils simple; pseudostipules small, not foliaceous;

inflorescence an axillary or terminal thyrse; calyx closed in bud, variously ruptured

in anthesis, campanulate, thin, truncate or bilabiate, the upper conic portion

sometimes pushed off by the expansion of the bud; corolla campanulate-funnel-

form, densely pubescent outside; stamens included, the anthers densely villous;

disc none; ovary oblong, densely pubescent, villous at the base; capsule linear,

compressed, smooth, septicidally dehiscent, each valve with an elevated longi-

tudinal median nerve; seeds narrowly transverse-oblong, with hyaline wings.

About 12 species, in tropical America. Two other species have

been reported from southern Central America.

Lundia dicheilocalyx Blake, Contr. Gray Herb. 52: 94. 1917.

Moist or wet forest or thickets, 225 m. or less; British Honduras

(type from Toledo, Peck 495~) ; Nicaragua.

Vines 9-12 m. long, the stems as much as 5 cm. in diameter, the branchlets

tomentulose, drying blackish, with numerous pale elevated lenticels; leaves drying

brownish black, the leaflets ovate, abruptly acuminate, obtuse to cordate at the

base, sparsely puberulent above and with a few minute impressed glands, minutely

glandular-lepidote beneath and sparsely pubescent, barbellate beneath in the

nerve axils; pseudostipules very small, triangular-dentiform, 1 mm. long; calyx

bilabiate, 4-6.5 mm. long, closed in bud; corolla rose, 4-4.5 cm. long, tomentulose.

MACFADYENA A. DC.

Woody vines, the branchlets slender, subterete, with conspicuous interpetiolar

gland fields at the nodes; leaves bifoliolate, terminated by a trifid tendril, its

branches uncinate and claw-like; pseudostipules small, subulate-lanceolate;

inflorescence an axillary contracted few-flowered thyrse, cyme, or raceme, or the

flowers solitary; calyx spathaceous and cleft to about the middle, or bilabiate, the

costa of the posterior lobe produced into a short incurved hook; corolla funnelform,

yellow, glabrous, the tube expanded at the base; stamens included, the anthers

glabrous; disc flattened, pulvinate or cupuliform; ovary oblong or linear, glandular-

FIG. 33. Lundia dicheilocalyx. A, habit, X

C, calyx and style, X 3; D, anther, X 5.

; B, corolla dissected, X

198

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 199

lepidote; capsule oblong-linear or linear, compressed, smooth, septicidally de-

hiscent, the valves with an elevated longitudinal median nerve, thin-coriaceous;

seeds with a truncate base and curved upper margin, the wings firm, dark and

opaque or hyaline.

About five species, in tropical America. Only the following are

known from Central America.

Wings of the seeds hyaline; branchlets verrucose-lenticellate M. unguis-cati.

Wings of the seeds firm, dark and opaque.

Leaflets punctate beneath ; inflorescence a contracted thyrse or cyme, or a single

flower in the axils of the leaves on the main branches M. uncata.

Leaflets pubescent beneath; inflorescence of short axillary racemes terminating

short lateral branches M. mollis.

Macfadyena mollis (Sond.) Seem. Journ. Bot. 1: 227. 1863.

Spathodea mollis Sond. Linnaea 22: 561. 1849. M. guatemalensis

Blake, Contr. U. S. Nat. Herb. 24: 24. 1912 (type from banks of

Rio Izabal, Izabal, Blake 7845).

Moist or wet forest, 300 m. or less; Alta Verapaz; Izabal. Colom-

bia to Brazil.

Branchlets sub terete, striate, puberulent, sometimes with gland-tipped hairs;

petioles and petiolules densely puberulent; leaflets oval to broadly ovate, 9-14 cm.

long, 3.5-6.5 cm. broad, acutely acuminate, cuneate to rounded at the base,

glandular-lepidote above, densely puberulent beneath and somewhat glandular-

lepidote; inflorescence of 1-2 short racemes or contracted thyrses terminating short

lateral branches; calyx membranaceous, 1.5-2.5 cm. long, subspathaceous, usually

somewhat lobate, each lobe terminated by a short hook, sparsely and minutely

pubescent and with scattered plate-shaped glands near the base; corolla drying

blackish, 5.5-9 cm. long; disc thick, pulvinate or somewhat cupuliform; ovary

densely glandular-lepidote.

Macfadyena uncata (Andr.) Sprague & Sandw. Recueil Trav.

Bot. Ne'er!. 34: 215. 1937. Bignonia uncata Andr. Bot. Rep. 8: t.

530. 1908. B. uncinata G. F. W. Mey. Prim. Fl. Esseq. 210. 1818.

M. uncinata DC. Prodr. 9: 180. 1845.

Moist or wet thickets or forest, sometimes in mangrove swamps,

1,000 meters or less; Pete"n; Izabal; Escuintla; Retalhuleu; San

Marcos. Southern Mexico; British Honduras to Panama; southward

to Brazil.

Large or small vines, the branchlets minutely pubescent or lepidote; leaves

bifoliolate, terminated by a trifid tendril having hooked branches; petioles and

petiolules minutely puberulent when young; leaflets elliptic to oblong-lanceolate,

long-acuminate, cuneate-obtuse at the base, glandular-lepidote on both sides, with

a few impressed plate-shaped glands beneath near the base on each side of the

FIG. 34. Macfadyena uncata. A, habit, X Yi; B, corolla dissected, X 1; C,

calyx and style, X 1; D, capsule, X }/%', E, seed, X 1.

200

STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 201

costa; inflorescence few-flowered in the axils of the leaves on the main branches;

calyx 1-2 cm. long, subspathaceous, cleft to about the middle, terminated by a

short hook, glabrous, with a few plate-shaped glands; corolla bright yellow,

5-7 cm. long, glabrous; ovary minutely glandular-lepidote; capsule gradually

narrowed at each end, 25 cm. long and 2 cm. broad or smaller, drying blackish,

glabrous; seeds about 1.5 cm. long and 2.5 cm. broad, with firm blackish wings.

Known in Honduras by the names "una de gato," "chinacla,"

and "coral"; "bejuco verde," "bejuco bianco," and "una de iguana"

(Tabasco).

Macfadyena unguis-cati (L.) A. Gentry, Brittonia 25: 236.

1973. Bignonia unguis-cati L. Sp. PI. 623. 1753. B. unguis L. ex DC.

Prodr. 9: 146. 1845. B. unguis var. guatemalensis Schum. & Loes.

Bot. Jahrb. 23: 130. 1896 (type from Retalhuleu, Bernoulli & Cario

2057}. Doxantha unguis-cati Render, Mitt. Deutsch. Dendr. Ges.

262. 1913. Una de gato; bejuco azucena; pega-palo (fide Aguilar).

Common in moist or wet thickets or forest, sometimes in man-

grove swamps, 1,500 m. or less; Alta Verapaz; Izabal; Santa Rosa;

Escuintla; Guatemala; Solola; Suchitepe"quez ; Retalhuleu; Quezal-

tenango; San Marcos; Huehuetenango. Mexico; El Salvador;

Honduras; Nicaragua; Costa Rica; Panama; West Indies; South

America.

Small or large vines, the branchlets verrucose-lenticellate, glabrate; tendrils

with short branches, these uncinate and thickened at the apex; leaflets 2, lance-

oblong to oblong-ovate, acute or short-acuminate, acute to truncate at the base,

often drying blackish, sparsely pubescent or almost glabrous, minutely glandular-

lepidote above; flowers solitary or in fascicles of 2-3, long-pedicellate; calyx

membranaceous, crenate-lobate, glabrous, 1-2 cm. long; corolla yellow, 5-8 cm.

long, usually blackish when dried; ovary minutely glandular-lepidote; capsule

60 cm. long or less, 1-1.5 cm. broad, with numerous small pale lenticels.

Maya names of Yucatan are recorded as "xcanloac," "xanicab,"

and "ec-ki-xilac." The variety published by Schuman and Loesener

is said to differ in having more numerous flowers, evidently not a

character of importance. Plants called Doxantha unguis-cati var.

dasyonyx (Blake) Siebert, Carnegie Inst. Wash. Publ. 522: 422.

1940 (Bignonia dasyonyx Blake, Contr. Gray Herb. 52: 93. 1917; D.

dasyonyx Blake, Journ. Bot. 61: 192. 1923) differs in having some-

what more puberulent leaves than the usual form, and a puberulent

rather than glandular-lepidote ovary. It is known only from the

type, from Toledo, British Honduras (M.E. Peck 919}. This vine is

sometimes cultivated in Guatemala because of its exceptionally

showy, bright yellow flowers, produced in great abundance on the

larger vines. The species is most often represented by abundant ju-

202 FIELDIANA: BOTANY, VOLUME 24

venile plants, which can be associated with adult ones only because of

the presence of the characteristic claw-like tendrils. These young

plants creep closely along rough tree trunks by the innumerable aerial

rootlets, and often form dense mats covering the trunks. These mats

frequently are inhabited by large ants that bite severely. The leaves

of juvenile plants are very small and vary greatly in shape, being

often orbicular, and broadly rounded at the apex. It is possible that

these juvenile forms are often confused with similar juvenile forms

of Macfadyena uncata.

MARTINELLA Baillon

Woody vines, the branchlets subterete, with short spreading capitate-glandular

hairs or almost glabrous, the nodes with interpetiolar ridges; leaves bifoliolate, the

tendrils trifid; pseudostipules obsolete; inflorescence an axillary or rarely terminal,

flexuous raceme; flower buds ovoid, the calyx closed, apiculate; calyx in anthesis

tubular-campanulate, deeply and irregularly 2-3-lobate, the lobes apiculate;

corolla funnelform or campanulate-funnelform, thin, glabrous or minutely lepidote,

the tube broadened at the base, gradually constricted above the disc; stamens

included, attached high in the elongate tube; disc pulvinate or patelliform; ovary

cylindric, minutely glandular-lepidote; capsule compressed, elongate-linear, thin,

septicidally dehiscent., attenuate at each end, the valves slightly thickened on the

margins, with an indistinct median nerve; seeds transversely oblong, with broad

membranaceous wings.

About six species have been reported, all in tropical America,

some of them perhaps referable to other genera or to synonomy.

Only one occurs in North America. This genus is closely related to

Arribidaea and perhaps should include Scobinaria, which see.

Martinella obovata (HBK.) Bur. & Schum. in Mart. Fl. Bras.

8, pt. 2: 161, t. 84. 1896. Spathodea obovata HBK. Nov. Gen. & Sp.

PL 3:115. 1819.

Wet forest, at or near sea level; Izabal. British Honduras;

Costa Rica; Panama; southward to Brazil.

Branchlets terete, striate, becoming costate, with spreading gland-tipped hairs

or glabrate, the nodes with broad flat interpetiolar ridges; tendrils trifid, the

branches recurved and somewhat uncinate; leaflets 2, ovate to ovate-elliptic or

ovate-lanceolate, attenuate-acuminate, rounded or subcordate at the base, glab-

rous; flowers on slender, flexuous or deflexed pedicels; calyx 12-17 mm. long,

minutely puberulent or glabrate, with a few impressed glands; corolla 5-7 cm.

long, rose-purple, minutely lepidote; disc broad, pulvinate; ovary constricted

above the disc,, minutely lepidote; capsule drying brown, 80 cm. long or less, about

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 203

2 cm. broad, glandular-lepidote; seeds 10-17 mm. long and 4-6 cm. broad, the

wings yellowish white or yellowish brown.

MELLOA Bureau

Woody vines, the branchlets subterete, striate, with interpetiolar ridges at the

nodes; leaves bifoliolate, terminated by a trifid tendril, its branches uncinate;

pseudostipules subulate; inflorescence terminal or axillary, usually on short lateral

branchlets, thyrsoid or reduced to a single flower, some of the flowers often aborted;

calyx ampliate, membranaceous, closed in bud, spathaceously cleft in anthesis,

recurved-apiculate; corolla elongate, tubular-funnelform, almost glabrous, the

lobes ciliate; stamens included, the anthers glabrous; disc large, annular-pulvinate;

ovary oblong, glabrous; capsule oblong, compressed, ligneous, smooth, septicidally

dehiscent, the valves very thick, with a longitudinal median nerve; seeds trans-

verse-oblong, with broad hyaline wings.

One other species has been described, from South America.

Melloa quadrivalvis (Jacq.) A. Gentry, Brittonia 25: 237. 1973.

Bignonia quadrivalvis Jacq. Fragm. 37. 1801. B. populifolia DC.

Prodr. 9: 159. 1845. Melloa populifolia Britton, Ann. N. Y. Acad.

7: 188. 1893. Sinaca.

Moist or dry thickets, 500-1,000 m.; El Progreso; Jutiapa.

Southern Mexico; South America, southward to Argentina.

Branchlets becoming verrucose-lenticellate, glabrous; tendrils usually cadu-

cous; leaflets 2, ovate to elliptic or oblong, acute or obtuse, mucronulate, rounded

at the base, glabrous but with a few impressed plate-shaped glands; pseudostipules

subulate, 3 mm. long or less; inflorescence usually a thyrse in which the 2 lateral

flowers of each cyme are aborted; calyx thin, ampliate, cleft one-third its length

on one side, recurved-apiculate, 1.5-2 cm. long, 1-1.5 cm. broad, glabrous or

nearly so; corolla 5-7 cm. long, bright yellow, minutely lepidote or glabrous; ovary

glabrous; capsule 8-12 cm. long, 4-4.5 cm. broad, the valves thick, woody, with

numerous pale lenticels; seeds 1.5-4.5 cm. broad.

Both the Index Kewensis and Seibert in his account of the Mayan

Bignoniaceae give the author of the combination Melloa populifolia

as Bureau, but both are in error, since no such name is published at

the place they cite.

MUSSATIA Bureau

Woody vines, the branchlets tetragonous, with distinct ribs on the angles, the

nodes with interpetiolar ridges; leaves opposite or nearly so, 2-3-foliolate, often

terminated by a simple tendril; pseudostipules usually foliaceous; inflorescence a

terminal thyrse or dichasium; calyx very short, spreading-cupular or campanulate,

truncate, somewhat lobate or cleft; corolla funnelform, bilabiate, thin, densely

stipitate-glandular-lepidote; stamens included, the anthers glabrous; disc short,

204 FIELDIANA: BOTANY, VOLUME 24

carnose, shallowly cupular or pulvinate; ovary oblong, sulcate, densely glandular-

lepidote; capsule elongate-linear, compressed, the valves without an elevated

median nerve.

About four species, in tropical America. Only the following are

found in Central America.

Pseudostipules small, not foliaceous; corolla white or cream-colored; fertile stamens

2, with 2 short sterile ones and 1 staminode M. caudiculata.

Pseudostipules foliaceous; corolla purple or yellowish with purple stripes; fertile

stamens 4, a staminode also present M. hyacinthina.

Mussatia caudiculata (Standl.) Seibert, Carnegie Inst. Wash.

Publ. 522: 418. 1940. Petastoma caudiculatum Standl. Field Mus.

Bot. 11: 141. 1932 (type from British Honduras, Schipp S297).

Wooded swamps or wet forest, 700 m. or less; Alta Verapaz;

Izabal; Retalhuleu. British Honduras.

Large vines, the young branches tetragonous, striate, some of the striations

becoming rib-like in age, the stems then multiangular, glabrous, ochraceous, the

nodes somewhat dilated., the interpetiolar ridges often interrupted in the middle;

leaves 2-3-foliolate, the leaflets narrowly elliptic-oblong to obovate-oblong, 4-12

cm. long, 2-6 cm. broad, acute or acuminate, obtuse to truncate at the base,

lustrous and glabrous above, minutely and sparsely glandular-lepidote beneath

and becoming sparsely punctate; pseudostipules subulate, 3 mm. long; inflores-

cence a small terminal thyrse; calyx campanulate, spreading in anthesis, 4 mm.

long, truncate or short-dentate, minutely lepidote; corolla white or cream-colored,

bilabiate, broadly campanulate, 2.5 cm. long, densely stipitate-glandular-lepidote;

ovary oblong to obovate, costate; capsule 1.5 cm. broad, lustrous, glabrous.

Mussatia hyacinthina (Standl.) Sandwith, Recueil Trav. Bot.

Ne'er!. 34: 218. 1937. Tynanthus hyacinthinus Standl. Carnegie Inst.

Wash. Publ. 461:87. 1935.

Moist or wet forest, 300 m. or less; Pete"n; Alta Verapaz. British

Honduras (type from Jacinto Creek, Schipp 5661} ; Mexico (Chiapas) ;

Panama; northern South America.

Large vines as much as 18 m. long, the branches tetragonous, striate, distinctly

4-costate, the nodes with interpetiolar ridges, these usually interrupted in the

middle; leaves often subopposite, especially on younger branches, bifoliolate;

leaflets ovate to elliptic or oblong-ovate, 8-16 cm. long, 5-8 cm. broad, acute or

short-acuminate, rounded or truncate at the base, papyraceous, with numerous

plate-shaped glands on the lower surface, the nerve axils barbellate; pseudostipules

foliaceous, ovate, 18 mm. long or less; inflorescence a terminal many-flowered

thyrse or dichasium; calyx broadly campanulate, spreading, 1.5-2.2 mm. long,

truncate or somewhat lobate, glandular-lepidote or sparsely short-pubescent;

corolla 1.5-2 cm. long, purple or yellow streaked with brown, brownish-glandular-

lepidote, the glands short-stipitate; ovary oblong, acute, glandular-lepidote.

FIG. 35. Mussatia hyacinthina. A, habit, X 1A; B, flower, X 2; C, corolla

dissected showing stamens and staminode, X 2; D, calyx and style, X 2; E, stig-

ma, X 8.

205

206 FIELDIANA: BOTANY, VOLUME 24

This species was collected some 170 years ago by Sesse" and

Mocino.

NEOMACFADYA Baillon

Woody vines, the branchlets subterete, striate, with very small interpetiolar

gland fields at some of the nodes; leaves opposite or subopposite, 1-2-foliolate,

sometimes with simple tendrils; pseudostipules inconspicuous; inflorescence of

terminal few-flowered racemes on short lateral branchlets; calyx membranaceous,

cleft on one side, spathaceous; corolla campanulate, somewhat bilabiate, the tube

expanding from a very narrow base, minutely and sparsely pubescent and glandu-

lar-lepidote; stamens included, the anthers glabrous; disc annular; ovary densely

glandular-lepidote; capsule linear, compressed, smooth, septicidally dehiscent, the

valves with a slightly elevated longitudinal median nerve; seeds transversely

oblong, with translucent membranaceous wings.

The genus consists of a single species, and that exceedingly rare.

Neomacfadya podopogon (DC.) Baill. Hist. PI. 10: 26. 1888.

Spathodea podopogon DC. Prodr. 9: 205. 1845. Phryganocydia

brevicalyx Standl. Field Mus. Bot. 4: 261. 1929.

Dry or moist forest, little above sea level; Pete"n. British Hon-

duras (type of P. brevicalyx from Tower Hill, Orange Walk District,

Karling 39} ; Mexico (Yucatan) ; Cuba.

Branchlets pubescent or glabrate, with pale elevated lenticels; leaflets ovate-

oblong or elliptic-oblong, obtuse to short-acuminate, cuneate to cordate at the

base, subcoriaceous, glandular-lepidote, especially beneath, becoming punctate,

the nerve axils barbellate beneath; calyx closed in bud, cleft on one side halfway

to the base in an thesis, 16-18 mm. long; corolla pinkish lavender, 4-5 cm. long;

capsule about 11 cm. long and 1.3 cm broad, sparsely glandular-lepidote.

The plant is unusual in not having glandular fields at every node,

but often has indistinct interpetiolar ridges where these fields are

lacking.

PACHYPTERA DC.

Woody vines, the branches more or less tetragonous, costate and striate, the

nodes with conspicuous interpetiolar gland fields, similar fields also present at the

nodes of the inflorescence and at the apex of the petiole; leaves 2-3-foliolate, the

tendrils trifid; axillary buds usually 3, arranged serially and forming a conspicuous

row of superposed subulate pseudostipules; inflorescence an axillary or terminal

raceme; calyx narrowly campanulate, truncate, lobate or denticulate, densely

puberulent, with large plate-shaped glands on the upper half; corolla narrowly

funnelform, thin, puberulent, the lobes with rows of glands outside; stamens

included, the anthers densely villous; disc pulvinate; ovary elliptic-oblong or

oblong-linear, 4-angulate, pubescent; capsule oblong or oblong-linear, attenuate

or apiculate, thick and subcylindric, loculicidally dehiscent; capsule valves smooth,

with or without a distinct longitudinal nerve, puberulent and dotted with immersed

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 207

plate-shaped glands; seeds transversely oblong, the wings somewhat corky, of the

same color as the body, with a narrow hyaline margin.

Perhaps five species, in tropical America. Only the following

occurs in Central America.

Pachyptera kerere (Aubl.) Sandwith, Recueil Trav. Bot. Ne'er!.

34: 219. 1937. Bignonia kerere Aubl. PL Guian. 644, t. 260. 1775.

P. foveolata DC. Prodr. 9: 175. 1845. Tanaecium zetekii Standl.

Contr. Arnold Arb. 5: 140, t. 19. 1933.

Wet thickets, at or little above sea level; Izabal. British Hon-

duras; Costa Rica; Panama; southward to Amazonian Brazil.

Branchlets puberulent when young, glabrate in age, the nodes with gland

fields and with a straight interpetiolar ridge above the glands; leaves 2-3-foliolate,

the leaflets ovate to ovate-lanceolate, 9-18 cm. long, 3-11 cm. broad, usually

acuminate, rounded and slightly cordate at the base, sparsely and minutely

glandular-lepidote beneath, minutely puberulent on the veins; pseudostipules

subulate, sharp-pointed, serially arranged in 3 pairs, 7 mm. long or less; inflores-

cence short, nodose, about 5 cm. long; calyx 7-11-mm. long, densely puberulent,

with large plate-shaped glands in vertical rows on the upper half; corolla pale

pink, white, or creamy white, 5.5-8 cm. long, minutely and rather densely puberu-

lent, the lobes with 2 irregular rows of conspicuous glands near the base; anthers

densely white-villous; ovary elliptic, 4-angulate, densely and minutely pubescent;

capsule 10-25 cm. long, 2-4 cm. broad, densely and minutely pubescent and with

scattered immersed glands.

The flowers have a strong odor resembling that of roach powder.

PANDOREA Spach

Slender woody vines, the stems striate, somewhat dilated at the nodes, with

interpetiolar lines; leaves opposite, pinnate, the leaflets mostly 5 or more, entire

or dentate; tendrils none; inflorescence a terminal thyrse, often leafy at the base;

calyx small, campanulate or cupular, truncate or with 5 short teeth; corolla

tubular-funnelform, pubescent or almost glabrous, the limb somewhat bilabiate;

stamens 4 and didynamous, the fifth rudimentary; ovary oblong or oval; capsule

elliptic-oblong, acuminate, cuneate at the base, the valves coriaceous; seeds with

a broad membranaceous wing.

About seven species, in Australia and the Netherlands East

Indies. Cultivated widely in tropical regions.

Pandorea jasminoides (Lindl.) Schum. in Engler & Prantl,

Pflanzenf. 4, 3b: 230. 1894. Tecoma jasminoides Lindl. Bot. Reg. 23:

t. 2002. 1837.

Native of Australia; cultivated frequently for ornament in

Central America, and seen occasionally in Guatemala, especially

about Coban and Guatemala City.

208 FIELDIANA: BOTANY, VOLUME 24

Large vines, glabrous or nearly so; leaflets 5-9, short-petiolulate, subcoriaceous,

ovate to lance-oblong, mostly 4-5.5 cm. long, acute or obtuse, entire; panicles

few-flowered, the flowers long-pedicellate; calyx campanulate, 6-7 mm. long, very

shallowly lobate and with broad subapiculate lobes; corolla 4.5-5.5 cm. long, white

or pale pink, rather densely puberulent outside with rather thick, spreading,

short hairs.

Called "jazmin de Italia" in El Salvador, because the foliage

and flowers do suggest those of some species of Jasminum, although

the flowers are much larger than in that genus.

PARAGONIA Bureau

Woody vines, the branchlets subterete, lenticellate, the nodes with interpetiolar

ridges; leaves bifoliolate, the tendrils 2-3-fid, often deciduous; petioles with plate-

shaped glands along the upper side, especially near the upper end; pseudostipules

small, subulate-lanceolate; inflorescence usually a many-flowered terminal dicha-

sium, rarely a simple thyrse; calyx campanulate, truncate or slightly lobate;

corolla funnelform, velutinous-tomentose outside, rather thick; stamens included,

the anthers glabrous; disc large, cupular; ovary subcylindric, glandular-lepidote;

capsule elongate-linear, subcylindric, acuminate,minutely and densely tuberculate,

septicidally dehiscent, the valves with an elevated longitudinal costa; seeds

narrowly transverse-oblong, the wings broad, membranaceous, translucent,

brownish.

The genus consists of probably a single species.

Patagonia pyramidata (L. Rich.) Bureau, Vid. Medd. Naturh.

For. 104. 1894. Bignonia pyramidata L. Rich. Act. Soc. Hist. Nat.

Paris 110. 1792.

Moist or rather dry thickets or in forest, 1,200 m. or less; Pete"n;

Alta Verapaz; Izabal; Jutiapa; Escuintla; Solola; Retalhuleu. British

Honduras to Panama; southward to Peru and Brazil.

Often large vines, the branchlets with numerous elevated pale lenticels,

glandular-lepidote; leaflets ovate to obovate, elliptic, or oblong-elliptic, 8-17 cm.

long, 3-9 cm. broad, short-acuminate, obtuse or rounded at the base, subcoria-

ceous, minutely glandular-lepidote; pseudostipules 5-7 mm. long, usually longi-

tudinally striate in age; calyx campanulate, 5-9 mm. long, densely and minutely

tuberculate, appearing coarsely and densely tomentose, with a few impressed

plate-shaped glands; corolla deep rose or pinkish purple, drying chocolate brown,

3-6 cm. long, densely velutinous-tomentose; ovary glandular-lepidote; capsule 45

cm. long or shorter, 1.5 cm broad; seeds about 1.5 cm. long and 4.5 cm. broad,

the wings pale brownish.

Called "tietie" in British Honduras and "bejuco de casa" in El

Salvador. Guatemalan specimens have been reported as Bignonia

laurifolia Vahl.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 209

PARMENTIERA DC.

Reference: Louis 0. Williams, Parmentiera, Fieldiana, Bot. 36 (4) :

27-29. 1973.

Small or medium-sized trees, the branches subterete, usually armed with a

short spine at each node, the nodes ampliate to form the spines; leaves opposite

or subopposite, often alternate on young branchlets, often fasciculate in the axils

of the spines, trifoliolate or simple on young shoots, the petioles usually narrowly

winged; at least near the apex; flowers large or small, greenish, solitary or fasciculate

from nodes on old wood, or terminating the branchlets; calyx spathaceous, closed

in bud, cleft on one side in anthesis and soon deciduous, glandular-lepidote; corolla

campanulate and somewhat curved, the limb somewhat bilabiate; stamens slightly

exserted, the anthers glabrous; disc large, pulvinate; ovary oblong, glandular-

lepidote, 2-celled; fruit elongate, subcylindric, indehiscent, with fleshy pericarp,

smooth or costate; seeds small, numerous, not winged, imbedded in pulp.

There are seven species of the genus, all in tropical North America;

one of these is the "candle tree" of Panama, P. cerifera Seeman,

which bears a large number of showy smooth yellowish fruits 30-120

cm. long, strongly suggestive of wax candles. One other species,

Parmentiera millspaughiana L. Wms. occurs in the Yucatan Peninsula

and may eventually be found in Pete"n.

Calyx 2.5-3.5 cm. long; corolla 5-6.5 cm. long; fruits prominently costate, 11-17

cm. long P. aculeata.

Calyx 1-1.5 cm. long; corolla about 1.5 cm. long; fruits smooth or essentially so,

up to 6 cm. long P. parvi flora.

Parmentiera aculeata (HBK.) L. Wms., Fieldiana, Bot. 36:

27. 1973. Crescentia aculeata HBK. Nov. Gen. & Sp. PI. 3: 158.

1819. Parmentiera edulis DC. Prodr. 9: 244. 1845. Crescentia edulis

Moc. ex DC. I.e., nomen nudum. Parmentiera aculeata Seeman, Bot.

Voy. Herald 183. 1854, nomen nudum, non Crescentia aculeatum of

HBK. Cuajilote; caiba; coxluto (Chimaltenango, fide Tejada); ixlut

(Huehuetenango, fide Tejada).

Moist or dry thickets or lowland forest, often along rocky water-

courses, chiefly at 1,200 m. or less; planted commonly about dwellings

in the drier regions; Peten; Alta Verapaz; Baja Verapaz; El Progreso;

Chiquimula; Santa Rosa; Guatemala; Suchitepe"quez ; Retalhuleu;

San Marcos; El Quiche". Southern Mexico; British Honduras; El

Salvador; Honduras.

Small or sometimes rather large trees with usually broad, dense crowns,

the trunk short and thick, the bark pale, the branches with short, stout, somewhat

incurved spines at the nodes; leaves minutely lepidote, the petioles long, winged;

leaflets 3, entire, elliptic to obovate, acute or obtuse, cuneate at the base, 4-8 cm.

FIG. 36. Parmentiera aculeata. A, habit with flower and immature fruit, X %;

B, terminal growth, X H? C, fruit, X K; D, corolla dissected to show stamens

and staminode, X 1; E, calyx, disk, and pistil, X 1.

210

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 211

long, the nerve axils barbate beneath; flowers fasciculate on old wood or axillary

at or near the ends of the branches; calyx 2.5-3.5 cm. long, green; corolla green

with brown-purple lines, 5-6.5 cm. long; fruit pale yellow, 11-17 cm. long, 2-3 cm.

or more in diameter, often curved, conspicuously costate, acute or acuminate.

Called "cow okra" in British Honduras; "pepino de arbol"

(Yucatan); "cat"; "caat" (Yucatan, Maya). The usual name in

Guatemala, "cuajilote," is of Nahuatl origin, and signifies maize-ear

tree. Cuajilote is the name of a village in Jalapa. The tree is

abundant on the dry hills between El Rancho and Salama, and less

frequent in other dry regions. It is widely spread in cultivation in

Guatemala, but the trees are not very numerous, since the fruit is

not highly esteemed except in the very dry regions where there is

often a shortage of fruits and other food. The fruit is fried or other-

wise cooked before being eaten, and it is often stuffed with meat or

other articles. It is said to be eaten greedily by pigs where available.

At Aguacatan it was stated that the Indians employ the ripe fruit

for making sweets.

Parmentiera parviflora Lundell, Lloydia 3: 211. 1940.

Wet forest and on stream banks to 300 m.; Pete"n (Tun 835).

Mexico (type from Chiapas, Matuda 3210).

Shrubs or small trees, a meter or taller, the branches slender, terete, obscurely

puberulent at first, with 1 or 2 short porrect spines at the nodes, these sometimes

bearing small leaflets; leaves 1 or more from the nodes, opposite or alternate,

trifoliolate or simple, with sparse peltate scales, especially below, — leaves when

simple 2-4 at a node, oval, about 1 cm. long, — when trifoliolate the mid-leaflet

lanceolate or ovate-lanceolate, acuminate, (2-)3-7 cm. long, the lateral leaflets

smaller, suborbicular to ovate, apex rounded or acute, 1-2 cm. long or even less;

petioles slender, canaliculate, 1-3 cm. long (or even less); inflorescence terminal,

a single short pedicellate flower; calyx spathaceous, about 1-1.5 cm. long; corolla

narrowly campanulate-tubular, about 1.5 cm. long, obscurely bilabiate; anthers

reaching the throat; fruits terete, smooth or obscurely costate, somewhat arcuate,

fusiform, up to 6 cm. long and 1.5 cm. in diameter; seeds bilobate, about 8 mm.

long and 9 mm. broad.

This is the species with the smallest flowers and fruits of the tree

species known in the Guatemalan- Yucatecan area and is closely

related to P. millspaughiana L. Wms.

PITHECOCTENIUM Martius

Woody vines, the branchlets angulate, with ridges or 6-8 detachable costae,

the nodes with interpetiolar ridges; leaves 2-3-foliolate, the tendrils trifid or twice

trifid; pseudostipules often evident, oblong or spathulate; inflorescence a terminal

raceme or thyrse; calyx coriaceous, campanulate, truncate and often denticulate;

212 FIELDIANA: BOTANY, VOLUME 24

corolla subcoriaceous, campanulate-funnelform, densely tomentose or stellate-

furfuraceous except near the base of the tube; anthers glabrous; disc conspicuous,

pulvinate; ovary ellipsoid, contracted above the disc., densely and softly spinulose;

capsule ellipsoid, compressed, ligneous, septicidally dehiscent, densely echinate to

smooth; seeds transverse-oblong, wih a broad thin body, the wings very broad,

hyaline.

About 20 species in tropical America.

The genus Phadranthus Miers is not considered distinct and the

species known only from cultivation in Guatemala is placed here.

Corolla 4-6 cm. long; capsule echinate; common native plant P. echinatum.

Corolla 9-11 cm. long; capsule not echinate; cultivated plant. . .P. buccinatorium.

Pithecoctenium buccinatorium P. DC. in DC. Prodr. 9: 195.

1845. Phadranthus buccinatorius Miers, Proc. Hort. Soc. Lond. 3:

182. 1863.

Cultivated, probably, Guatemala (Aguilar 206}. Mexico.

Large and attractive vines, the branchlets at first sparsely pilose; leaflets

oblong to ovate, oval, or oval-elliptic, 5-9 cm. long, acute to rounded at the apex,

rounded to truncate or subcordate at the base, glabrous above, densely covered

beneath with rather large, pale scales; pseudostipules broadly ovate, obtuse or

acute, to 1.5 cm. long or more; calyx 1-1.5 cm. long, 5-dentate, densely tomentose;

corolla 9-11 cm. long, rather densely tomentose, the tube narrow for some distance

above the base, then gradually dilated to a throat almost 2 cm. broad, the lobes

about 2 cm. long; capsules not seen.

The plant is sometimes cultivated for ornament in Mexico and

California. It is probable that the Guatemalan specimens were

taken from a vine in cultivation at Finca La Aurora in Guatemala

City.

There is no record of this plant having been found as a wild plant

anywhere in Guatemala. It is curious that this very attractive

plant, if really native of Mexico, has not been found there in more

than 100 years.

Pithecoctenium echinatum (Jacq.) Schum. in Engler & Prantl,

Pflanzenf. 4, 3b: 218. 1894. Bignonia echinata Jacq. Enum. PI.

Carib. 25. 1760. P. muricatum Mocifio ex P. DC. Prodr. 9: 194.

1845. P. hexagonum P. DC. I.e. 195. Cucharo; cucharillo; lengua de

vaca; petaquillas (seeds) ; shape (Huehuetenango) .

Dry or moist thickets, 1,500 m. or less; Baja Verapaz; Izabal;

Santa Rosa; Escuintla; Guatemala; Huehuetenango. Southern

Mexico; ranging southward to Panama; southward to Brazil.

FIG. 37. Pithecolobium echinatum. Famous Sesse & Mocino plate 217 in the

Delessert Herbarium, Geneva, made in Mexico before 1800. Field Museum neg.

30767.

213

214 FIELDIANA: BOTANY, VOLUME 24

Large or small vines, the branchlets hexagonal, somewhat pubescent and

glandular-lepidote; leaves mostly trifoliolate, the tendrils trifid or twice trifid;

leaflets ovate to suborbicular, acuminate, often abruptly so, rounded or usually

cordate at the base, 5-14 cm. long, 4-10 cm. broad, papyraceous, glandular-

lepidote, pubescent or glabrate above, usually more densely pubescent beneath;

pseudostipules oblong-linear or spathulate, 7-9 mm. long; inflorescence a long

narrow raceme; calyx 7-10 mm. long, densely tomentulose. with gland fields on

the upper half; corolla white or yellowish, 4-6 cm. long, densely tomentose except

near the base, there glabrous; capsule about 16 cm. long and 6 cm. broad, covered

with hard prickles 3-4 mm. long; seeds about 3 cm. long and 8 cm. broad.

Often called "peine de mico" in El Salvador and other parts of

Central America, also "pico de pato" and "bateita"; "xachextabay,"

"netoloc," "xtabay" (Yucatan, Maya). The vine is common in

many parts of the Central American lowlands, especially in rather

dry areas, and is easy of recognition because of the distinctive

capsules, whose valves are hard and woody, and covered with hard

prickles. These valves are often used as rasps for scouring and

cleaning various objects, or as graters. The tough stems are used

for tying, like those of many other Bignoniaceae. The Maya name

"xachextabay" is derived from xach or xachah, to comb, and xtabay,

an apparition in the form of a woman, dressed as a mestizo,, who

appears, combing her beautiful hair with a pod of this vine, in isolated

spots of the Yucatan villages. The generic name Pithecoctenium

signifies "monkey comb." In Huehuetenango a decoction of the

leaves is said to be used as a lotion for treating cutaneous diseases.

PODRANEA Sprague

Shrubs or vines (ours) with opposite, simple imparipinnate leaves. Inflores-

cence paniculate, of pink or white flowers; calyx regular, campanulate, 5-dentate,

inflated; corolla campanulate, narrowed to a cylindric tube below, limb slightly

bilabiate, lobes spreading, subequal; stamens 4, included, anthers divaricate when

mature, the cells free except at the apex; disc cupular; capsule linear, slightly

compressed, bilocular.

An African genus of two species of which the following is found

in cultivation in America.

Podranea ricasoliana (Tanf.) Sprague in Dyer, Fl. Cap. 4, 2:

450. 1904. Tecoma ricasoliana Tanf. Bull. Soc. Tosc. Ort. 17, tt. 1-2.

1887. Pandorea ricasoliana Baillon, Hist. PI. 10: 40. 1888. Linda;

agradecido; mirame-linda.

An African plant commonly cultivated for ornament in Guatemala

at middle and low elevations.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 215

Large or small, slender vines, glabrous throughout or nearly so; leaflets

usually 7-11, long-petiolulate, lanceolate, 3.5-5.5 cm. long, with a very narrow,

long acumination, usually serrate or serrulate, at least some of them; panicles

terminal, lax, mostly few-flowered; calyx lax and somewhat inflated, about 2 cm.

long, with a few large depressed glands above, lobate for about a third its length,

the lobes broadly ovate, ending in a long subulate tip; corolla pink, glabrous, 4.5

cm. long; capsule linear, terete, 25-30 cm. long.

This is a handsome vine with pretty pink flowers, a favorite in

many parts of Central America. In Guatemala it is much planted

in parks and gardens, as about Guatemala City, Coban, Huehue-

tenango, Puerto Barrios, and elsewhere. It is especially plentiful

about Coban, where it has a tendency to become naturalized in

the hedges.

PSEUDOCALYMMA Sampaio & Kuhlmann

Woody vines, often with the odor of garlic, the branchlets subterete, often

becoming angulate in age, with conspicuous interpetiolar gland fields at the nodes;

leaves bifoliolate, the tendrils trifid, often caducous; petioles usually with gland

fields at the apex; leaflets mostly somewhat 3-nerved, the pseudostipules small,

subulate; inflorescence a terminal or axillary raceme or thyrse; calyx campanulate,

truncate, usually minutely denticulate, without plate-shaped glands; corolla

funnelform, the tube glabrous or minutely glandular-lepidote, the lobes densely

pubescent outside; stamens included, the anthers glabrous; disc annular or pul-

vinate; ovary oblong, lepidote or glandular-lepidote; capsule elongate-linear,

compressed, smooth, septicidally dehiscent, the valves thin, with an elevated

longitudinal median nerve; seeds transverse-oblong, the wings broad, mem-

branaceous, hyaline.

About eight species, in tropical America. Only the following

are known from North America.

Calyx 15-18 mm. long, about 15 mm. broad P. sagotii var. macrocalyx.

Calyx 5-9 mm. long, about 5 mm. broad P. sagotii.

Pseudocalymma sagotii (Bur. & Schum.) Sandwith, Rec. Trav.

Bot. Ne"erl. 34: 210. 1937; L. Wms. Phytologia 25: 458. 1973.

Adenocalymma sagotii Bur. & Schum. in Mart. Fl. Bras. 82: 110.

1896. A. macrocarpa Donn.-Sm. Bot. Gaz. 40: 9. 1905. Pseudo-

calymma macrocarpum Sandwith, L c. P. alliacea var. microcalyx

Sandwith, Kew Bull. 1953: 467. 1954. Bejuco de ajo.

Moist or dry thickets, 1,400 m. or less; Zacapa; Santa Rosa;

Escuintla; Quezaltenango ; Huehuetenengo ; doubtless also in San

Marcos, since collected at Tapachula, Chiapas. Western and

southern Mexico; El Salvador to Panama; South America.

216 FIELDIANA: BOTANY, VOLUME 24

Branchlets becoming shallowly sulcate but not distinctly angulate; petioles

with inconspicuous gland fields at the apex or with a few submersed glands;

leaflets ovate to broadly ovate or rounded, 6-9 cm. long, 4-8 cm. broad, abruptly

short-acuminate or sometimes obtuse, rounded to shallowly cordate at the base,

with a few impressed plate-shaped glands beneath in the axils of the 2 basal

nerves; pseud ostipules small and inconspicuous; calyx 5-8 mm. long, minutely

puberulent or almost glabrous, the margin ciliolate; corolla lavender or rose,

5-6.5 cm. long; ovary lepidote; capsule 25-35 cm. long, 1.5-2 cm. broad, con-

spicuously punctate; seeds 12-18 mm. long, 6-7 cm. broad.

Mexican and Guatemalan material has been determined as P.

laevigatum (Bur. & Schum.) Sampaio & Kuhlmann.

Pseudocalymma sagotii var. macrocalyx (Sandwith) L. Wms.

Phytologia 25: 458. 1973. P. alliacea var. macrocalyx Sandwith.

Kew Bull. 1953: 468. 1954. P. standleyi Steyerm. Field Mus. Bot,

23: 235. 1947 (type from Quezaltenango, Steyermark 33533}.

Known in Guatemala only from lower slopes of Volcan Santa

Maria, 1,300-1,400 m.; Quezaltenango. British Guiana; Brazil.

Large glabrous woody vines with coarse stout tendrils, the branches terete,

ochraceous, the young ones green, somewhat obtusely tetragonous; leaves large,

the petioles stout, 2.5-4 cm. long; leaflets 2, thick-chartaceous or subcoriaceous,

on petiolules 1.5-2 cm. long, broadly elliptic or ovate-elliptic, 15-18 cm. long,

9-11 cm. broad, abruptly short-acuminate, obtuse or somewhat rounded at the

base, slightly paler beneath, densely and conspicuously impressed-glandular in

the axils of the lowest nerves, 3-nerved from the base, the lateral nerves about 6

on each side, the veins prominulous, laxly reticulate; inflorescences axillary and

lateral, thyrsoid, 18-27 cm. long, 6-8 cm. broad, many-flowered, the branches

compressed, the flowers long-pedicellate; calyx broadly campanulate and somewhat

inflated, 15-23 mm. long, 15-21 mm. broad, rounded at the base, the margin very

shallowly 5-lobate, the lobes somewhat unequal, about 2.5 mm. long, 4-7 mm.

broad; corolla pale lilac (on the tube) and rose-lavender, about 7 cm. long, the

tube glabrous outside, 4.7 cm long, 2 cm. broad, the lobes ovate-rounded, rounded

at the apex, 2.3 cm. long and broad, glandular-lepidote outside; stamens included,

the f-laments glabrous; anthers glabrous, the cells divaricate, 4.5-4.8 mm. long;

disc annular; ovary oblong, minutely glandular-lepidote.

PYROSTEGIA Presl

Woody vines, the branchlets angulate, with 6-8 small costae, with interpetiolar

ridges at the nodes; leaves 2-3-foliolate, the tendrils trifid, the leaflets somewhat

pellucid-punctate; pseudostipules conic, small, caducous; inflorescence a terminal

or axillary thyrse, dense and many-flowered; calyx campanulate, truncate and

denticulate, glandular-lepidote; corolla orange, narrowly long-tubular, gradually

dilated from the base, the lobes narrow, subvalvate, tomentulose outside, the

greater part of the corolla glabrous; stamens exserted, the anthers glabrous; disc

annular; ovary 4-angulate, linear, glandular-lepidote; capsule linear, compressed,

smooth, loculicidally dehiscent, acute or attenuate, the valves subcoriaceous, with

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 217

an indistinct median longitudinal nerve; seeds transverse-oblong, the wings thin,

opaque, with hyaline margins.

About four species, in South America, one of them often cultivated

in other tropical lands.

Pyrostegia venusta (Ker) Miers, Proc. Roy. Hort. Soc. 3: 188.

1863. Bignonia venusta Ker, Bot. Reg. 3: L 2^9. 1818. B. ignea

Veil. Fl. Flum. 244. 1825; 6: 1. 15. 1827. P. ignea Presl, Bot. Bemerk.

93. 1843. Chiltote; chorro de oro; chorro.

Native of Brazil and Paraguay; planted commonly for ornament

in Guatemala, at low and middle elevations; frequent in gardens of

the central region, also in the Oriente, about Coban, and in the

lowlands of San Marcos, as well as in other areas.

Branchlets somewhat glandular-lepidote and punctate, slightly pubescent at

the nodes; leaves mostly trifoliolate, the leaflets ovate, acuminate, truncate or

shallowly cordate at the base, 4-8 cm. long, 2-5 cm. broad, densely punctate

beneath and with scattered plate-shaped glands; pseudostipules 3 mm. long or

less, caducous; calyx denticulate, pubescent and glandular-lepidote; corolla orange,

5.5-7.5 cm. long, sparsely glandular-lepidote, tomentulose on the upper part of

the lobes; capsule 25 cm. long, 1-1.5 cm. broad, attenuate at the apex, sparsely

glandular-lepidote; seeds 1 cm. long and 3.5 cm. broad, the wings dark brown

except for the hyaline margins.

Called "San Carlos" in El Salvador and "triquitraque" in Costa

Rica. This is one of the most showy vines of the Bignoniaceae, and

is grown for ornament in many tropical regions of the earth. It is

common through much of Central America, at least in the lowlands,

and has become common also in Florida.

ROSEODENDRON Miranda

Large trees with the young branches somewhat quadrangular, often pubescent

with branched farinaceous hairs; leaves opposite, digitate, 5-7-foliolate, the leaflets

entire or dentate, long petiolulate, often with branched hairs and subpeltate

circular scales; inflorescence paniculate, cymose, the flowers large and yellow;

calyx membranaceous, yellowish, obconic-cylindric, tightly enclosing the base of

the corolla, without ribs, bilobulate or irregularly divided; corolla tubular-in-

fundibuliform, the limb obscurely bilabiate, 5-lobate with lobules semiorbicular;

stamens 4, didynamous, included, the anthers divaricate at maturity; staminode

small; disc suburceolate, entire; capsule linear, with 8-12 longitudinal costae,

loculicidally bivalvate, the valves subcoriaceous; seeds with a wide hyaline wing.

There are three species of the genus, two from Mexico and

northern Central America, the other from Venezuela. The genus

was named by Dr. Faustino Miranda and dedicated to the memory

of Joseph Nelson Rose, botanist at the Smithsonian Institution who

218 FIELDIANA: BOTANY, VOLUME 24

described a species of this genus, as Tabebuia, but called attention

to its differences with Tabebuia.

Roseodendron donnell-smithii (Rose) Miranda, Bol. Soc.

Bot. Mex. 29: 43, fig. 1965. Tabebuia donnell-smithii Rose in Bonn. -

Sm. Bot. Gaz. 17: 418, t. 26. 1892 (type from Cuyuta, Escuintla,

Donnell- Smith 2070). Cybastax donnell-smithii Seibert, Carnegie

Inst. Wash. Publ. 522: 392. 1940. Palo bianco; copal (fide Donnell-

Smith); cortez; cortez bianco; primavera (in U.S. lumber trade).

Forested plains little above sea level; Chiquimula; Santa Rosa;

Escuintla; Suchitepe"quez; Retalhuleu; San Marcos. Possibly south-

ern Mexico; El Salvador; Honduras; possibly nearly exterminated

as a wild plant for its valuable wood, cultivated in plantations

according to F. B. Lamb (Caribbean Forester 21: 47. 1960).

Tall trees, sometimes 35 m. high, the trunk often 60-100 cm. in diameter,

usually buttressed, the crown rounded or spreading, the bark light brown to gray,

fairly smooth or sometimes with large scales, the inner bark white or pale brown;

young branchlets minutely and sparsely pubescent with stellate hairs; leaves

deciduous, mostly 7-foliolate, sometimes 5-foliolate; leaflets long-petiolulate,

membranaceous, oblong to ovate, acuminate, rounded or subcordate at the base,

5-25 cm. long, often irregularly serrate, pubescent above and beneath along the

nerves, glabrate in age; inflorescence a large open panicle, rather densely pubescent

with glandular-capitate hairs; calyx membranaceous, bilabiate, deeply lobate,

12-15 mm. long, covered with very short gland-tipped hairs; corolla bright yellow,

4.5-6 cm. long, pubescent with gland-tipped hairs; ovary densely glandular-

lepidote; capsule 30-45 cm. long, 2-3 cm. broad, oblong in cross-section, each

valve with 5-6 costae, sparsely pubescent with short white hairs.

Called "cortez" in Honduras and El Salvador; "primavera"

(Oaxaca, Veracruz, fide Standley). The wood is pale yellow or

almost white to light yellowish brown, without distinctive odor or

taste; rather light in weight but firm, the specific gravity 0.45; grain

fairly straight to roey; texture medium to rather coarse; fairly

strong, easy to work, finishes smoothly, does not warp or check

badly, does not appear durable. The wood is exported from Guate-

mala and Mexico to the United States under the names "primavera"

and "white mahogany," as much as 300,000 board feet having been

imported in a single year. It is used principally for making furniture,

largely in the form of veneer. During the first world war it was

utilized for making airplane propellers. In Mexico and Guatemala

it is employed for making furniture, for house construction, and for

general purposes. This was one of the important lumber trees of the

Pacific plains of Guatemala. The trees bloom when leafless and

are exceptionally showy when covered with the large panicles of

FIG. 38. Roseodendron donnell-smithii. Plate XXVI from the Botanical

Gazette for the year 1892, used by permission of the University of Chicago. Mag-

nifications may be calculated from the description.

219

220 FIELDIANA: BOTANY, VOLUME 24

flowers. The usual Guatemalan name of palo bianco refers to the

whitish bark, which makes the tree easily recognizable from a

distance.

SCOBINARIA Seibert

Woody vines, the branchlets subterete, striate, conspicuously lenticellate,

dilated at the nodes and compressed, furnished there with gland fields composed

of numerous, very small glands; leaves bifoliolate, the tendrils caducous; pseu-

dostipules small, subulate; inflorescences terminal and axillary, thyrsoid; calyx

narrowly tubular-campanulate, glabrate, membranaceous, shallowly bilabiate or

lobulate-serrate; corolla rose-purple, funnelform, pilose; stamens included, the

anthers glabrous; disc annular; ovary oblong, tetragonous; capsule elongate-linear,

compressed, densely covered with rather long tubercles; seed oblong, the wings

broad, thin, with hyaline margins.

Scobinaria japurensis (DC.) Sandw. Kew Bull. 1958: 440.

1958. Tabebuia japurensis DC. Prodr. 9: 214. 1945. Adenocalymma

verrucosum Standl. Field Mus. Bot. 4: 323. 1929 (type collected near

Tela, Honduras). Arrabidaea belizensis Standl. Field Mus. Bot. 8:

48. 1930 (type from Middlesex, British Honduras, Schipp 284).

Martinella verrucosa Standl. Contr. Arnold Arb. 5: 138. 1933.

Scobinaria verrucosa Seibert, Carnegie Inst. Wash. Publ. 522: 408.

1940.

Moist or wet forest, 200 m. or less; Huehuetenango (between

Ixcan and Rio Ixcan, Steyermark 49314). British Honduras; Atlantic

coast of Honduras; Costa Rica; Panama.

Small or large vines; leaflets elliptic or broadly elliptic, 5-12 cm. long, 2.5-7

cm. broad, acute or short-acuminate, obtuse to rounded at the base, subcoriaceous,

darkening when dried, minutely glandular-lepidote, barbate beneath in the nerve

axils; inflorescence an axillary or terminal, rather few-flowered thyrse; calyx 1.5-3

cm. long, gradually expanding upward and about 14 mm. broad at the throat,

colored like the corolla; corolla rose-purple, 5-7 cm. long, sparsely short-pilose;

ovary slightly pubescent near the apex; capsule 30-40 cm. long, 1.5-2 cm. broad,

densely covered with short spine-like tubercles; seeds 1.5 cm. long, 4.5 cm. broad.

The genus is easy to recognize in fruit, since the long linear

capsule, densely covered with tubercles, is quite unlike that of any

other member of the family. The similarity of this genus to Mar-

tinella, other than the verrucose capsules, is rather great and it may

be doubted that they are really very different.

SPATHODEA Beauvois

Large trees; leaves large, pinnate, the leaflets petiolulate, entire; flowers large

and showy, red or orange, arranged in short dense racemes; calyx tomentose,

closed in bud and long-acuminate, spathaceously cleft in anthesis; corolla glabrous,

broadly ventricose-campanulate, oblique, the limb somewhat bilabiate, the lobes

broad and rounded, with undulate margins; stamens 4, didynamous, short-exserted,

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 221

FIG. 39. Scobinaria japurensis. A, habit, X K; B, fruit, X K; C, flower dis-

sected, X 13^; D, corolla from a bud, X 1.

the anthers glabrous; disc pulvinate; capsule oblong-lanceolate, compressed,

loculicidally dehiscent, the valves thick-coriaceous, cymbiform or finally explanate;

seeds compressed, with a broad hyaline wing.

Three species, in tropical Africa.

222 FIELDIANA: BOTANY, VOLUME 24

Spathodea campanulata Beauv. Fl. Owar. 1: 47, t. 27, 28.

1805.

Native of tropical Africa, now grown for ornament in many

tropical regions; a few trees are planted in Guatemala, especially in

the Atlantic lowlands and about Guatemala City.

Tall trees when fully grown, with dense rounded crowns; leaves large, the

leaflets 9-19, short-petiolulate, oval to elliptic or lance-oblong, 5-10 cm. long,

rounded and cuspidate at the apex or obtuse, obtuse or rounded at the base, entire,

glabrous above or nearly so, somewhat paler beneath, puberulent or glabrate, with

2-3 large glands at the base; racemes short, dense, with few-many flowers on thick

pedicels; calyx strongly curved, long-acuminate, 4-5 cm. long, densely fulvous-

tomentose; corolla scarlet, 10 cm. long, very broad in the throat; capsule glabrous,

about 20 cm. long.

Called "tulipan" in El Salvador, and "tulip tree" in Panama.

The tree is of recent introduction into Central America and probably

was planted first in the Canal Zone 40 or 50 years ago.

The tree is one of the finest ornamental trees introduced in Central

America. It is suitable and does well at elevations up to 1,000 m.

Thousands of small trees have been distributed through Central

America since 1950 by Escuela Agricola Panamericana.

STIZOPHYLLUM Miers

Woody vines, the branchlets terete, striate, with obscure interpetiolar ridges

at the nodes; leaves 2-3-foliolate, the tendrils simple or trifid; leaflets coarsely and

conspicuously glandular-lepidote beneath, entire or dentate; pseudostipules

inconspicuous; inflorescence a short, axillary or terminal raceme; calyx campanu-

late, membranaceous and somewhat inflated, pubescent, irregularly 5-lobate and

bilabiate, costate; corolla campanula te-funnelform, pubescent and glandular-

lepidote; stamens included, the anthers glabrous; disc thick, depressed-pulvinate;

capsule elongate-linear, compressed, smooth, septicidally dehiscent, gradually

attenuate to the apex, the valves with an elevated longitudinal median nerve;

seeds transverse-oblong, with broad membranaceous wings.

Two or three species in tropical America, only the following in

North America.

Stizophyllum perforatum (Cham.) Miers, Proc. Roy. Hort.

Soc. 3: 198. 1863. Bignonia perforate/, Cham. Linnaea 7: 667. 1832.

Adenocalymma flos-ardeae Pittier, Contr. U. S. Nat. Herb. 18: 256.

1917 (type from Panama). A. punctifolium Blake, Contr. U. S. Nat.

Herb. 24: 22. 1922 (type from Quebradas, Izabal, H. Pittier 8570).

S. punctifolium Sandw. Recueil Trav. Bot. Ne"erl. 34: 212. 1937.

Wet forest or thickets, 350 m. or less; Pete"n; Alta Verapaz;

Izabal. Mexico (Oaxaca; Yucatan); British Honduras; Panama;

southward to Brazil.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 223

Large or small vines, the branchlets densely tomentulose at first; leaflets

entire or often dentate, ovate to oval-ovate, acute to abruptly short-acuminate,

more or less cordate at the base, densely dotted, especially beneath, with depressed

saucer-shaped pellucid glands, pubescent beneath, especially along the nerves;

panicles tomentulose; calyx 10-16 mm. long, densely fulvous-puberulent, with

sparse yellowish glands near the apex; corolla cream-colored or lavender, 4.5-6 cm.

long, papillose-puberulent and sparsely glandular-lepidote; ovary densely glandu-

lar-lepidote, becoming tomentulose; capsule 30-60 cm. long, 7-8 mm. broad,

densely puberulent and somewhat glandular-lepidote.

This plant is easy to recognize because of the many large yellow-

ish glands on the lower leaf surface.

TABEBUIA Gomes

Reference: Alwyn H. Gentry, A revision of Tabebuia in Central

America, Brittonia 22: 246-264. 1970.

Large or medium-sized trees, deciduous; leaves palmately 3-7-foliolate in

Central American species, in others sometimes simple or unifoliolate, the leaflets

petiolulate, entire or dentate, the lateral ones smaller; inflorescence terminal,

loosely paniculate to capitate; calyx campanulate or tubular, unequally bilabiate,

usually 5-lobate; corolla straight or slightly curved, the limb spreading; stamens

included, the anthers glabrous, their cells divergent; staminode small; disc annular-

pulvinate or cupuliform; ovary linear or linear-oblong, glabrous or glandular-

lepidote, sometmes pubescent; capsule elongate-linear or oblong-linear, sub terete,

pendent, loculicidally dehiscent; seeds broadly winged, with hyaline or almost

opaque wings or wing sometimes reduced.

Perhaps 100 species, in tropical America. Two other species are

known in Central America and one of these, T. neochrysantha A.

Gentry, may be expected in Guatemala.

Calyx and leaflets glandular-lepidote, never with spreading or stellate hairs;

corolla purple to rose or almost white T. rosea.

Calyx and leaflets with pubescence of spreading or stellate hairs.

Corolla lavender or purple; densely puberulent or tomentulose T. palmeri.

Corolla bright yellow, glabrous or nearly so.

Young branchlets glandular-lepidote; inflorescence rather open; flowers all

open at the same time; calyx sparsely stellate-puberulent . .T. guayacan.

Young branchlets densely stellate-pubescent, often stellate-pilose; inflores-

cence dense and congested; flowers not all open at the same time; calyx

densely covered with barbate hairs and with short stellate hairs.

T. chrysantha.

Tabebuia chrysantha (Jacq.) Nicholson, Diet. Gard. 4:1. 1889.

Bignonia chrysantha Jacq. Hort. Schoenbr. 2: 45, t. 211. 1797.

Tecoma chrysantha DC. Prodr. 9: 221. 1845. Tecoma evenia Donn.-

Sm. Bot. Gaz. 20: 8. 1895, in part (type from Santa Rosa, Heyde &

224 FIELDIANA: BOTANY, VOLUME 24

Lux 3110} . Tecoma palmeri Kraenzl, in Fedde, Repert. Sp. Nov.

17: 220. 1921. Matilisguate (Alta Verapaz).

Moist or rather dry forest, often on open hillside, sometimes on

limestone, 750 m. or less; Izabal; Alta Verapaz; Baja Verapaz; El

Progreso; Chiquimula; Jutiapa; Santa Rosa. Mexico; British

Honduras to El Salvador and Panama; Colombia and Venezuela.

A medium-sized or often small tree with pale bark; young branchlets densely

stellate-tomentose; leaflets usually 5, broadly lanceolate to obovate or elliptic-

oblong, 10-18 cm. long, abruptly acuminate, obtuse at the base, usually entire

sometimes coarsely dentate, densely stellate-tomentose to glabrate beneath, less

abundantly pubescent above, sparsely glandular-lepidote; inflorescence congested

and headlike, the flowers numerous, opening at different times; calyx 8-14 mm.

long, 5-dentate, costate, densely covered with long or short, more or less branched,

fulvous hairs; corolla bright yellow, 5.5-7 cm. long, almost glabrous; ovary sparsely

stellate-pubescent; capsules 20-30 cm. long, 1.3 cm. thick, the valves thick,

somewhat rugose when dry, stellate-tomentose.

Known in El Salvador as "cortez," "cortez amarillo," "cortez

coyote," "cortez negro," and "cortez prieto;" "ahau-che/' "ha-

hauche" (Yucatan, Maya). The wood is brown, hard, heavy, strong,

durable, and finishes smoothly. It is employed for tool handles and

for general construction. The tree is a showy and rather handsome

one, blooming when devoid of leaves, in the spring months. It is

particularly plentiful on the low dry hills along the road between

El Rancho and Salama. The type material of Tecoma evenia consists

of a mixture, the flowers being those of Tabebuia chrysantha, while

the leaves are referable to T. rosea.

Tabebuia guayacan (Seem.) Hemsl. Biol. Cent.-Am. Bot. 2:

495. 1882. Tecoma guayacan Seem. Bot. Voy. Herald 180. 1854.

Cortez; corteza; guayacan.

Wet to rather dry forest, often in open places, 1,200 m. or less;

Pet&i; Alta Verapaz; El Progreso; Izabal; Chiquimula; Santa Rosa.

Mexico (Oaxaca, Vera Cruz, Tabasco, Chiapas); British Honduras

to Panama.

Large trees, sometimes 40 m. high, the trunks to 60 cm. or more in diameter,

often buttressed, the crowns narrow or rounded, the bark light tan to buff or gray-

brown, scaly, the inner bark light brown; young branchlets minutely glandular-

lepidote; leaflets usually 5, the lateral ones sometimes parted, ovate to oblong-

elliptic, acuminate, rounded to subtruncate at the base, minutely glandular-

lepidote, especially beneath, glabrate or somewhat persistently pubescent beneath,

the axils of the lateral nerves barbate; inflorescence rather open, few-flowered,

stellate-tomentulose, the flowers all open at the same time, bright yellow, appearing

STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 225

when the tree is leafless; calyx 1-1.5 cm. long, costate, sparsely and minutely

stellate-puberulent; corolla 7-9.5 cm. long, glabrous or nearly so; ovary glabrous.

The wood of this species is highly esteemed where available, and

is considered excellent for construction purposes, especially where

durability is important. The Cathedral of Panama Vieja contained

beams of this guayacdn which were reported to be perfectly sound

after having been exposed to the weather since the destruction of

the city, some 250 years ago. Called "yellow mayflower" in British

Honduras. Seibert states that the calyx is usually found punctured

at the base, possibly because hummingbirds thus obtain nectar from

them. Very closely allied to the preceeding species.

Tabebuia palmeri Rose, Contr. U. S. Nat. Herb. 1: 109, t. 11.

1891. T. nicaraguensis Blake, Contr. Gray Herb. 52: 95. 1917.

Cortex Colorado.

Moist forest, 600 m. or less; Chiquimula; El Progreso; Zacapa.

Western and southern Mexico; El Salvador; Nicaragua; Panama.

Trees of 10-15 m. or more, the branchlets glabrous except at the tips, there

minutely stellate-puberulent; leaflets usually 5, oval-oblong to oval-lanceolate or

oblong-obovate, abruptly short-acuminate, cuneate to rounded at the base, entire,

glandular-lepidote, somewhat puberulent beneath, especially along the veins and

in the nerve axils; inflorescence densely farinose-tomentulose, appearing when the

tree is leafless; calyx 5-8 mm. long, stellate-tomentulose; corolla pinkish lilac or

purple, 5-7 cm. long, densely puberulent or tomentulose; ovary glabrous; capsules

35 cm. long or shorter, about 2 cm. thick, glabrous; seeds 1.5 cm. long and 5 cm.

broad.

The wood is dark reddish brown, very hard, and heavy; durable

in contact with the soil. It is used in Mexico for railroad ties and

general construction.

Tabebuia rosea (Bertol.) DC. Prodr. 9: 215. 1845. Tecoma

rosea Bertol. Fl. Guatemalensis 25. 1840 (type, from near Escuintla,

Velasquez). Couralia rosea Donn.-Sm. Bot. Gaz. 20: 9. 1895. Tecoma

evania Donn.-Sm. I.e. 8, in part. Matilisguate (the most usual name) ;

maqueliz (Peten); matilishuate', mano de leon (perhaps an erroneous

name); macuelizo; macueliz; fresno (Huehuetenango) .

Common in moist or rather dry forest, often in open fields or

along roadsides, most abundant on the Pacific plains, but often on

steep hillsides, 1,200 m. or less; Pete"n; Alta Verapaz; Baja Verapaz;

El Progreso; Izabal; Zacapa; Jutiapa; Santa Rosa; Escuintla;

Guatemala; Solola; Suchitep£quez; Retalhuleu; San Marcos; Hue-

226 FIELDIANA: BOTANY, VOLUME 24

huetenango. Mexico; British Honduras to El Salvador and Panama;

southward to Venezuela.

Large trees, sometimes 30 m. tall, with thick straight trunks as much as a

meter in diameter, often buttressed, the crown spreading or rounded, the bark

light brown, with rather coarse, long, vertical fissures, the inner bark medium

brown; branchlets glandular-lepidote; leaflets usually 5, mostly on very long

petiolules, subcoriaceous, 10-25 cm. long, elliptic-oblong to elliptic-ovate or some-

times obovate, acute or acuminate, acute to rounded at the base, entire, densely

glandular-lepidote, the axils of the lateral nerves beneath with plate-shaped glands,

never barbate; inflorescence large and open, glandular-lepidote; calyx bilabiate,

1.5-2 cm. long, closed in bud, splitting in an thesis, densely glandular-lepidote;

corolla 6-8 cm. long, varying in color from deep rose-purple to rarely white,

glabrous; ovary glandular-lepidote; capsules about 30 cm. long and 12 mm. thick,

attenuate to each end, densely glandular-lepidote.

Called "may flower" and "May bush" in British Honduras;

"hocab" (Yucatan, Maya); "roble" (British Honduras); "roble

bianco" (Honduras); "maquiligua," "maculigua," "matilisguat" (El

Salvador); "puntilla" (Honduras). The name "matilisguate," most

usual for the species in Guatemala, is evidently of Nahuatl origin,

and probably signified "hand tree," in allusion to the fine leaflets

composing each leaf. In Guatemala the name of the tree appears as

a place name, Matilishuate, a village in Jutiapa. The wood is one

of the most important ones of Central America, being used for a

great variety of purposes — heavy construction, furniture and cabinet-

work, interior finish, boat building, carts, and many other kinds of

work. It is rather dull grayish brown with fine striping of deep

brown, often in conspicuous pattern on the tangential surface,

without distinctive odor or taste; moderately light and soft to rather

hard and heavy; specific gravity 0.62; grain usually straight, some-

times roey or wavy; texture medium; easy to work, finishes smoothly,

seasons without difficulty, is fairly durable. It is said that if the

wood is cut green and stacked to dry, it acquires a dark color,

without losing its striping, and is then of different appearance from

lumber treated in the more usual manner. A large part of the cheaper

chairs of Central America are made from this wood. When in flower,

the tree has few equals among Central American trees for beauty.

The trees blossom when leafless, mostly in the spring months,

toward the end of the dry season. They are then huge bouquets of

flowers, which vary in shade from deep rose-purple to pure white

(rarely), in their range of tints strongly suggesting the multicolored

and lovely Japanese cherry trees. They are abundant on the Pacific

plains, where from a slight elevation one may sometimes overlook a

widely spread display of beautiful color provided by matilisguate

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 227

trees. There is a fine lot of planted trees in the Parque Central of

Guatemala, in full blossom about the end of April. The blooming

season is somewhat extended and variable, and some trees may be in

flower as early as January. Some handsome trees stand near the

river at the entrance into Coban.

This species, more often than not, has been confused with the

West Indian Tabebuia pentaphylla (L.) Hemsl.

TECOMA Jussieu

Shrubs or small trees, the nodes of the branches with indistinct interpetiolar

ridges; leaves odd-pinnate, rarely simple, the leaflets serrate; pseudostipules small

and inconspicuous; inflorescence a terminal raceme or panicle; calyx campanulate,

the lobes triangular, often apiculate; corolla campanula te-funnelform or tubular-

funnelform, yellow in North American species, usually glabrous except on the

margins of the lobes; stamens included, the anthers pilose; disc shallowly cupuli-

form; ovary oblong, glandular-lepidote; capsule linear, slightly compressed, smooth,

loculicidally dehiscent, the valves subcoriaceous; seeds oblong, with broad mem-

branaceous whitish wings.

Perhaps four to five species, all except the following in South

America.

Leaflets glabrous or nearly so T. stows.

Leaflets whitish-tomentose beneath T. stans var. velutina

Tecoma stans (L.) HBK. Nov. Gen. & Sp. PL 3: 112. 1819.

Bignonia stans L. Sp. PI. ed. 2. 871. 1763. Stenolobium stans Seem.

Journ. Bot. 1: 88. 1863. Timboco; timboque; chacte (Zacapa; also

the Quecchi name); San Andres; barreto (Jutiapa).

Wet or dry thickets or open forest, frequent on dry open rocky

hillsides, often planted for ornament, 1,500 m. or less, chiefly below

1,000 m.; Baja Verapaz; El Progreso; Zacapa; Chiquimula; Jalapa;

Jutiapa; Santa Rosa; Escuintla; Guatemala; Sacatep^quez ; Suchi-

tep£quez; Retalhuleu; Quezaltenango ; San Marcos. Southwestern

United States through Mexico; El Salvador to Panama; West Indies;

South America.

Shrubs or small trees, rarely 12 m. high with trunks 25 cm. in diameter, the

branchlets terete, with numerous pale elevated lenticels, glandular-lepidote when

young; leaflets usually 7, sessile or short-petiolulate, lanceolate to ovate-lanceolate,

4-10 cm. long, attenuate-acuminate, cuneate at the base, serrate, glabrous or

nearly so, punctate beneath and sparsely pilose along the veins, the axils of the

lateral nerves barbellate; panicles often large and many-flowered; calyx 4-7 mm.

long, sparsely glandular-lepidote; corolla bright yellow, funnelform, 3.5-5 cm. long,

228 FIELDIANA: BOTANY, VOLUME 24

sparsely glandular-lepidote in bud, the lobes white-ciliate; anthers pilose; capsule

linear, attenuate to each end, brownish, 10-20 cm. long, lustrous, lenticellate.

The Maya names of Yucatan are reported as "canlol" and

"canlol-che" ; "flor amarilla" (Yucatan); "borla de San Pedro,"

"candox" (Chiapas); "San Andre's," "marchucha," "tagualaishte,"

"tache," "tacho" (El Salvador); "sardinillo" (Honduras); "sauco

amarillo" (Yucatan). This is one of the common shrubs or small

trees through most of the lower regions of Central America, perhaps

more plentiful along the dry Pacific slope. It is planted generally

about the houses for ornament, especially in regions where it does

not grow naturally, and many of the plants found apparently wild

probably are really escapes from cultivation. The flowers are showy

and handsome, and may be found at almost any season of the year.

Neither the species nor its variety were observed in the Coban

region, although the plant must be in cultivation in some of the

fincas. It is common in fincas of Retalhuleu and Mazatenango, but

apparently is not native in that region. It, or the variety, is culti-

vated as high as Quezaltenango and San Marcos. The generic

name Tecoma is derived from the Nahuatl word signifying "trumpet."

Tecoma stans var. velutina DC. Prodr. 9: 224. 1845. T. mollis

HBK. Nov. Gen. & Sp. PI. 3: 112. 1819. Stenolobium molle Seem.

Journ. Bot. 1: 90. 1863. Timboco.

Dry or moist, brushy, often rocky slopes, often in hedgerows,

1,500-2,500 m., frequently planted for ornament; El Progreso;

Guatemala; Sacatepe"quez ; El Quiche", Huehuetenango; Solola;

Quezaltenango. Southern Mexico; western South America.

Differing from the typical form of the species only in having

more abundant pubescence; young branchlets densely pubescent;

leaflets whitish-tomentose beneath, sometimes glabrate in age.

We have seen no Central American specimens of the variety

except from Guatemala, and there the variety is limited in range,

being found chiefly in the western highlands. It grows wild at a

much higher elevation than the typical form, and is apparently the

only form occurring wild in the higher mountains and in the center

and west of Guatemala. The abundant pubescence gives both

growing plants and herbarium specimens an appearance substan-

tially different from that of the species. The variety often has been

treated as a distinct species, but there is too much variation in

pubescence, apparently, to warrant the recognition of T. mollis as a

distinct species.

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 229

TECOMARIA Spach

Woody vines, the branchlets sub terete; leaves odd-pinnate, the leaflets small,

serrate; inflorescence terminal, racemose or thyrsoid, the lateral flowers of each

cymule reduced or abortive; calyx campanulate, 5-dentate; corolla somewhat

curved, with a long narrow tube; stamens exserted, the anthers glabrous; disc

annular-cupuliform; ovary glabrous; capsule oblong-linear, loculicidally dehiscent,

the valves thin-coriaceous, with a conspicuous longitudinal median costa; seeds

transverse-oblong, with broad hyaline wings.

Probably two species, native in South Africa, one of them often

planted for ornament in tropical regions.

Tecomaria capensis (Thunb.) Spach, Hist. Veg. Phan. 9: 137.

1840; Seibert, Carnegie Inst. Wash. Publ. 522: 395. 1940. Bignonia

capensis Thunb. Prodr. PI. Cap. 105. 1800. Tecoma capensis Lindl.

Bot. Reg. 13: t. 1117. 1828. Julia; flor defuego.

Native of South Africa, cultivated occasionally in Guatemala, as

in Guatemala City, Panajachel, and Coban.

Large or small vines; leaflets 7-9, ovate to broadly elliptic, 1.5-3 cm. long,

acute or short-acuminate, coarsely serrate, somewhat pubescent, pilose beneath

on the veins, stellate-tomentose along the costa; calyx 4-5 mm. long, puberulent,

costate; corolla orange-red, 4-5 cm. long, glabrous, the lobes small and short;

capsule linear, about 11 cm. long and 8 mm. broad, attenuate to each end.

Seibert states that this species is "cultivated in the higher alti-

tudes of tropical America, where it occasionally has become natural-

ized." We have seen no evidence from field work or herbarium

material that the vine has become naturalized anywhere in tropical

America. In Central America it is infrequent, and in Guatemala it

is rarely seen.

TOURRETIA Fougeroux

Herbaceous vines with tendrils, the branchlets tetragonous, the nodes with

interpetiolar ridges; leaves opposite, 2-3-ternate, the tendrils trifid, each arm of

the tendril usually dichotomous; leaflets thin, small, coarsely serrate; inflorescence

a terminal raceme, the flowers small, short-pedicellate, dimorphous, the upper

ones sterile and deciduous; calyx 2-parted, deciduous, the segments entire; corolla

slender-tubular, unequally bilabiate, the tube shorter than the calyx; fertile

stamens 4, no staminode present, the anthers glabrous; disc annular; ovary ovoid,

densely covered with short spines; capsule ovoid, densely covered with long stiff

uncinate spines, septicidally dehiscent; seeds somewhat oblong, small, with a

narrow membranaceous wing.

The genus consists of a single species.

Tourretia lappacea (L'Kter.) Willd. Sp. PI. 3: 263. 1800.

Dombeya lappacea L'He"r. Stirp. Nov. 33, t. 17. 1784. T. volubilis

230 FIELDIANA: BOTANY, VOLUME 24

J. F. Gmel. Syst. 2: 940. 1791. Guisquilito; guisquilillo; guisquilete;

amarillo.

Wet thickets or forest, 650-2,000 m.; Santa Rosa; Guatemala;

Sacatepe"quez; Chimaltenango; Suchitepe'quez; Retalhuleu; Quezal-

tenango; San Marcos; Huehuetenango. Honduras; Nicaragua;

Costa Rica; western South America.

Slender and often very large, herbaceous vines, covering large shrubs and

often good-sized trees, somewhat succulent; leaflets ovate or rhombic, acute or

acuminate, acute or obtuse at the base, 3-9 cm. long, conspicuously serrate,

glabrous, generally with scattered impressed plate-shaped glands on the upper

surface; calyx usually bright red, sometimes pale red, 10-14 mm. long, somewhat

villosulous; corolla 12-20 mm. long, sparsely villosulous; capsules 3-4 cm. long,

1.5-2 cm. thick, densely covered with long hard uncinate spines.

Called "pegapega" in El Salvador. The plant is a most abnormal

member of the Bignoniaceae, having no close resemblance to any

other American genus of the family. A botanist seeing it for the

first time would almost certainly refer it to the Cucurbitaceae. Its

fruits densely covered with long, hard, hooked spines that can pene-

trate the flesh and that closely resemble those of some cucurbits or

a large cocklebur (Xanthiuni). In Guatemala the vine is plentiful

in many regions, usually in wet forest or thickets along ravines, but

is particularly abundant in the barrancos of San Marcos, where with

Hidalgoa it often forms a dense mantle over shrubs and small trees

or along cliffs growing so densely that it is impossible to penetrate

the tangles. In this respect also it suggests some of the Cucurbita-

ceae, such as Momordica. The small red flowers are not at all

conspicuous, and the plant is a decidedly weedy one.

The species was originally described from Peru and occurs in all

the Andean countries northward. It is said to occur in Mexico but

I have seen no specimens, nor have specimens been seen from Panama.

The curious spiny fruit may account for this curious distribution.

TYNANTHUS Miers

Woody vines, the branchlets subterete, striate or subangulate, the nodes with

interpetiolar, often V-shaped ridges; leaves 3-2-foliolate, the tendrils simple;

pseudostipules often foliaceous, soon deciduous; inflorescence of axillary or terminal

thyrses or dichasia; calyx very small, turbinate or campanulate, truncate, denticu-

late; corolla very small, conspicuously bilabiate, curved, funnelform, densely

tomentulose; stamens slightly exserted, the anthers glabrous; disc very small;

ovary conic, pubescent; capsule elongate-linear, compressed, septicidally dehiscent,

the valves smooth, with an elevated longitudinal median nerve or the ligneous

STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 231

valves each with two coriaceous wings on the margins; seeds small, with broad

hyaline wings.

About 12 species, in tropical America, only three in continental

North America.

Tynanthus guatemalensis Donn-Sm. Bot. Gaz. 18: 6. 1893.

Moist thickets and forest, at or little above sea level; Pete"n;

Izabal; Quezaltenango (type from Rio Ocosito, 75 m. Donnell-Smith

1488). Mexico (Tabasco; Yucatan); British Honduras.

Large vines, the branchlets subterete or obscurely angulate, striate, minutely

glandular-lepidote, pubescent at the nodes at first, the nodes with a more or less

distinct V-shaped interpetiolar ridge; leaflets elliptic to ovate, 4-10 cm. long,

2-6.5 cm. broad, acuminate, obtuse or rounded at the base, thin, glabrous except

on the veins, with scattered plate-shaped glands beneath on each side of the costa,

the axils of the lateral nerves pitted beneath ; pseudostipules f oliaceous, caducous,

5-15 mm. long, ovate, acute; inflorescence of axillary thyrses, sometimes terminal,

the flowers cream-colored, fragrant; calyx campanulate, truncate, almost glabrous,

2 mm. long; corolla 6-8 mm. long, conspicuously bilabiate, tomentulose, the lobes

tomentulose within; stamens slightly exserted; ovary densely pubescent; capsule

unknown.

The name "cafe"-ac" is reported from Yucatan. The vine is

noteworthy for its extremely small flowers, smaller than those of all

other Central American Bignoniaceae.

XYLOPHRAGMA Sprague

Woody vines, the branchlets subterete, with interpetiolar gland fields at the

nodes; leaves 3-5-foliolate, or bifoliolate and terminated by a simple tendril,

petiolar scars conspicuous; inflorescence of dense thyrses borne in the axils of

fallen leaves; calyx campanulate, somewhat turbinate, truncate or subulate-

dentate, pubescent; corolla membranaceous, campanulate-funnelform, minutely

stellate-pubescent, the tube straight; stamens included, the anthers glabrous; disc

annular-pulvinate, somewhat lobate; ovary tetragonous, glandular-lepidote;

capsule elongate or oblong, somewhat compressed, thick, rugose, septicidally

dehiscent, the valves ligneous; seeds broadly transverse- oblong, with hyaline wings.

About six species, in tropical America, only the following in

Central America.

Xylophragma seemanniana (0. Kuntze) Sandw. Kew Bull.

1953: 469. 1954. Saldanhaea seemanniana O. Kuntze, Rev. Gen.

PI. 2: 489. 1891. Distichtis rovirosana Donn.-Sm. Bot. Gaz. 20: 7.

1895 (type from Mexico).

Wet or dry thickets or forest, 800 m. or less; Pete"n; Izabal;

Escuintla ; Huehuetenango (?) . Mexico (Tabasco) ; British Honduras ;

Costa Rica; Panama; Colombia; Venezuela; and Trinidad.

232 FIELDIANA: BOTANY, VOLUME 24

Small or large vines, usually without tendrils, the branchlets somewhat

tetragonous when young but soon becoming subterete, stellate-tomentulose, soon

glabrate; leaves long-pedunculate, the leaflets usually 3, oblong-ovate to obovate

or oval, abruptly acuminate, obtuse to subcordate at the base, thin, sparsely

stellate-tomentulose when young, glabrate in age except along the veins, pseu-

dostipules 3 mm. long or less, triangular; calyx 5-7.5 mm. long, truncate, 5-costate,

the costae extended as small teeth, somewhat tomentulose; corolla orchid-pink,

4.5-5.5 cm. long, stellate-pubescent; ovary ovoid, slightly pubescent near the base,

glandular-lepidote above; capsule oblong, 9-12 cm. long, 5 cm. broad or less,

smooth or when dry somewhat rugose; seeds 2.5 cm. long, 4.5 cm. broad, the

body pubescent.

The name "pie de gallo" is recorded from Tabasco. This has

been reported from Pete"n under the name Saldanhaea costaricensis

Kraenzl.— Sandwith has commented (Kew Bull. 22: 407. 1968) that

"if the winged border of the valves is not to be regarded as of generic

significance in this family then Saldanhaea must surely be reduced

to the earlier described Cuspidaria DC." Dr. Alwyn Gentry has

told me (in lit.} that he considers the species to belong to Xylophragma.

PEDALIACEAE. Sesame Family.

DOROTHY NASH GIBSON

Annual or perennial herbs or rarely shrubs, often with mucilaginous sap, the

stems erect; leaves opposite or the upper ones alternate; flowers axillary, usually

solitary, sometimes cymose, perfect, zygomorphic; calyx segments 5; corolla

gamopetalous, the tube usually broad, the limb usually short, somewhat bilabiate,

the lobes 5; stamens included, usually 4 and didynamous, rarely 2, the anthers

dorsifixed, the thecae parallel or sometimes discrete; a staminode often present;

disc annular; ovary mostly superior, rarely inferior, bicarpellate, the locules 2 or

4, placentation axile, the ovules several or numerous, anatropous; style filiform;

stigmas 2; fruit capsular or indehiscent, often beaked or barbed; seeds few or

numerous, smooth, the endosperm thin, the embryo straight.

About 14 genera, natives of the Old World tropics.

SESAME L.

Erect herbs, the lower leaves opposite, the upper ones alternate, or all the

leaves sometimes alternate; flowers short-pedicellate, solitary in the leaf axils;

calyx segments 5; corolla tube oblique at the base, somewhat gibbous, the limb

somewhat bilabiate, the 5 lobes spreading; stamens 4, didynamous, inserted near

the base of the corolla tube, the anthers sagittate; ovary bilocular, the ovules

numerous; fruit capsular, oblong, somewhat 4-angulate, loculicidally dehiscent.

About 12 species, in tropical Africa and Asia.

GIBSON: FLORA OF GUATEMALA 233

Sesamum indicum L. Sp. PL 634. 1753. S. orientate L., I.e.

Ajonjoli sesame.

Native of the East Indies; cultivated widely in tropical regions

for its seeds, and often more or less naturalized; cultivated rather

extensively in Guatemala and often found as a weed in rice fields or

other cultivated ground.

Erect annuals to about a meter tall, the stems simple or branched, sparsely

pilose or glabrate; leaves petiolate, the petioles usually more or less pilose, the

blades variable, the upper ones commonly lanceolate or ovate-lanceolate, the

lower ones often broadly ovate and sometimes trilobate, mostly 7-15 cm. long,

acuminate, acute at the base, the margins entire, coarsely dentate or sinuate-

dentate, glabrous or glabrate above, the lower surfaces rather densely punctate

with minute, 4-celled, mucilaginous hairs; leaves of inflorescence mostly reduced;

flowers axillary, the pedicels mostly 2-5 mm. long; calyx parted nearly to the base,

the segments linear to linear-lanceolate, 5-8 mm. long, villous outside; corolla

pink, lilac, white, or cream, 2-3 cm. long, more or less villous outside, the throat

1-1.5 cm. broad; capsule oblong, short-rostrate, 4-sulcate, 2-3 cm. long, rather

densely pubescent; seeds obovate, somewhat compressed, about 3 mm. long.

The plants are said to be known in Yucatan by the Maya name

"zicilpuuz."

In Mexico and Central America Sesamum is grown more com-

monly than in the United States, the seeds used for flavoring candy

and other confections as well as for baked goods. It has been grown

on a fairly large scale on the Pacific plains of Guatemala. The plants

are somewhat mucilaginous and have emollient properties.

MARTYNIACEAE. Unicorn Plant Family.

DOROTHY NASH GIBSON

References: G. P. Van Eseltine, A Preliminary Study of the

Unicorn Plants (Martyniaceae), N. Y. Agr. Exp. Sta. Tech. Bull.

149, 1-41. 1929; George H. M. Lawrence, Proboscidea and Other

Unicorn Plants (Martyniaceae), Baileya 5: 127-132. 1957; Richard

H. Hevly, Nomenclatural History and Typification of Martynia and

Proboscidea (Martyniaceae), Taxon 18: 527-534. 1969.

Coarse, usually viscid-pubescent herbs; leaves simple, opposite or alternate,

petiolate, the blades entire, undulate, or dentate, sometimes lobate; inflorescences

terminal and /or axillary, racemose; the flowers pedicellate; calyx with 5 free

sepals or spathelike and with 5 lobes on the dorsal side and split to the base on

the ventral side; bracts at the base of the calyx 1-2, often becoming thick and

fleshy in age; corolla gamopetalous, the tube cylindric at the base, then campanu-

late or funnelform above, often ventricose and oblique, the limb zygomorphic,

234 FIELDIANA: BOTANY, VOLUME 24

usually somewhat bilabiate, the lobes 5, the upper 2 exterior in bud; fertile stamens

4, didynamous, or 2, the second pair reduced to staminodes, with the rudiments

of a fifth stamen sometimes obvious, all inserted on the corolla tube near the base

of the expanded portion; anthers bithecous, the thecae divergent, the anthers of

each pair of stamens initially coherent; disc annular; ovary superior, bicarpellate,

the carpels united to form the unilocular ovary with 2 parietal placentae; ovules

few to many, anatropous; style slender, the stigma bilobate; fruit a horned capsule,

the exocarp somewhat fleshy and deciduous, the endocarp ligneous; seeds few-

many, black, sculptured, almost oblong, often irregularly compressed; endosperm

thin or none; embryo straight, the cotyledons large, carnose.

Although Van Eseltine recognized five genera, the family as

treated by most authors today consists of three genera, native to

the New World tropics, and cultivated in many parts of the world

for their ornamental fruits. Two occur in Guatemala.

Calyx spathelike, shallowly lobate above, cleft to the base below; fertile stamens

4; horns of capsule about equalling or surpassing the body Proboscidea.

Calyx not spathelike, the 5 sepals free; fertile stamens 2; horns of capsule much

shorter than the body Martynia.

MARTYNIA L.

Erect, often branching, viscid annuals, glandular-pubescent to glandular-

villous throughout; leaves opposite, petiolate, the blades more or less angular-ovate

to broadly triangular, cordate and palmate-nerved at the base; inflorescences

terminal, racemose, the flowers bibracteate, pedicellate; calyx segments 5, free,

irregular; corolla rather abruptly campanulate from the short-cylindrical basal

portion of the tube, ventricose, the limb somewhat bilabiate, oblique, the lobes 5;

fertile stamens 2, staminodes 2, with the fifth stamen very rudimentary or absent;

anthers bithecous; ovary superior; style slender; stigma bilobate; fruit a horned

capsule, the body ovoid, somewhat compressed dorsiventrally, the exocarp glandu-

lar-tomentose, soon separating from the hard, woody endocarp, this with 4 stout

ribs above and 4 below, the central fissure revealing a woody, pectinate crest, the

stout, recurved horns much shorter than the body.

A monotypic genus varying in flower size and color and in leaf

form.

Martynia annua L. Sp. PL 618. 1753, non L. Syst. Nat. 1113.

1759. M. diandra Glox. Obs. Bot.: 14. 1785. M. angulosa Lam.

Encycl. Meth. Bot. 2: 112. 1786. Carpoceras angulata A. Rich. Bull.

Sci. Nat. Geol. 21: 98. 1830. Disteira angulosa Raf. Fl. Telluriana:

68. 1836. Vatkea diandra 0. Hoffm. Verh. Bot. Brandenb. 73: 45.

1881; Linnaea 43: 554. 1882. Una de gato (Zacapa, Guatemala);

chiche de gata (Jutiapa); catsclaw (British Honduras); hoja de pulga

and casan pulga (Chiapas).

FIG. 40. Martynia annua. A, habit, X Yz', B, flower with corolla partly opened

to show stamens and style (stamens and staminodes opposite); X 1%; C, detail

of stigma, X 5; D, detail of anthers (dorsal view), X 4; E, three views of cap-

sule, X %.

235

236 FIELDIANA: BOTANY, VOLUME 24

Damp or dry thickets, fields, hedgerows, and clearings, sea level

to 2,400 m.; Guatemala; Huehuetenango ; Jutiapa; Pete"n; El Quiche;

Santa Rosa; Suchitepe'quez ; Zacapa. Mexico; British Honduras to

El Salvador and Nicaragua; West Indies.

Coarse, erect, often much-branched annuals to 1.5 m. tall, viscid throughout

with short or long, spreading, glandular hairs; leaves opposite, on petioles mostly

3-15 cm. long, the blades angular-ovate, angular-cordate, or broadly triangular,

subacute, cordate at the base, the margins irregularly dentate, sometimes obscurely

3-5-lobate, mostly 5-23 cm. long, 5-22 cm. broad, sparsely or densely glandular-

pubescent on both surfaces; racemes lax, mostly 10-20-flowered, the pedicels

1.5-3 cm. long, the bracts 1-1.5 cm. long, oblong-ovate to ovate, often asym-

metrical, obtuse, pubescent, ciliate; calyx about 1.5 cm. long, the upper 3 sepals

oblanceolate to elliptic, the middle one considerably longer than the 2 lateral ones,

the 2 lower sepals broader, ovate to obovate or oblanceolate; corolla 4-5.5 cm.

long, white or pinkish, with rose or purple spots on the inside of each lobe, the

spots usually surrounded by yellow; stamens included; capsule ovoid, somewhat

compressed dorsiventrally, the body of the fruit 2-3 cm. long, the stout, recurved

horns 0.5-1 cm. long.

PROBOSCIDEA Schmidel

Erect, annual or perennial herbs, sometimes suffruticose, viscid, more or less

glandular-pubescent to glandular-villous throughout, the stems stout; leaves

mostly opposite but sometimes alternate above, petiolate, the blades entire or

3-7-lobate, commonly broadly triangular to broadly ovate, sometimes suborbicular,

nearly entire or sinuate or more or less dentate; inflorescences terminal, racemose,

the flowers bracteate, pedicellate; calyx viscid-pubescent, spathelike, split ventrally

to the base, the 5 lobes shallow, the margins glandular-ciliate; corolla tube with

short, cylindrical basal portion, rather abruptly becoming campanulate and often

ventricose above, the limb somewhat bilabiate, the 5 lobes rounded, spreading;

stamens 4, didynamous, with a rudimentary fifth stamen usually obvious; ovary

unilocular; stigma bilobate; fruit capsular, the body more or less ovoid, the exocarp

viscid-pubescent, deciduous, the endocarp ligneous, deeply sculptured, crested

dorsally and sometimes ventrally, the horns about equalling or longer than the

body; seeds few-many, black.

Perhaps six or seven valid species, with only one in Guatemala.

Proboscidea triloba (Cham. & Schlecht.) Dene. Ann. Sci. Nat.

5(3) : 326. 1865. Martynia triloba Cham. & Schlecht. Linnaea 5: 121.

1830. P. botterii Dene. Ann Sci. Nat. 5(5) : 327. 1865. M. botterii

Hemsl. Biol. Cent. Am. Bot. 2: 499. 1882. P. confusa Van Eseltine,

N. Y. Agric. Exp. Sta. Tech. Bull. 149: 13, /. 4. 1929. Martynia

confusa Standl. & Steyerm. Field Mus. Bot. 23: 86. 1944.

From 1,400 to about 1,500 m.; Chimaltenango; Guatemala;

Sacatepe"quez. Mexico.

FIG. 41. Proboscidea triloba. A, habit, X KJ B, corolla opened to show sta-

mens and staminode, X 1; G, calyx opened to show pistil, X 1; D, side view of

calyx, with bracts, X 1; E, capsule opened vertically to show seeds and crest

within, X %; F, detail of leaf from lower stem, X Yt-

237

238 FIELDIANA: BOTANY, VOLUME 24

Erect, perennial herbs, sometimes suffruticose, viscid-villous throughout, the

stems stout; leaves opposite or the upper ones sometimes alternate, on petioles

mostly 3-12 cm. long, the blades mostly 3-10 cm. long, simple or shallowly 3-5-

lobate, cordate or truncate at the base, when lobate the terminal lobe acute or

obtuse, rarely acuminate, the lateral ones rounded or subacute, viscid-pubescent

on both surfaces, more densely so beneath, the margins somewhat undulate, ir-

regularly crenate, and coarsely and obscurely denticulate; racemes 4-20 cm. long,

pedicels mostly 1-3 cm. long; bracts 2, elliptic-oblong to broadly triangular,

rounded or obtuse, 0.5-1 cm. long; calyx spathelike, 1-2 cm. long, cleft to the base

ventrally, shallowly 5-lobate dorsally with the uppermost lobe longer than the

others, glandular-pubescent outside, glabrous inside, the margins glandular-ciliate ;

corolla pale yellow to brownish or purplish outside, usually white or yellowish

inside with purple and yellow markings, 3-4 cm. long, the limb spreading, the 5

lobes rounded; stamens 4, didynamous; body of capsule 4-5 cm. long, the dorsal

crest prominent, the horns curved, slightly longer than the body.

OROBANGHACEAE. Broom-rape Family.

DORTOHY NASH GIBSON

References: Guenther Beck von Mannagetta, Orobanchaceae, in

Engler, Pflanzenr. IV. 261: 1-348. 1930; John W. Thieret, The

Genera of Orobanchaceae in the Southeastern United States, Journ.

Arn. Arb. 52: 404-434. 1971.

Plants without chlorophyll, parasitic on the roots of other plants, annual or

perennial, often succulent, pubescent or glabrous; stems slender or stout, the

flowering stems arising from a thickened or swollen base; leaves reduced to scales,

alternate; inflorescences simple or branched, spicate or racemose, or the flowers

rarely solitary; flowers sessile or pedicellate in the axil of a bract, perfect; calyx

tubular or campanulate or often appearing spathelike and anteriorly cleft, the

limb with 2-5 lobes or teeth, these usually very unequal; corolla sympetalous, the

limb zygormorphic, usually bilabiate, the lobes 5, the tube curved or straight;

stamens 4, included or exserted, alternate with the corolla segments, didynamous,

inserted on the corolla tube, the filaments free; anthers dorsifixed, bithecous, free

or coherent, the thecae parallel or divergent, both or only one fertile; connective

conspicuous, sometimes apiculate; ovary superior, 2-3-4-carpellate, unilocular,

the placentae parietal; ovules numerous, anatropous; style one, the stigma usually

capitate or bilamellate; fruit capsular, loculicidally 2-3-valvate; seeds numerous,

small.

Seventeen genera, most of the species in temperate regions of

both hemispheres and most numerous in the Old World. Only the

following genus is represented in Central America.

CONOPHOLIS Wallroth

Reference: Robert R. Haynes, A Monograph of the Genus Cono-

pholis (Orobanchaeceae), Sida 4: 246-264. 1971.

GIBSON: FLORA OF GUATEMALA

239

D

FIG. 43. Conopholis alpina. A, habit, X 1A\ B, corolla opened to show sta-

mens, X2; G, calyx opened to show pistil, X 2; D, flower complete with bracteole,

X 2.

Plants parasitic on oak roots, the annual flowering stalks arising from peren-

nial swollen bases, the stems erect, commonly unbranched, grooved, glabrous to

sparsely glandular-pubescent, covered with sessile scales (reduced leaves), im-

bricate below, alternate above; inflorescences spicate, the flowers solitary, sessile

or subsessile in the axil of a bract (reduced scale-leaf), each flower subtended by

one or two small bracteoles arising from the base of the calyx; calyx tubular,

usually cleft anteriorly to below the middle and appearing spathelike, the limb

irregularly dentate or lobate; corolla sympetalous, the tube somewhat ventricose

near the base, the limb bilabiate, the upper lip arched, nearly entire or notched

or rarely shallowly 3-4-lobate, the lower lip usually trilobate, the lobes subequal,

short, usually spreading; stamens 4, exserted, the filaments inserted below the

middle of the corolla tube, the anthers bithecous, the thecae slightly divergent,

basally attenuate, glabrous or sparsely pilose; ovary superior, 4-carpellate, unilocu-

lar, the 4 placentae parietal; ovules numerous, anatropous; style one, the stigma

240 FIELDIANA: BOTANY, VOLUME 24

capitate; fruit capsular, bivalvate, when immature often partly enclosed by the

persistent corolla; seeds numerous.

Haynes recognizes two closely related species, C. americana (L.)

Wallroth, which he states is limited to the eastern United States and

Canada, and C. alpina Liebmann, of Mexico and Central America.

Conopholis alpina Liebmann, Forh. Skand. Naturf. Mode 4:

184. 1847. C. sylvatica Liebmann, I.e. 185. C. mexicana Gray ex

Watson, Proc. Am. Acad. Sci. 18 : 131. 1883. C. panamensis Woodson,

Ann. Mo. Bot. Gard. 25: 835. 1935. C. alpina var. mexicana

Haynes, Sida 3: 347. 1969.

Parasitic on roots of oak, 1,300-2,200 m.; Jalapa (Volcan de

Jumay, Steyermark 32326). Western United States; Mexico to

Panama.

Plants glabrous, the swollen basal portions sometimes 15 cm. in diameter;

stems erect, 8-28 cm. tall, to 1.2 cm. in diameter; scale leaves broadly triangular

to lanceolate, appressed and imbricate, fleshy at first, becoming coriaceous, mostly

10-20 mm. long, acute to acuminate, the veins invisible to obscure in young leaves,

in age often becoming more or less prominent; spikes densely flowered; bracts

glabrous or minutely glandular-pubescent, 10-22 mm. long, narrowly lanceolate

to narrowly triangular, acute to acuminate; bracteoles 1 or 2 or none, mostly 2-6

mm. long; calyx 4-13 mm. long, the tube more or less cylindrical, usually split

anteriorly from about half-way from the base, sometimes split to the base (rarely

not split) ; the limb irregularly dentate or lobate, the teeth or lobes acute or round-

ed; corolla yellowish white to very pale yellow, glabrous, 8-20 mm. long; stamens

exserted, the filaments pilosulous above or glabrous, the anthers glabrous or

sparsely pilose; style 6-12 mm. long, the stigma capitate; capsule broadly ovoid

to ellipsoidal, 8-16 mm. long.

Haynes distinguishes C. alpina var. mexicana (Gray ex Watson)

Haynes, as limited to the western United States and northern

Mexico, and states that it differs from the typical variety "by the

veins of its leaves and bracts being invisible, by its bracts mostly

not concealing the calyx, and by its pubescence being mostly glandu-

lar." The single Guatemalan collection of the genus, Steyermark

32326, annotated by Haynes in 1968 as C. alpina var. mexicana is a

poor specimen, lacking corollas, and all the calyces and bracts are

at least partially broken away. Veins can be seen on the lower

scale leaves.

GESNERIACEAE. Gesneria Family

DOROTHY NASH GIBSON

References: Johannes von Hanstein, Die Gesneraceen des Kon.

Herbariums und der Garten zu Berlin, Linnaea 26: 145-216, 1853;

GIBSON: FLORA OF GUATEMALA 241

27: 693-785, 1854; 29: 497-592, 1857-1858; 34: 225, 462, 1865-1866;

Gesneriaceae, in Martius, Fl. Bras. 8(1): 341-448. 1864. Anders

Oersted, Centralamericas Gesneraceer, et systematisk, plante-

geographisk Bidrag til Centralamericas Flora, 78 pp. 1858. Karl

Fritsch, Gesneriaceae, in Engler & Prantl, Pflanzenf. IV (3b) : 133-

144. 1893; 145-185. 1894; Nachtrage 1: 299-300. 1897; 2: 71. 1900;

3: 317-319. 1908; Beitrage zur Kenntnis der Gesnerioideae, in Engler

Bot. Jahrb. 50: 392-439. 1913. Paul C. Standley, Gesneriaceae, in

Trees and Shrubs of Mexico, Contr. U. S. Nat. Herb. 23: 1325-1331.

1926. Conrad V. Morton, Gesneriaceae, in Standley, Flora of Costa

Rica, Field Mus. Bot. 18: 1137-1187. 1938. Hermano Alain,

Gesneriaceae, in Leon y Alain, Flora de Cuba 4: 451-472. 1957.

Harold E. Moore, Jr., African Violets, Gloxinias, and their Relatives,

323 pp. 1957. A. J. M. Leeuwenberg, The Gesneriaceae of Guiana,

Acta Bot. Neerl. 7: 291-444. 1958.

Terrestrial or epiphytic herbs and shrubs from scaly rhizomes, stolons, or

tubers, erect, pendent, or more or less scandent, usually pubescent, rarely glabrous,

the indument commonly of multiseptate hairs; stems simple or branched; leaves

usually opposite, sometimes whorled, sometimes in basal rosettes., rarely alternate,

petiolate, those of a pair often unequal or subequal, the blades entire, serrate,

dentate, or crenate; inflorescences axillary, the often bracteate pedicels solitary,

geminate, or clustered in the upper leaf axils (when the upper leaves are reduced,

the areas of inflorescence appearing racemose), or the pedicels arising from a com-

mon peduncle and the cymose inflorescence then appearing subumbelliform;

flowers perfect; calyx free or more or less connate with the ovary, often colored,

the 5 lobes free or connate, usually valvate, often unequal or subequal, entire or

incised; corolla gamopetalous, erect, oblique, or horizontal in the calyx, the tube

often saccate or calcarate at the base, cylindric, ventricose, or ampliate above, the

limb bilabiate or almost regular, the lobes spreading or erect, often broad, entire

or denticulate or fimbriate; stamens 4 (in ours), didynamous, a staminode some-

times present, the filaments inserted near the base of the corolla, often connate;

anthers commonly quadrate to oblong, sometimes obovate, subglobose, heart-

shaped, or reniform, frequently coherent, the thecae discrete or confluent, dehiscent

by longitudinal slits for all or only part of their length; ovary superior or partly or

wholly inferior, unilocular, the 2 parietal placentae bilobate; ovules very numerous,

anatropous, on funicles that are often fleshy, borne on the inner surface of the

placentae or on both the inner and outer surfaces; style simple, elongated; stigma

usually bilobate or stomatomorphic, rarely capitate; disc annular or of distinct or

united glands, rarely absent; fruit capsular and variously dehiscent, often bivalvate,

or indehiscent and berry like; seeds numerous, minute, usually more or less fusiform.

Nearly 100 genera, widely distributed in tropical regions of the

world, with about half that number in the New World. Sixteen are

known from Guatemala. Chrysothemis friedrichsthaliana (Hanst.)

Moore (Baileya 2: 86. 1954) has been omitted from the flora. Al-

though based upon specimens supposed to have been collected in

242 FIELDIANA: BOTANY, VOLUME 24

Guatemala by Friedrichsthal, the plant has not been found later in

Guatemala, but has been collected frequently in Costa Rica and

Panama where many of Friedrichsthal's "Guatemalan" plants are

known to have been collected.

Ovary superior or nearly so.

Leaves alternate; flowers secund in lax, terminal racemes; corollas bright

blue Rhynchoglossum.

Leaves opposite, whorled, or crowded at the apex of the stem and appearing

rosulate; flowers axillary, solitary, or disposed in fascicles or cymes, these

sometimes pedunculate and subumbelliform, or the inflorescence rarely

appearing racemose as in Rechsteineria but then flowers opposite and the

racemes very leafy; corollas variously colored but not blue.

Calyx lobes shorter than the tube (in ours) ; leaves often in whorls.

Rechsteineria .

Calyx lobes longer than the tube; leaves never in whorls.

Plants acaulescent or with a very short, naked stem below a rosette-like

cluster of leaves; corollas about 12 mm. long; disc absent Napeanthus.

Plants with elongated leafy stems, the leaves never in a rosette-like cluster ;

corollas 2-8 cm. long; disc present.

Anthers with thecae confluent at the apex; disc annular or rarely semi-

annular; plants always terrestrial Besleria.

Anthers with thecae distinct; disc reduced to 1, 3, 4, or rarely 5 glands;

plants epiphytic or terrestrial.

Corollas conspicuously bilabiate, the upper lip trilobate, the middle

lobe galeiform with 2 lateral lobes spreading from it ... Columnea.

Corollas nearly regular or if bilabiate, the upper lip never galeiform.

Anthers with thecae separated by a broad connective; leaves small,

mostly 2-6 cm. long, always glabrous Codonanthe.

Anthers with thecae not separated by a broad connective; leaves

mostly 6-45 cm. long, usually more or less pubescent (rarely

glabrous).

Anthers narrowly oblong, appearing sagittate at the base, united

by their inner faces with the pores (at the base of the thecae)

uppermost until pollen is shed; corolla tube ampliate upward,

not contracted below the lobes, the limb more or less bilabiate

and somewhat spreading Drymonia.

Anthers suborbicular, not sagittate at the base, united by their

tips, thecae fully dehiscent; corolla tube usually somewhat

ventricose and more or less contracted below the lobes of the

nearly regular limb (except in A. cucullatus, when the corolla

is somewhat ampliate in the throat, with an obliquely bilabiate

limb) Alloplectus.

Ovary wholly or partly inferior.

Corollas subrotate, the tube 1-2 mm. long, much shorter than the limb.

Anthers obovate to subglobose; thecae distinct Phinaea.

Anthers oblong; thecae confluent at the apex Niphaea.

Corollas tubular to campanulate, the tube equalling or longer than the limb.

Fruiting calyx tube long-cylindric; disc absent Monopyle.

Fruiting calyx tube turbinate or hemispheric; disc present.

Inflorescence racemose; corollas 1-1.4 cm. long (in ours); disc glands

linear Diastema.

Inflorescence not racemose; corollas 1.5-6.5 cm. long; disc glands not linear.

GIBSON: FLORA OF GUATEMALA 243

Plants epiphytic and scandent (in ours); corolla pale green, spotted

with maroon Campanea.

Plants terrestrial, erect; corollas never green.

Calyx tube prolonged far above the ovary Solenophora.

Calyx tube not prolonged above the ovary.

Corolla limb broad, the lobes broad and spreading; disc annular,

usually entire and cuplike (rarely divided and more or less

bilobate in A. erecta) Achimenes.

Corolla limb narrow, the lobes short, not spreading; disc usually of

5 glands, or 2 of the glands connate, or rarely all united to form

an entire, crenate annulus, but this never cuplike. . . .Kohleria.

ACHIMENES Persoon

Perennial, terrestrial, pubescent or glabrous herbs from scaly rhizomes, the

stems simple; leaves opposite or ternate, petiolate, those of a pair subequal or

unequal, the blades usually thin, dentate or serrate; inflorescences axillary, the

pedicels solitary or geminate or one to several on a common peduncle; calyx tube

turbinate, adherent to the ovary, the 5 lobes free, entire; corolla usually salverform

funnelform, sometimes almost campanulate, the tube erect or oblique in the calyx,

often saccate or calcarate at the base, the limb almost regular or somewhat bila-

biate, the 5 lobes broad; stamens 4, didynamous, the filaments inserted near the

base of the corolla tube, the anthers united to form a square, the thecae distinct;

a small staminode often present; disc annular, often cuplike, usually entire but

rarely divided and appearing lobate, glabrous; ovary half or almost wholly inferior,

the style elongated, the stigma stomatomorphic or more or less distinctly bilobate;

fruit capsular, convex or short-conic at the apex, bivalvate; seeds small, numerous.

About 30 species, all in tropical America, with eight in Guatemala.

Corolla oblique in the calyx, the tube conspicuously saccate or calcarate at the base.

Corolla purple or red-violet, not spotted, the tube slender, scarcely ampliate

upward, calcarate at the base, the spur about half as long as the calyx lobes,

the limb commonly 3-5 cm. in diameter A. grandiflora.

Corolla orange-red, rose, yellow, or straw-colored, more or less spotted or

marked, at least in or near the throat, the tube gradually ampliate upward,

saccate at the base, the limb commonly 1.5-2.5 cm. in diameter.

Leaves mostly 5-10 cm. long; corolla finely puberulent to glabrate, the tube

orange-red or rose; stigma bilobate.

Calyx lobes oblong-ovate to broadly triangular, 3-4 mm. long; corolla

bright orange-red, the limb 1.5-2 cm. in diameter, prominently spotted

or marked with darker red, the lobes 3-7 mm. long, entire or nearly

so A. pedunculata.

Calyx lobes lanceolate to linear-lanceolate, 5-7 mm. long; corolla rose, the

limb 2-2.5 cm. in diameter, only minutely red-spotted near the throat,

the lobes 6-10 mm. long, somewhat crenate or minutely denticulate.

A. skinneri.

Leaves mostly 1.5-6 cm. long; corolla pilose with multi-septate hairs, the

tube pale yellow or straw-colored with broken maroon or reddish lines

outside; stigma stomatomorphic A. antirrhina.

Corolla erect in the calyx, the tube only a little dilated at the base, neither saccate

nor calcarate.

244 FIELDIANA: BOTANY, VOLUME 24

Corolla commonly 4.5-6.5 cm. long A. longiflora.

Corolla commonly 0.7-2.5 cm. long.

Leaves with 3-5 pairs of lateral veins; corolla commonly bright red, rose, or

pink, the tube nearly cylindric, scarcely ampliate in the throat, the limb

flat or nearly so; stigma bilobate A. erecta.

Leaves with 6-10 pairs of lateral veins; corolla white, usually with lavender,

purple, maroon, or brownish markings, the tube gradually ampliate

upward, the limb not flat; stigma stomatomorphic.

Stems densely glandular-pilosulous or glandular- villous; calyx lobes about

2 mm. wide at base; corolla tube with 2-3 pouches on underside.

A. miser a.

Stems glabrate or only sparsely villous; calyx lobes about 1 mm. wide at

base; corolla tube lacking pouches on underside A. Candida.

Achimenes antirrhina (DC.) Morton, Kew Bull. 1936: 11.

1936. Gloxinia antirrhina A. DC. in DC. Prodr. 7: 534. 1839.

Trevirana maculata Mart. & Gal. Bull. Acad. Brux. 9(2): 36. 1842.

A. foliosa Morren, Ann. Soc. Gand. 2: 403. t. 91. 1846.

Damp or wet mountain forest, often on rocks in ravines, 1,300-

3,000 m.; Huehuetenango. Mexico (Chiapas).

Slender, erect herbs 5-45 cm. tall, the stems simple, more or less pilose; leaves

opposite, those of a pair unequal or subequal, on petioles mostly 0.5-3 cm. long,

the blades ovate to lance-ovate, mostly 1.5-6 cm. long, acute to acuminate,

rounded to subcordate and usually oblique at the base, the margins serrate,

sparsely or rather densely pilose or pilosulous on both surfaces, often reddish

beneath and more densely pilose on veins and costae beneath, the lateral veins

5-8 pairs; peduncles axillary, 0.5-2.5 cm. long (rarely obsolete), the flowers solitary

on bracteate pedicels 1-2 cm. long; calyx lobes oblong-ovate to oblong-lanceolate,

5-7 mm. long, 2-3 mm. broad, pilose; corolla oblique in the calyx, 3-3.5 cm. long,

the tube pale yellow or straw-colored with broken maroon or reddish lines outside,

saccate at the base, pilose with multiseptate hairs, the limb more or less bilabiate,

the lobes yellow outside, bright red inside, the 2 upper lobes shorter, reflexed,

5-6 mm. long, the lower ones commonly 7-10 mm. long and spreading; stigma

stomatomorphic.

Achimenes Candida Lindl. Journ. Hort. Soc. London 3: 317.

1848. Diastema gracilis Regel, Flora 32: 180. 1849 (type said to

have been collected in Guatemala between Guatemala and Zacapa,

by Warscewicz) . A. gracilis Warsc. ex Regel, I.e. Dicyrta Candida

Hanst. Linnaea 27: 714, 754. 1854. Kohleria saxicola Brandegee,

Univ. Calif. Publ. Bot. 6: 66. 1914. A. saxicola Morton, Kew Bull.

1936:14. 1936.

Usually in dense wet, mountain forest, sometimes in second-

growth thickets, often on rocks, 1,200-2,500 m. (type said to have

been collected in Guatemala by Skinner} Chimaltenango; Huehue-

GIBSON: FLORA OF GUATEMALA 245

tenango; Quezaltenango ; El Quiche"; Sacatepe"quez ; Santa Rosa;

Suchitepe"quez. Mexico; Costa Rica; Panama.

Slender herbs, erect or ascending, 6-45 cm. tall, the stems simple, glabrate

or only sparsely villous, often red or purplish; leaves opposite, those of a pair

usually very unequal, on petioles mostly 0.5-4 cm. long, the blades thin, ovate to

elliptic or lance-oblong, mostly 3-8 cm. long, 1.5-5 cm. wide, acuminate or long-

acuminate, obliquely rounded or cuneate at the base, the margins serrate, sparsely

villous above, more or less villous beneath, especially on costae and veins, rarely

almost glabrous, lateral veins 6-10 pairs; inflorescence axillary, the pedicels solitary

or several on a very short, bracteate peduncle or the peduncle obsolete; calyx

pilosulous or almost glabrous, the lobes linear or linear-lanceolate, 4-6 mm. long,

about 1 mm. wide at base; corolla 15-25 mm. long, the tube white outside, some-

times marked with maroon, yellow within throat and often flecked with red or

maroon, the lobes white, the tube erect in the calyx, somewhat dilated at the base

but not saccate nor calcarate, glabrous or nearly so, the limb 12-18 mm. broad,

the lobes undulate and almost entire to erose-denticulate; filaments of stamens

glabrous; style glabrous or nearly so, the stigma stomatomorphic.

Achimenes erecta (Lam.) H. P. Fuchs, Act. Bot. Neerl. 12:

15. 1963. Columneo, erecta Lamarck, Encycl. Method. Bot. 2(1): 66.

Oct. 1786. Buchnera coccinea Scopoli, Del. Fl. et Faun. 2: 10. t. 5.

Nov. 1786. C. humilis Meerburg, PL Rar., ed. 2: 54. /. 2. 1789.

CyriUa pulchella L'Her. Stirp. Nov., Fasc. 6: 147. t. 71. 1791. A.

coccinea Pers. Syn. PL 2: 169. 1807. A. tenella A.DC. in DC. Prodr.

7: 536. 1839. A. rosea Lindl. in Edwards' Bot. Reg. 27: t. 65. 1841

(based on plants grown in England, sent from Guatemala by Hart-

weg). ?A. autumnalis Hanst. Linnaea 27: 724, 744. 1854 (believed

to have been of Guatemalan origin). A. pulchella Hitchcock, Ann.

Rept. Mo. Bot. Gard. 4: 114. 1893. Clarincillo (Huehuetenango) ;

flor de piedras, British Honduras.

Damp or wet, shaded, shrubby slopes, river bluffs, along road-

banks, and in forest, frequently in rocky places, 900-1,970 m.;

Chiquimula; Guatemala; Huehuetenango; Jalapa; El Quiche"; Solola;

Zacapa. Mexico; British Honduras; Honduras; Costa Rica; Panama;

West Indies.

Slender herbs, commonly 5-20 cm. tall, rarely to 40 cm., erect or ascending,

the stems villous or pilosulous; leaves opposite or ternate, short-petiolate, those of

a pair subequal, the blades ovate to elliptic, mostly 1-5 cm. long, acute or sub-

obtuse, acute or rounded at the base, thinly or densely villous or pilose on both

surfaces, often reddish or purplish beneath, especially on the costae and veins,

the margins serrate-dentate with only 5-10 teeth on each side, the lateral veins

3-5 pairs; pedicels axillary, solitary or geminate, 1-3 cm. long, the common

peduncle usually obsolete and the pedicels ebracteate, or rarely the bracts minute

and inconspicuous; calyx densely pilosulous, the tube 1-2 mm. long, the lobes

246 FIELDIANA: BOTANY, VOLUME 24

2-5 mm. long, linear to linear-lanceolate; corolla commonly red or rose (rarely

pink or white with red, orange, or purplish markings), puberulent or almost

glabrous, commonly 15-25 mm. long, erect in the calyx, the tube nearly cylindrical,

slightly dilated at the base but neither saccate nor calcarate, little ampliate above,

the lobes spreading, the limb usually flat or nearly so, 8-18 mm. in diameter; disc

entire or sometimes divided and bilobate; stigma shallowly bilobate.

The flowers of the form with small white corollas (7-15 mm. long),

the tube variously marked with color, superficially resemble the

flowers of A. misera Lindl. or A. Candida Lindl., especially since the

bilobate stigma is not easily seen (the stigmas of A. misera and A.

Candida are stomatomorphic) . However, the leaves of these two

species are larger, usually with 6-8 pairs of lateral veins and the

margins with 15-30 or more teeth on each side.

Achimenes grandiflora (Schiede) A. DC. in DC. Prodr. 7: 536.

1838; Hooker in Curtis' Bot. Mag. 69: t. 1+012. 1843. Trevirana

grandiflora Schiede, Linnaea 8: 247. 1833. A. grandiflora var. incisa

Hanst. Linnaea 27: 710. 1854. A. incisa Klotzsch ex Oersted,

Vidensk. Selsk. Skr. V. 5: 19. 1858. Flor de Jesus; ji-tit (Huehue-

tenango); maravilla (Guatemala).

Damp or wet, mixed forest or pine forest, often on bluffs or rocky

cliffs, 700-2,400 m.; Alta Verapaz; Chimaltenango; Chiquimula;

Guatemala ; Huehuetenango ; Jalapa ; Santa Rosa. Southern Mexico ;

Honduras; Costa Rica.

Erect herbs to 60 cm. tall, the stems simple, often red or purplish, thinly

villous; leaves opposite, those of a pair subequal, on petioles 0.5-6 cm. long, the

blades ovate to broadly lanceolate, mostly 3-12 cm. long, acuminate, usually

oblique at the base, cuneate, rounded, or subcordate, the margins coarsely serrate,

more or less short-villous on both surfaces, usually sparsely so above, paler beneath

or sometimes reddish, lateral veins 6-10 pairs; inflorescences axillary, the common

peduncle obsolete, or if present not more than 1 cm. long, the pedicels solitary,

bracteate, usually shorter than the leaves; calyx tube villous or pilose, often densely

so, the lobes linear, acute, mostly 5-12 mm. long, more or less villous or sometimes

scabrous; corolla purple or red-violet, not spotted, oblique in the calyx, 3.5-6 cm.

long, conspicuously calcarate at the base, the spur about half as long as the calyx

lobes, the tube pilose, slender, scarcely ampliate upward, abruptly expanded into

the broad limb, this commonly 3-5 cm. in diameter, the lobes spreading, entire or

crenulate; style pilosulous or puberulent; stigma bilobate.

Plants with considerably smaller corollas, the limb only 2-2.5 cm.

in diameter, are infrequently seen, but appear to conform in other

respects.

Achimenes longiflora A. DC. in DC. Prodr. 7: 536. 1839;

Lindley in Edwards' Bot. Reg. 28: t. 19. 1842; Moore, African

GIBSON: FLORA OF GUATEMALA 247

Violets, Gloxinias, and their Relatives: 107. 1957. A. kleei Paxton,

Mag. Bot. 16: 289. 1849 (described from plants cultivated in England

from material sent from Guatemala by Skinner). A. longiflora var.

alba Haage ex v. Houtte, Fl. Serres 5: t. 536. 1849. A. longiflora var.

latifolia Hanst. Linnaea 27: 706. 1856. A. longiflora var. nobilis

Hanst. I.e. A. longiflora vars. albida, ilicifolia, jauregiana, margaritae,

rosea, sessilis Hanst. op. cit. 708. A. pauciflora Oersted, Vidensk.

Selsk. Skr. V. 5: 17. 1858. Corazon de roca; maravilla celeste (Guate-

mala) ; pensamiento de monte (Huehuetenango) ; flor de pajaro ( Jalapa) .

Usually on damp or wet, steep, shaded banks and cliffs, sometimes

on rocks, in forest, often in pine forest, 250-2,600 m.; Chimaltenango,

Chiquimula; Escuintla; Guatemala; Huehuetenango; Jalapa; Jutia-

pa; El Progreso; Quezaltenango ; El Quiche"; Sacatepe"quez; Santa

Rosa; Solola; Suchitepe"quez ; Zacapa. Southern Mexico; Honduras

and El Salvador to Panama.

Erect or ascending, slender herbs, the stems simple, 10-40 cm. tall, often

reddish, villous; leaves opposite or ternate, short-petiolate, the blades ovate to

lanceolate, mostly 2-8 cm. long, acute, cuneate at the base, thinly or densely

short-villous, paler beneath or often reddish or purplish, margins serrate, the

lateral veins 3-5 pairs; flowers axillary, solitary, subsessile or on pedicels 2-10 mm.

long; calyx tube densely villous, the lobes linear-lanceolate, 7-12(15) mm. long,

long-ciliate but otherwise nearly glabrous or only sparsely pubescent; corolla

commonly purple or blue, but in some forms rose or white, or the tube white with

purple lobes, usually white in the throat, commonly 3.5-6.5 cm. long, the slender

tube erect in the calyx, neither saccate nor calcarate at the base, glabrous, not

dilated above, abruptly expanded into the flat limb, this commonly 4-6 cm. broad,

the lobes entire; style more or less puberulous; stigma bilobate.

This is probably the most common and certainly the most

attractive species found in Central America.

Achimenes misera Lindl. Journ. Hort. Soc. London 3: 319.

1848; Morton, Baileya 10: 150. 1962. Dicyrta warszewicziana Regel,

Flora 32: 181. 1849 (type a cultivated plant grown from material

sent from Guatemala by Warscewicz}. A. warszewicziana E. Otto

ex Regel, I.e. nomen, in syn. A. warszewicziana H. E. Moore,

Baileya 10: 45. 1962; The Gloxinian 12(6): 149. 1962. A. calderonii

Standley, Field Mus. Bot. 8: 150. 1930.

Damp or wet, shaded banks, often on rocks, 300-2,000 m.,

Chiquimula; Escuintla; Huehuetenango; Jalapa; Santa Rosa; Za-

capa Mexico; El Salvador; Honduras.

Slender, weak, usually erect herbs, the stems simple, mostly 10-30 cm. tall,

rather densely glandular-villous; leaves opposite, or sometimes ternate or quater-

248 FIELDIANA: BOTANY, VOLUME 24

nate, those of a pair unequal or subequal, on petioles mostly 0.3-2.5 cm. long, the

blades very thin, elliptic-oblong, ovate-elliptic, oblong-ovate, or almost lanceolate,

mostly 2.5-8 cm. long, 1-4 cm. wide, acute to long-acuminate, acute to rounded at

the base and asymmetric, the margins serrate or crenate, sparsely or densely

villosulous above, usually more densely so beneath and more or less glandular,

often densely viscid-villosulous on the veins and costae beneath, the lateral veins

mostly 6-8 pairs; flowers axillary, usually solitary, sometimes geminate, the

pedicels slender, longer than or about equalling the petioles; calyx glandular-

villosulous, the tube at anthesis little more than 1 mm. long, but 2-4 mm. long in

fruit, the lobes narrowly ovate to oblong-lanceolate, 4-7 mm. long, almost 2 mm.

wide at base, acute or acuminate, often recurving in fruit; corolla 9-14 mm. long,

the tube white with maroon or purplish spots or flushed with lavender, narrow but

gradually ampliate toward the throat, the lobes white, 2-4 mm. long; filaments of

stamens glabrous; stigma stomatomorphic.

Moore and Morton recognize A. warszewicziana as a distinct

species, based on whether the corolla tubes have two or three grooves

on the underside and on whether the color on the tubes is in spots or

in patches, but I am unable to distinguish either of these characters

on herbarium material. Of the grooves on the corolla tubes, Morton

states in part, "These characters are doubtless easily visible in fresh

flowers but are not readily seen in dried material. . ." Of the color,

he states, "The corollas of A. warszewicziana are strongly and

obviously purple-spotted on the tube, whereas those of A. misera

are essentially unspotted, at least in dried material." This statement

is followed by Moore's footnote, "The colors in living flowers are

more pronounced. The corolla of A. warszewicziana is spotted with

red or red-brown to maroon, that of A. misera is flushed with lilac

patches below the lobes on the inner surface of the tube." Morton

also stated that the leaf blades of A. warszewicziana are "strongly

resinous-glandular beneath, but those of A. misera are not, or at

least not obviously so." However, indument on leaves of specimens

annotated by both Morton and Moore varies considerably, even on

the same plant, and this character cannot be relied upon.

Achimenes pedunculata Benth. PI. Hartweg. 78. 1841 (type

from Santa Maria — perhaps Santa Maria de Je"sus, Sacatepe"quez —

Hartweg 5^6} ; Lindley in Edwards' Bot. Reg. 28: t. 31. 1842; Hooker

in Curtis' Bot. Mag. 70: t. -4077. 1844; Moore, African Violets,

Gloxinias, and their Relatives: 109. 1957. Locheria pedunculata

Regel, Flora 31: 252. 1848. Ji-lit (Huehuetenango) .

Damp or wet, shaded banks or rocks, usually in dense, mixed

forest, 1,200-2,000 m.; Chimaltenango; Guatemala; Huehuetenango;

B

FIG. 43. Achimenes pedunculata. A, habit, X 14; B, corolla opened to show

stamens and staminodes, X IK; C, calyx and pistil, X 11A.

249

250 FIELDIANA: BOTANY, VOLUME 24

Sacatepe"quez; Santa Rosa. Southern Mexico; Honduras; Costa

Rica; Panama; Colombia.

Erect herbs to nearly a meter tall, the stems pilosulous or sometimes glabrate,

often purplish or reddish; leaves opposite, those of a pair usually subequal, some-

times unequal, on petioles 0.5-4 cm. long, the blades ovate, mostly 4-14 cm. long,

abruptly acuminate, obliquely rounded or subcordate at the base, the margins

serrate, usually sparsely villous or villosulous above, paler beneath and puberulent,

the lateral veins (6)8-12 pairs; pedicels axillary and solitary or 1-several on a

common peduncle 2.5-8 cm. long, the pedicels 3-8 cm. long, bracteate, frequently

bearing small, bracteolate clusters of tiny, scale-like propagules (these also some-

times occurring in the upper leaf axils); calyx lobes oblong-ovate and abruptly

acuminate or broadly triangular, 3-4 mm. long, densely puberulent; corolla 2.5-

3.5 cm. long, the tube bright orange-red, sometimes orange or yellow on the under-

side beneath the throat, puberulous, oblique in the calyx, saccate at the base, the

sac about 1 mm. long, the tube gradually ampliate upward, yellow inside, the limb

oblique, the lobes 3-7 mm. long, orange-red, prominently lined and spotted with

darker red, rounded, entire or nearly so; stigma bilobate.

Achimenes skinneri Gordon ex Lindl. Journ. Hort. Soc.

London 2: 293, t. 4, f. 2. 1847; Moore, African Violets, Gloxinias, and

their Relatives: 110. 1957. A. hirsuta Lindl. in Edwards' Bot. Reg.

29: t. 55. 1843 (based on plants grown in England from material

sent from Guatemala by Skinner) ; Hooker in Curtis' Bot. Mag. 71 :

t. 41U- 1845, non A. hirsuta A. DC. in DC. Prodr. 7: 536. 1839.

Locheria hirsuta Regel, Flora 31: 252. 1848.

Damp or wet banks or slopes in forest, 1,000-1,700 m.; Alta

Verapaz; Baja Verapaz; Chiquimula; Sacatepequez. Honduras;

Costa Rica.

Erect or ascending herbs, the stems 15-60 cm. tall (in ours, but said to attain

almost a meter in height), more or less pilosulous or glabrate, often reddish; leaves

opposite, those of a pair usually subequal, sometimes unequal, on petioles mostly

0.5-4 cm. long, the blades broadly ovate to elliptic-ovate, mostly 3-9 cm. long,

abruptly acuminate, rounded and oblique at the base, the margins rather coarsely

serrate, sparsely or densely villous or villosulous above, paler beneath or sometimes

reddish, puberulent, the lateral veins 6-8 pairs; common peduncle obsolete (in

ours), the pedicels axillary, solitary, bracteate, pubescent, 2.5-8 cm. long, fre-

quently bearing, near the middle, small, linear, inconspicuous, often caducous

bracteoles, in the axils of which are borne clusters of tiny, scale-like propagules;

calyx lobes lanceolate to linear-lanceolate, 5-7 mm. long, densely pubescent;

corolla 3-4 cm. long, the tube rose, usually flushed with yellow beneath the throat

more or less puberulous, oblique in the calyx, saccate to very short-calcarate at

the base, the sac or spur only 1-2 mm. long, the tube gradually ampliate upward,

yellowish inside and only minutely red-dotted near the throat, the limb oblique,

the lobes clear rose to purplish rose, 6-10 mm. long, rounded, somewhat crenate or

minutely denticulate; stigma bilobate.

GIBSON: FLORA OF GUATEMALA 251

ALLOPLECTUS Martius

Reference: W. T. Steam, The Jamaican Species of Columnea and

Alloplectus (Gesneriaceae), Bull. Brit. Mus. Bot. 4: 181-236. 1969.

Shrubs or herbs, usually epiphytic, occasionally terrestrial; leaves opposite,

those of a pair equal, subequal, or unequal, petiolate, the blades entire or more or

less serrate or denticulate; inflorescences axillary, bracteate, the flowers usually

pedicellate, often fasciculate, sometimes short-racemose, rarely solitary; calyx

often colored, the 5 lobes free, entire, incised, or dentate; corolla erect or oblique

in the calyx, gibbous at the base, the tube cylindrical or more or less ventricose,

usually somewhat contracted in the throat, very rarely a little ampliate upward,

the limb narrow, nearly regular, the 5 lobes short, rounded, commonly entire;

stamens 4, didynamous, the filaments connate at the base, the anthers first united

in pairs by their tips, later free, thecae parallel, suborbicular (in ours), fully

dehiscent; disc gland large, usually solitary, sometimes smaller glands present;

ovary superior; stigma usually bilobate; fruit typically a bivalvate capsule but

sometimes berry like and indehiscent; seeds minute, shining, more or less ellipsoidal,

striate.

About 80 species, all in tropical America, with five in Guatemala.

Plants always terrestrial; calyx segments concave and cucullate at the apex;

corolla not ventricose, somewhat ampliate in the throat, the limb oblique.

Stems and lower leaf surfaces pilose A. cucullatus.

Stems and lower leaf surfaces more or less strigose A. cucullatus

var. substrigosus.

Plants either epiphytic or terrestrial; calyx segments not as above; corolla more

or less ventricose, more or less constricted in the throat, the limb nearly

regular.

Leaves of a pair very unequal, the larger ones 9-10 times longer than the

smaller ones A. viridis.

Leaves of a pair mostly subequal to equal.

Leaves densely velutinous-sericeous on both surfaces; calyx segments ovate,

entire A. calochlamys.

Leaves not velutinous-sericeous; calyx segments not ovate, the margins more

or less subulate-dentate.

Leaves with about 5 pairs of lateral veins; calyx segments spathulate-

oblong A. calotrichus.

Leaves with 7-9 pairs of lateral veins; calyx segments linear-lanceolate.

A. vinaceus.

Alloplectus calochlamys Donn.-Sm. Bot. Gaz. 27: 437. 1899.

Columnea calochlamys Morton, Bol. Soc. Venez. Cienc. Nat. 23:

78. 1962.

Often on limestone in open places or thickets, rarely epiphytic in

forest, 500-1,350 m.; Alta Verapaz (type from Sacoyoju, Tuerckheim

450).

Terrestrial or epiphytic, shrubby plants, usually erect and 30-60 cm. tall, the

stems stout, simple, densely pilose with long, appressed, white hairs; leaves of a

FIG. 44. Alloplectus calochlamys. A, habit, X K; B, calyx opened to show

corolla in early anthesis, X 1J4; G, corolla opened to show pistil and stamens, X

3; D, detail of corolla lobe, X 6.

252

GIBSON: FLORA OF GUATEMALA 253

pair subequal or unequal, the petioles stout, 0.5-3 cm. long, the blades thick,

oblong-lanceolate, mostly 6-20 cm. long, 1.5-6 cm. wide, acute, attenuate to the

base and often oblique, velutinous-sericeous on both surfaces, the margins serrate;

inflorescence axillary, bracteate, the bracts linear, mostly 1-1.5 cm. long; pedicels

1-3 in leaf axils, 0.5-4 cm. long; calyx 1.5-2 cm. long, densely sericeous, the

segments more or less ovate, usually red, obtuse or acute, entire; corolla red, but

so densely sericeous with light tan to nearly white hairs that the color may be

seen only on the tips of the lobes, the tube cylindric, slightly gibbous at the base,

appearing straight due to the dense indument but actually slightly ventricose,

commonly 2-2.5 cm. long, narrowed below the short lobes; anthers coherent in

pairs, thecae parallel, suborbicular, fully dehiscent; staminode often present; ovary

villous; disc gland solitary, bifid; fruit not seen.

Alloplectus calotrichus (Donn.-Sm.) Stearn, Bull. Brit. Mus.

Bot. 4(5): 189. 1969. Columnea calotricha Donn.-Sm. Bot. Gaz. 40:

9. 1905; Leeuwenberg, Acta Bot. Neerl. 7: 385. 1958.

Known in Guatemala only from the type, Alta Verapaz, Cubil-

guitz, 350 m., Tuerckheim 85^.2. Costa Rica; Suriname; French

Guiana.

Epiphytic or terrestrial, usually shrubby plants, more or less pilose through-

out, 10-30 cm. tall (in ours), the stems ascending, simple or branched, densely

hirsute with reddish, multiseptate hairs; leaves of a pair usually subequal (rarely

those of an occasional pair conspicuously unequal), the petioles pilose, 0.5-2.5 cm.

long, the blades oblong-elliptic, mostly 6-12 cm. long, 2-5 cm. wide, acute or

obtuse, cuneate and usually oblique at the base, pilose on both surfaces, red or

purplish beneath, the lateral veins about 5 pairs, the margins obscurely crenulate;

inflorescences axillary, the pedicels usually fasciculate, rarely solitary, 3-10 mm.

long; bracts linear or oblong, acute, 5-9 mm. long; calyx 12-20 mm. long, the

segments reddish, spathulate-oblong, acute, subequal, usually very sparsely and

irregularly subulate-dentate, more or less pilose on both surfaces, especially near

the apex; corolla orange-yellow, when young often appearing red due to the dense

red indument on the outside, 2-3 cm. long, the tube slightly ventricose, gibbous at

the base, only a little narrowed or contracted in the throat, the lobes 3-4 mm. long;

anthers coherent in pairs, the thecae parallel, suborbicular, fully dehiscent; disc

gland solitary; fruit said to be red, ovoid, about 16 mm. long.

Alloplectus cucullatus Morton, Contr. U. S. Nat. Herb. 29:

36. 1944.

Dense, wet, mixed forest, 750-2,000 m.; Alta Verapaz; El Quiche

(type from forest in the Zona Reina, Skutch 1812); Huehuetenango.

Mexico (Chiapas); Nicaragua.

Shrubs or herbs 1-3.5 m. tall, erect or ascending, usually simple, the young

stems densely villous; leaves of a pair subequal to equal, long-petiolate, the blades

elliptic or obovate-elliptic, mostly 12-30 cm. long, 5-13 cm. wide, abruptly short-

acuminate, obliquely cuneate at the base, appressed-pilose above, pilose beneath

with spreading or subappressed hairs, more densely so on costae and veins, the

FIG. 45. Alloplectus cucullatus. A, section of flowering branch, X K; B,

flower, X 1H; G, corolla opened to show stamens, X 1J^; D, calyx and pistil, X

1^2 (with insert showing large disk gland); E, detail of leaf, X %•

254

GIBSON: FLORA OF GUATEMALA 255

lateral veins 8-9 pairs, the margins finely serrate; pedicels mostly 2-5 in the leaf

axils, commonly 1-4 cm. long; bracts ovate or lance-ovate, mostly 5-8 mm. long,

caducous; calyx red, the segments unequal, 4 of them suborbicular and about 2 cm.

long, the posterior one smaller, all concave, curving outward and cucullate at the

apex, pectinate-dentate, yellow-hirsute at the middle, outside and within; corolla

red, 4-5 cm. long, densely pilose outside, oblique in the calyx, saccate at the base,

the tube cylindrical but somewhat ampliate toward the throat, the limb oblique

and shallowly bilabiate, the short lobes glabrous within; anthers coherent in pairs

by their tips, the thecae suborbicular, fully dehiscent; ovary densely hirsute;

disc gland solitary, large, bilobate; fruits not seen.

Corollas of these plants have been variously described by col-

lectors as "red," "pink," "orange-red," and "flesh-colored," perhaps

because of the heavy yellowish indument.

The corollas of A. cucullatus resemble those of Drymonia as the

tubes are neither ventricose nor constricted in the throat as in typical

Alloplectus species, and the oblique limb appears somewhat bilabiate.

However, Drymonia anthers are oblong, sagittate at the base, and

are united face to face, not by the tips as are those of this species.

A. cucullatus var. substrigosus Morton, Contr. U. S. Nat.

Herb. 29: 37. 1944.

Dense, wet forest, 1,100-1,600 m., Alta Verapaz (type from

Pansamala, Tuerckheim 733).

Differs from the typical form only in the indument of the stems

and lower leaf surfaces, which is more or less strigose rather than

pilose.

Alloplectus vinaceus (Morton) D. Gibson, Phytologia 23: 334.

1972. Columnea vinacea Morton, Contr. U. S. Nat. Herb. 29: 38.

1944.

On trees in wet, mixed forest, 1,250-3,000 m.; San Marcos;

Solola (type from southern slopes of Volcan de Atitlan, Steyermark

-47408); Suchitepe"quez.

Shrubby epiphytes, the stems stout, hirsute at the apex; leaves of a pair equal

or subequal, on petioles 0.5-2 cm. long, the blades elliptic-oblong to ovate-oblong,

mostly 7-19 cm. long, 3-9 cm. broad, acute or acuminate, cuneate at the base and

usually oblique, green and more or less pilose above, red or wine-colored beneath

(sometimes mottled or speckled with red) and hirsute with subappressed hairs,

lateral veins 7-9 pairs, the margins conspicuously serrulate; pedicels 1-2 cm. long;

bracts of inflorescence caducous, lanceolate, densely appressed-pilose, about

0.5 cm. long; calyx 10-12(15) mm. long, the segments subequal, linear-lanceolate,

acuminate, subulate-dentate, pilose; corolla dull yellow or greenish-yellow, densely

pilose outside, 3-4 cm. long, the tube very slightly saccate at base, somewhat

ventricose above, contracted in the throat, the limb almost regular, the lobes

256 FIELDIANA: BOTANY, VOLUME 24

subequal, to 5 mm. long, suborbicular, more or less undulate; anthers coherent in

pairs, thecae suborbicular, fully dehiscent; ovary densely pilose; disc glands 5, with

the 2 posterior ones connate, or 4, with the anterior one sometimes bifid or bilobate;

fruits not seen.

A collection from Quezaltenango (Roe, Roe, and Mori 723} said

to have reddish flowers, appears much like A. vinacea but the calyx

lobes are somewhat reflexed, and the plants have considerably less

indument. The three sheets of this number which I examined all

lacked corollas.

Alloplectus viridis (Morton) D. Gibson, Phytologia 23: 335.

1972. Columnea viridis Morton, Contr. U. S. Nat. Herb. 29: 38. 1944.

Usually in dense, wet, mixed forest, epiphytic or terrestrial, often

on limestone, near sea level to 2,000 m.; Alta Verapaz (type from

Sepacuite", Goll 187); Izabal; Huehuetenango. Mexico (Chiapas);

British Honduras; Honduras.

Epiphytic or terrestrial, shrubby plants to 1.5 m., simple or sparsely branched

the stems erect, densely hirsute; leaves of a pair very unequal, the larger ones, on

petioles 0.3-1.5 cm. long, the blades oblanceolate, mostly 10-25 cm. long, 3-9 cm.

wide, abruptly short-acuminate, attenuate to the oblique base, hirsute on both

surfaces, more densely so beneath, especially on costae and veins, the hairs spread-

ing, multicellular, the lateral veins 8-9 pairs, the margins inconspicuously denticu-

late, the smaller leaves subsessile, only 1.5-2.5 cm. long, lanceolate, long-attenuate,

oblique; flowers 1-3 in an axil, subsessile or very short-pedicellate; bracts ovate-

lanceolate or lanceolate, 1.5-2 cm. long, long-acuminate, subulate-dentate, green,

long-hirsute; calyx 1.5-2 cm. long, the segments subequal, linear-lanceolate, long-

acuminate, subulate-dentate, green, densely long-hirsute; corolla about 2 cm. long,

pale yellow or greenish-yellow but appearing red to orange-red due to the dense,

indument of long, dark red hairs outside, glabrous within, the tube cylindric,

ecalcarate but gibbous at the base, slightly ventricose but this hidden due to

heavy indument, a little narrowed in the throat, the limb nearly regular, the lobes

ovate, acute, about 2 mm. long; anthers coherent in pairs, thecae suborbicular,

fully dehiscent; ovary hiruste; disc gland solitary, truncate; fruits white, sparsely,

hirsute.

These plants are very similar to A. cubensis (Urban) Stearn, the

only difference apparent in specimens available to me being that the

indument on the lower leaf surfaces of A. cubensis is appressed

rather than spreading. However, our West Indian material is sparse

and none of our specimens from that area have mature corollas.

BESLERIA L.

Reference: C. V. Morton, A revision of Besleria, Contr. U.S. Nat.

Herb. 26:395-474. 1939.

GIBSON: FLORA OF GUATEMALA 257

Shrubs or perennial herbs, terrestrial (in ours), the stems terete or tetra-

gonous; leaves opposite, petiolate, subequal or unequal, the blades more or less

elliptic, acute or acuminate, membranaceous to coriaceous, dentate to subentire;

inflorescences axillary, often appearing subumbellate when a common peduncle is

present or well developed, but appearing fasciculate or cymose when the peduncle

is obsolete or inconspicuous, the flowers pedicellate, the pedicels ebracteate, solitary

or several; calyx more or less campanulate or urceolate, often brightly colored, the

5 segments imbricate, free or partially connate, usually subequal, entire or unevenly

serrulate; corolla tubular, erect, oblique, or horizontal in the calyx, sometimes

saccate or calcarate at the base, the tube usually ventricose, the limb regular or

irregular, terminal or lateral; fertile stamens 4, didynamous, a staminode sometimes

developing, the filaments broad, free, inserted on the base of the corolla tube,

usually glabrous; anthers quadrate, coherent when young, bithecous, the thecae

confluent at the apex; ovary superior, unilocular, the 2 parietal placentae ovulifer-

ous only on the inner surface; style elongated, the stigma bilobate or stomato-

morphic; disc annular or semiannular, glabrous or pubescent, often brightly

colored, entire; fruit berry like, globose, the exocarp thick, verrucose or almost

smooth; seeds numerous, minute, red, spirally striate.

About 140 species, all in tropical America, with only four in

Guatemala. Other species besides the following occur in other parts

of Central America, from Honduras to Panama.

Pedicels solitary in the leaf axils, the common peduncle present but inconspicuous,

indicated only by a node or joint; disc densely pubescent; corolla 3-3.5 cm[

long, the limb lateral B. pansamalana.

Pedicels not solitary, the inflorescence appearing fasciculate, umbelliform, or

cymose; disc glabrous; corolla 1.5-2.5 cm. long, the limb terminal.

Pedicels sessile or subsessile, the inflorescence appearing more or less fasciculate,

the common peduncle obsolete or nearly so; calyx lobes not mucronate;

corolla yellow.

Calyx lobes ovate or oblong; corolla about 1.5 cm. long, horizontal in the

calyx, the tube conspicuously saccate or short-calcarate at the base.

B. flava.

Calyx lobes orbicular; corolla about 2 cm. long, erect in the calyx, the tube

only slightly swollen at the base, neither saccate nor calcarate.

B. conspecta.

Pedicels pedunculate, the inflorescence appearing umbelliform or cymose; calyx

lobes mucronate; corolla orange or orange-red B. laxiflora.

Besleria conspecta Morton, Contr. U. S. Nat. Herb. 26: 424.

1939.

Dense, wet, mixed forest, 1,000-2,000 m.; Huehuetenango ;

Quezaltenango (type from Palmar, A. F. Skutch 11+13}; San Marcos;

Solola; Suchitepe"quez. Mexico (Chiapas).

Slender shrubs (reported also as a small tree) or suffruticose plants, commonly

1-3 m. tall, the branches densely strigose when young; leaves on petioles 1-10 cm.

long, the blades obliquely elliptic or oblong-elliptic, commonly 8-25 cm. long and

4-10 cm. wide, acuminate, acute or attenuate to the base, entire or minutely

denticulate, bearing a few scattered long hairs on the upper surface, paler beneath

FIG. 46. Besleria laxiflora. A, habit, X 1A; B, inflorescences, X 1; C, calyx

opened to show pistil, X 2; D, flower, X 2; E, corolla opened to show stamens

and staminode, X 2.

258

GIBSON: FLORA OF GUATEMALA 259

and sparsely appressed-pilose on the costae and veins, the lateral veins 9-11 (13)

pairs; inflorescence appearing fasciculate, the common peduncle obsolete or nearly

so, the few pedicels subsessile to sessile, 1-2.5 cm. long, glabrate; calyx pale green,

the lobes orbicular, about 5 mm. long, rounded at the apex, glabrous, the margins

suberose, ciliolate; corolla yellow, about 2 cm. long, erect, the tube only slightly

swollen at the base, about 4 mm. broad at the base, 8 mm. broad above, glabrous

outside, the limb terminal, about 8 mm. broad, the lobes about 2 mm. long, equal;

filaments of stamens glabrous; ovary and style glabrous; disc annular, glabrous;

fruit 4-6 mm. in diameter.

Besleria flava Morton, Contr. U. S. Nat. Herb. 26: 463. 1939.

Damp or wet, mixed forest, 850-1,150 m.; Alta Verapaz (type

from Quebradas Secas, Harry Johnson 283} collected also at Sepacuite"

and Pansamala.

Shrubs or herbs 30-60 cm. tall, possibly taller, at least sometimes branched,

the stems strigose; leaves on petioles 1.5-6 cm. long, the blades elliptic, commonly

6-18 cm. long and 3-7 cm. wide, acuminate, obliquely cuneate at the base, entire

or minutely denticulate, sparsely strigose above, paler beneath, more or less

strigose below, especially on the costae and veins, the lateral veins 9-11 pairs;

common peduncle obsolete, the pedicels sessile, few-several, 7-15 mm. long, pilose;

calyx lobes ovate to oblong, 4-5 mm. long, obtuse, membranaceous, scarious-

marginate, more or less puberulent on both surfaces, ciliolate; corolla yellow,

1.4-1.7 cm. long, conspicuously saccate or short-calcarate at the base, horizontal

in the calyx, glabrous outside, the tube ventricose, 5-7 mm. broad, the limb

terminal, the lobes about 1.5 mm. long, obtuse; filaments of stamens glabrous,

about equalling the corolla; ovary and style glabrous or nearly so; disc glabrous,

more or less divided into 2 glands.

This has been reported from Guatemala as B. glabra Hanst.

Besleria laxiflora Benth. Lond. Journ. Bot. 5: 361. 1846. B.

chiapensis Brandegee, Univ. Calif. Publ. Bot. 6: 64. 1914.

Damp or wet, usually dense, mixed forest, 35-1,300 m.; Alta

Verapaz; Huehuetenango; Izabal; Pete*n. Mexico (Chiapas) ; British

Honduras (Pueblo Vie jo) ; Honduras to Panama, along the Atlantic

coast; northern South America, southward to the Amazon.

Shrubs or suffrutescent herbs, 0.5-1.5 m. tall, sparsely branched, the young

branches densely strigose; leaves on petioles mostly 1.5-7 cm. long, the blades

elliptic to narrowly oblong, commonly 8-21 cm. long and 3-8.5 cm. wide, acuminate,

cuneate at the base, serrate, serrulate, or almost entire, paler beneath and sparsely

strigillose on the costae and veins, the lateral veins 5-10 (13) pairs; inflorescences

umbelliform or subcymose, the peduncles commonly 1.5-4.5 cm. long (rarely to

9 cm.), the flowers few, on pedicels 1-3.5 cm. long; calyx 5-14 mm. long, first pale

green or yellowish, later usually dull orange to orange-red, the lobes ovate or lanceo-

late, subcoriaceous, connate only at the base, usually minutely fimbriate, mucro-

nate; corolla orange or orange-red, erect in the calyx, not saccate at the base,

FIG. 47. Besleria pansamalana. A, habit, X 3^; B, corolla opened to show

stamens, X 1; C, stamen, X 2; D, pistil, X 2. (From Plate XVII, Botanical Ga-

zette, 1891.)

260

GIBSON: FLORA OF GUATEMALA 261

2-2.5 cm. long, slightly ventricose, glabrous outside, the limb terminal; ovary

glabrous, style puberulent; disc annular; fruits orange.

This species has a much wider range than most others in the

genus. It has been reported from Guatemala asB. mucronata Hanst.

and B. acutifolia Hanst.

Besleria pansamalana Donn.-Sm. Bot. Gaz. 16: 197. 1. 17. 1891.

Damp or wet, usually dense, mixed mountain forest, sometimes

on white sand slopes, 1,150-2,800 m.; Alta Verapaz (type from

Pansamala, Tuerckheim 196); Baja Verapaz; Chimaltenango; Que-

zaltenango; El Quiche"; Retalhuleu; San Marcos; Suchitep^quez.

Erect or ascending, usually simple, more or less succulent herbs, commonly

0.5-1 m. tall, or the main stem repent, elongating as much as 2.4 m., rooting at

the nodes and internodes, with lateral branches erect, nearly a meter tall, the

young stems densely canescent or tomentulose; leaves on petioles 1-3 cm. long,

the blades obovate-oblong or oblong-elliptic, mostly 7-20 cm. long and 4-9 cm.

wide, acute, cuneate at the base, conspicuously serrate, especially above the middle,

arachnoid-tomentulose beneath when young but soon glabrate except along the

costae and veins, the lateral veins 5-7 pairs; peduncles axillary, 2-8 cm. long, each

bearing a single pedicel 3-15 mm. long, the division between peduncle and pedicel

indicated by a node or joint; calyx parted nearly to the base, the lobes ovate,

acute, 8-13 mm. long, entire or unevenly serrulate, strigose outside, puberulent

within; corolla orange or red-orange, 3-3.5 cm. long, horizontal in the calyx, the

tube short-calcarate at the base, thinly puberulent outside, the limb lateral;

filaments glabrous; staminode often present; style bilobate; disc annular but

greatly enlarged posteriorly, densely pubescent.

A common species at many localities in the montane forests.

The plants are handsome with their large, brilliantly colored flowers

reminding one of the pouch of a Cypripedium flower.

CAMPANEA Decaisne

Herbaceous or suffruticose perennials from scaly rhizomes, usually epiphytic,

the stems usually scandent, more or less pubescent, often pendent; leaves opposite,

petiolate, those of a pair very unequal, the blades mostly oblong-lanceolate, often

asymmetric, oblique at the base, the margins crenate-serrate; peduncles usually

much elongated and commonly pendent, bearing 1-several long-pedicellate, rather

large flowers; calyx tube short, costate, pubescent, the lobes free, spreading, entire;

corolla open-campanulate, erect in the calyx, the tube ecalcarate, somewhat

ventricose, not contracted in the throat, the lobes spreading, broad, usually pale

green with conspicuous maroon spots within; filaments pilosulous, the anthers

coherent, thecae oblong, divergent, not confluent; ovary more than half inferior;

style stout, exserted, the stigma stomatomorphic; disc glands 5, glabrous; fruit

capsular, bivalvate.

262 FIELDIANA: BOTANY, VOLUME 24

The name Campanea is used instead of Capanea (Denham,

"Campanea, an Orthographic Error for Capanea (Gesneriaceae),"

Baileya 14: 37-39. 1966), as Denham's paper is based on an assump-

tion; I have been informed, in a letter from Mr. C. V. Morton, that

on the syntypes of Besleria grandiflora HBK. (the basionym of C.

grandiflora Dene.) in Paris, the name Campanea appears in the hand

of Decaisne.

About 10 species, in Central and northern South America, with

only one in Guatemala.

Campanea picturata Donn.-Sm. Bot. Gaz. 15: 28, t. 3. 1890.

Usually in wet, mixed, mountain forest, 1,450-2,500 m.; Alta

Verapaz (type from Chucaneb, J. D. Smith 1501); Chimaltenango ;

El Progreso; Quezaltenango ; San Marcos.

Usually small, herbaceous epiphytes less than 1 m. long, creeping on tree trunks

or over old logs, sometimes terrestrial and creeping, the stems, petioles, and

peduncles densely villous with long, weak, spreading, multiseptate dark hairs;

leaves on rather short, slender petioles, the blades thin, mostly oblong-lanceolate,

commonly 12-25 cm. long and 4.5-9 cm. wide, acuminate to long-acuminate, acute

or obtuse and very oblique at the base, the margins evenly crenate, abundantly

long-villous on both surfaces, somewhat paler beneath; peduncles suberect or

pendent, sometimes much exceeding the leaves, the 3-4 pedicels elongating,

commonly 2-7 cm. long; calyx lobes narrowly lanceolate, at anthesis about 2 cm.

long, long-attenuate, densely villous; corolla pale green or yellowish-green with

numerous maroon spots on the lobes within, 4-5 cm. long, abundantly villous

outside, the tube somewhat ventricose, the upper lip erect, about half as long as

the tube, bilobate, the lower lip shorter, trilobate; filaments conspicuously curved,

the anthers initially united in a disclike form.

The plants are usually abundant where they occur, but distribu-

tion is very local. All the species grow in places where there is a

continuously supply of moisture; in Guatemala, often in places where

dense fog prevails for much of the time. Dr. Standley has noted in

manuscript that if the stems are broken, the leaves collapse and

become limp almost immediately.

CODONANTHE Hanstein

Reference: Harold E. Moore, Jr., A Synopsis of the Genus Co-

donanthe, Baileya 19(1) : 4-33. 1973.

Epiphytic (in ours) or sometimes terrestrial, creeping or pendent, shrubby or

herbaceous plants growing mostly on ant nests in trees, the stems simple or

branched, glabrous or pubescent, usually rooting at some or all of the nodes;

leaves opposite, mostly short-petiolate, those of a pair equal or subequal, the

FIG. 48. Campanea picturata. A, habit, X M; B, flower with corolla opened

to show stamens and pistil, X 3^; G, detail of anthers, X 1; D, ovary dissected,

X 13/2; E, section of leaf, showing indument of lower surface, X 1; F, multiseptate

hair, X 18. (From Plate III, Botanical Gazette, 1890.)

263

264 FIELDIANA: BOTANY, VOLUME 24

blades broad, more or less fleshy, becoming coriaceous when dried, entire or serru-

late; flowers one to several in the leaf axils, on usually short pedicels; calyx seg-

ments 5 (in ours) or 2, slightly unequal, entire; corolla horizontal in the calyx, the

tube saccate or calcarate at the base, ventricose or merely ampliate upward, the

throat broad, the limb oblique, the 5 lobes rounded and spreading; stamens 4,

didynamous, included, a small staminode often present, the filaments glabrous or

pubescent, inserted below the middle of the corolla tube, connate or free at the

base, the anthers united by tips and sides, the thecae subglobose, dehiscent by

irregular pores, separated by a broad connective; ovary superior, the style in-

cluded, the stigma bilobate or stomatomorphic; disc reduced to a single posterior

gland; fruit berrylike, broadly ovoid to subglobose.

Possibly 15 species in tropical America, with only two in Guate-

mala. In addition to the two species treated here, a few plants from

other parts of Central America (Edwards P-390 from Honduras,

Bunting and Licht 1055 from Nicaragua, and Jimenez 3709, Valeria

929, and Williams et al. 284,81 from Costa Rica) were seen with

variable leaf forms, with unkeeled sepals as in C. crassifolia (Focke)

Morton, but with ventricose corollas 3-3.5 cm. long as in C. decurrens

I. M. Johnston. It may be that one or both of our species is more

variable than is known, or a third species may be involved.

Leaves mostly oblanceolate to elliptic, attenuate to the base; sepals more or less

ribbed or keeled near the base, at least one keel prominent and decurrent on

the pedicel; corollas 3-4 cm. long, the tube ventricose, the spur 4-5 mm.

long C. macradenia.

Leaves mostly ovate to lanceolate (rarely oval), the base usually rounded or acute;

sepals not keeled; corollas 1.5-2.5 cm. long, the tube strongly ampliate

upward, not ventricose, the spur about 2 mm. long C. crassifolia.

Codonanthe crassifolia (Focke) Morton, Field Mus. Bot. 18:

1159. 1938. Hypocerta crassifolia Focke, Tijdschr. Nat. Wetensch.

5: 199. 1852. Episcia hookeri Hanst. Linnaea 34: 350. 1865; Morton

in Fl. Trin. and Tob. 2(5) : 309. 1955; Leeuwenberg, Acta Bot. Neerl.

7: 381. 1958. C. confusa Sand with, Kew Bull. 492. 1931.

On earthen ant nests in trees in wet, often swampy forest, sea

level to 500 m.; Alta Verapaz; Izabal; Pete"n. British Honduras,

along the Atlantic coast to Panama; northern South America;

Trinidad.

Small epiphytic shrubs or suffrutescent plants, creeping, scandent, or sub-

scandent, often pendent, rooting at some or all of the nodes, the stems glabrous

and lustrous or minutely puberulent when young; leaves of a pair equal or nearly

so, short-petiolate, the blades coriaceous when dried, usually ovate to lanceolate,

sometimes oval, mostly 2-5 cm. long, 1-2 cm. wide, usually acute or obtuse,

sometimes short-acuminate, rounded or acute at the base, glabrous, the margins

entire; flowers 1-4 in the leaf axils, on pedicels 0.5-1.5 cm. long; calyx lobes linear-

FIG. 49. Codonanthe macradenia. A, flowering branch, X M; B, calyx opened

to show pistil with large disk gland, X 4^; C, corolla and side view of ovary with

disk gland, X 43^; D, corolla opened to show stamens, X 2^; E, detail of an-

thers, X 72/2J F, fruit opened to show seed, X 2J^ (with one seed greatly enlarged);

G, calyx with fruit, X 2K-

265

266 FIELDIANA: BOTANY, VOLUME 24

lanceolate, acuminate, puberulent, 4-9 mm. long, slightly unequal; corolla white

or yellowish, or white striped with pink, or the tube purplish and the lobes white,

puberulent or glabrous outside, 1.5-2.5 cm. long, the tube strongly ampliate

upward, not ventricose, the spur about 2 mm. long, the limb slightly irregular;

filaments of stamens glabrous; style glabrous or minutely puberulent; fruit pink

to pale purple, glabrous or nearly so, broadly ovoid to subglobose, 9-12 mm. long

and 7-9 mm. in diameter.

Codonanthe macradenia Donn.-Sm. Bot. Gaz. 25: 154. 1898.

C. decurrens I. M. Johnston, Sargentia 8: 275. 1949. Columnea calca-

rata Donn.-Sm. Bot. Gaz. 33: 254. 1902 (type from Alta Verapaz,

Tuerckheim 7645), non Codonanthe calcarata Hanst. (1865).

On earthen ant nests located in tree tops in damp or wet forest,

sea level to 350 m. ; Alta Verapaz; Izabal ; Pete"n. Mexico (Tabasco) ;

British Honduras; Nicaragua.

Small epiphytic, suffrutescent plants, creeping or scandent, rooting at the

nodes, the stems glabrous; leaves of a pair sometimes unequal, especially the lower

ones, short-petiolate, the blades carnose, succulent, paler beneath and often flushed

with pink, coriaceous when dried, oblanceolate to elliptic, mostly 3-6.5 cm. long,

1.5-3 cm. wide, short-acuminate, attenuate to the base and often decurrent on the

petiole, glabrous, the margins entire or obscurely serrulate; flowers solitary in the

upper leaf axils, short-pedicellate; calyx lobes narrowly lanceolate, acuminate,

puberulent or glabrate, unequal, 3-5 mm. long, usually more or less ribbed or

keeled near the base, at least one prominently so and decurrent on the pedicel;

corolla white, pale pink, or white striped with pink or lilac, 3-4 cm. long, glabrous

outside, the tube ventricose, the spur 4-5 mm. long; filaments of stamens pubescent;

style pubescent; fruit rose-red to purple, globose or subglobose, 10-14 mm. in

diameter.

COLUMNEA L.

Reference: W. T. Stearn, The Jamaican species of Columnea and

Alloplectus (Gesneriaceae), Bull. Brit. Mus. Bot. 4: 181-236. 1969.

Small shrubs, usually epiphytic, sometimes terrestrial, often subscandent or

pendent; leaves opposite, those of a pair subequal to conspicuously unequal,

usually short-petiolate, entire or somewhat serrate; inflorescence axillary; flowers

mostly large and showy, pedicellate, solitary or fasciculate, usually ebracteate;

calyx lobes 5, almost distinct, often colored, entire, dentate, or incised; corolla

erect in the calyx, the tube saccate at base but ecalcarate, gradually ampliate

upward, the limb conspicuously bilabiate, the upper lip trilobate, with the middle

lobe galeiform, the 2 lateral lobes spreading from it, the lower lip entire; stamens

4, didynamous, the filaments connate at the base, free above, the anthers oblong

(in ours), coherent, thecae parallel, fully dehiscent; staminode usually present;

disc gland large, posterior, usually solitary but sometimes accompanied by 2-4

smaller glands; ovary superior; stigma usually bilobate; fruit more or less globose,

fleshy and berry like; seeds minute, more or less ellipsoidal, shiny, striate.

GIBSON: FLORA OF GUATEMALA 267

A large genus, with more than 100 species, all in tropical America.

There are only five species known from Guatemala and they are never

found in the abundance that is commonplace in Costa Rica.

Leaves of a pair conspicuously unequal.

Calyx segments linear, entire; corollas yellow C. sulfurea.

Calyx segments more or less ovate, deeply laciniate-dentate; corollas red.

C. cobana.

Leaves of a pair subequal.

Leaves linear to linear-lanceolate, mostly 4-10 cm. long C. crassifolia.

Leaves neither linear nor linear-lanceolate, mostly 2-4 cm. long.

Leaf blades densely pilose on the upper surface; calyx segments ovate-oblong,

obtuse C. tuerckheimii.

Leaf blades glabrous on the upper surface; calyx segments linear-lanceolate,

acuminate C. guatemalensis.

Columnea cobana Donn.-Sm. Bot. Gaz. 57: 424. 1914, C.

venusta Standley, Field Mus. Bot. 17: 210. 1937 (type from Alta

Verapaz, Hatch & Wilson 154)-

On trees in wet forest, 900-2,500 m.; Alta Verapaz (type from

Coban, Tuerckheim 2475); El Progreso; Zacapa.

Shrubby epiphytes, erect, subscandent, sometimes pendent, the stems stout,

reddish or brownish, more or less villous with long, spreading, multiseptate,

reddish hairs; leaves of a pair very unequal, short-petiolate, the blades oblanceolate

or elliptic-lanceolate, the larger ones of each pair mostly 5-10 cm. long, the smaller

of a pair mostly 1.5-3 cm. long, acuminate to long-acuminate, usually very unequal

at the base, subobtuse or rounded on one side, cordate or auriculate on the other,

glabrous or sparsely strigose above, paler beneath and thinly strigose, often villous

with reddish hairs on the costae, veins, and margins; pedicels 2-6 cm. long, densely

villous; calyx red-villous outside, glabrous within, the segments about 2 cm. long,

more or less ovate, deeply laciniate-dentate, acuminate; corolla red, 6-7 cm. long,

pilose, the tube saccate at the base, slightly curved, the anterior lip triangular-

oblong, 1.5-2 cm. long, the galea 1.8-2.5 cm. long, about 1.5 cm. wide, broadly

rounded and subemarginate at the apex; anthers about 3 mm. long; disc gland

solitary, bifid; ovary pilose, the style sparsely villous; fruits unknown.

Columnea crassifolia Brongniart ex Lemaire, L'Hort. Univ. 6:

203, t. 7. 1844; Hooker in Curtis's Bot. Mag. 73: t. 4330. 1847. C.

stenophylla Standley, Journ. Wash. Acad. Sci. 16: 16. 1926 (type

from Chiapas). C. muenscheri Standley, Field Mus. Bot. 22: 106.

1940 (Type from Suchitepe"quez, Muenscher 124-49). Flor candelaria

romerio (Quezaltenango) .

On trees or near the bases of trees in wet forest, 500-2,500 m.;

Escuintla; Guatemala; Quezaltenango; San Marcos; Solola; Suchi-

tepe"quez. Southern Mexico.

268 FIELDIANA: BOTANY, VOLUME 24

Usually small, shrubby epiphytes, rarely terrestrial, the stems stout, reddish

or brown, when young pilose with stiff, appressed or ascending, multiseptate hairs;

leaves subsessile or very short-petiolate, those of a pair subequal, the blades

thick, linear or linear-lanceolate, mostly 4-10 cm. long and 0.5-1.5 cm. wide,

usually long-attenuate, obliquely acute at the base, glabrous above, sparsely

appressed-pilose beneath with lustrous hairs, the lateral veins inconspicuous, the

margins often revolute; pedicels solitary, 3-12 mm. long, pilose; calyx lobes

lanceolate or linear-lanceolate, 1.5-2 cm. long, about 3 mm. wide at base, long-

attenuate, entire, green or reddish, appressed-pilose; corolla bright red to orange-

red, mostly 6.5-8 cm. long, densely villous with long, slender, spreading red hairs,

the tube about 1 cm. broad in the throat, the galea 2-3 cm. long, more or less

rounded to nearly truncate at the apex, usually emarginate, the lateral lobes short,

triangular, obtuse; the lower lip triangular-oblong, about 1.5 cm. long, subacute;

anthers about 2.5 mm. long; ovary densely pilose; stigma shallowly bilobate; disc

gland solitary, broad, obscurely or shallowly bilobate; fruits not seen.

This species is similar to C. purpusii Standley (Chiapas and

Veracruz, Mexico) but C. purpusii has broader, more or less oblong

leaves to 2 cm. wide, with conspicuous lateral veins; the calyx lobes

are broader (5-7 mm. wide at base), and the hairs of the corolla tube

are shorter and usually gland-tipped.

Columnea guatemalensis Sprague, Kew Bull. 448. 1908.

On trees in dense, wet, mixed forest or thickets, 900-1,650 m.;

Alta Verapaz (type from Sarax, Tuerckheim 866}; Baja Verapaz;

Chiquimula; Huehuetenango. Mexico (Chiapas).

Sparsely branched epiphytes, the stems short or elongating and pendent from

trees, glabrous or more or less pilose; leaves of a pair subequal on short petioles

2-3 mm. long, the blades thick, subcoriaceous when dry, lanceolate to lance-oblong,

mostly 2-4 cm. long, acuminate-obtuse, obtuse or acute at the base, glabrous

above, sparsely strigose beneath, especially along the costae, or almost glabrous,

the margins entire, often revolute; flowers solitary, the pedicels commonly 5-9 mm.

long, appressed-pilose; calyx 1.5-2 cm. long, sparsely strigose or almost glabrous

outside, glabrous within, the segments linear-lanceolate, acuminate, entire, reflexed,

more or less tinged with red; corolla bright or pale red or brick red, 6-7 cm. long,

appressed-pilose outside, the anterior lobe narrowly oblong, 1.5-2 cm. long, the

galea about 2 cm. long and 1.5 cm. wide; anthers 2-3 mm. long; ovary villous;

style pubescent; fruits not seen.

This species has been confused with C. glabra (Oerst.) Hanst., of

Costa Rica, which has leaves attenuate to the base and calyx seg-

ments that are obtuse to subacute at the apex, never reflexed.

Further, the flowers of C. glabra specimens available to me for

examination often appear terminal, as they are usually in the upper-

most leaf axils, while those of C. guatemalensis are more often in the

middle or lower leaf axils.

FIG. 50. Columnea guatemalensis. A, habit, X 14; B, flower, X 11A; C,

corolla opened to show stamens, X 1>2; D, calyx with young fruit, X 11A.

269

270 FIELDIANA: BOTANY, VOLUME 24

Columnea sulfurea Donn.-Sm. Bot. Gaz. 31: 117. 1901.

On trees in wet forest, sea level to 400 m.; Alta Verapaz (type

from Cubilguitz, Tuerckheim 7646); Izabal; Pete"n. Mexico (Chia-

pas); British Honduras.

Small coarse epiphytes, the stems reddish or brown, thinly hirsute with long,

spreading, often reddish hairs; leaves of a pair unequal, short-petiolate, the blades

oblong-elliptic, elliptic, or rarely nearly obovate, the larger leaves of a pair mostly

6-11 cm. long, 2.5-4 cm. wide, the smaller leaves of the pairs mostly 2-5 cm. long,

obtuse to acuminate, narrowed to the very unequal, cuneate base, rather sparsely

pilose on both surfaces with very long and slender, spreading hairs, green above,

red or maroon beneath, the margins entire or somewhat crenate; pedicels solitary

or binate, longer than the petioles, mostly 8-16 mm. long; calyx long-pilose, the

segments green or red, linear, attenuate, 1.5-2.8 cm. long; corolla yellow, 5.5-7 cm.

long, thinly pilose with long spreading hairs, the tube slightly arcuate, 7-10 mm.

broad at the apex, gibbous at the base, the posterior lip 2-3 cm. long, the galea

1-1.5 cm. long and almost as wide, entire, the lower lip 10-12 mm. long; anthers

oblong, 2-3 mm. long; disc reduced to one bidentate gland; ovary densely pilose,

the style pubescent toward the apex; fruit white, about 12 mm. in diameter.

Columnea tuerckheimii Sprague, Kew Bull. 42. 1912. C. lutea

Donn.-Sm. Bot. Gaz. 57: 425. 1914 (type from Cubilguitz, Alta

Verapaz, 350 m., Tuerckheim 7930).

On trees in wet, mixed forest, 280-350 m.; apparently endemic

in Alta Verapaz (type from Cubilguitz, Tuerckheim 7640); also col-

lected southwest of Sibicte", Steyermark 44928.

Slender, shrubby epiphytes, the elongating, pendent branches terete, densely

fuscous-pilose with spreading hairs; leaves sessile or on petioles 1-4 mm. long,

those of a pair subequal, the blades 2-4 cm. long, elliptic-oblong, ovate-oblong,

or broadly falcate, obtuse or subacute, rounded or subcordate at the base, rather

densely hirsute on both surfaces, with 4-5 pairs of lateral veins conspicuous

beneath, the margins entire, revolute; pedicels pilose, 5-15 mm. long; calyx

segments ovate-elliptic to ovate-oblong, obtuse, narrowed at the base, 6-9 mm.

long at anthesis, 3-4 mm. broad, in fruit 10-12 mm. long, densely long-hirsute on

both surfaces, especially so within; corolla commonly deep red, 5.5-6.5 cm. long,

villous outside, the tube incurved, slightly saccate at the base, the anterior lobe

narrowly oblong or oblong-linear, 20-30 mm. long, 4-6 mm. wide, the galea

12-15 mm. long, 15-22 mm. wide, subtruncate or emarginate at the apex; filaments

of stamens glabrous, the anthers about 2 mm. long; ovary densely pilose, the style

puberulent; young fruit globose, pinkish or lavender, pilose.

Although the original description states that the calyx segments

are remotely denticulate, my examination of the type specimen as

well as other collections does not confirm this.

In his description of C. lutea, which is known only from the type

specimen with faded corollas, Donnell-Smith stated, "Corolla ex cl.

FIG. 51. Columnea sulfurea. A, habit, X M; B, flower with partly opened

corolla, X IK; C, corolla limb before anthesis, X 1/4', D, detail of anthers, X 3;

E, basal portion of corolla tube opened to show ovary, disk gland, and base of

filaments, X IK, with disk gland removed (foreground), X 1.

271

272 FIELDI AN A: BOTANY, VOLUME 24

repertore lutea . . .". The specimen differs from C. tuerckheimii in no

other respect.

These plants have been reported from Guatemala as C. microcalyx

Hanst., a Costa Rican species apparently closely related, differing in

its sparsely hirsute leaves and in its more narrow calyx lobes, 2-3 mm.

broad, that are acute to subacute at the apex and only sparsely

hirsute to nearly glabrous within.

DIASTEMA Bentham

Small, usually delicate herbs, perennial from scaly rhizomes, the stems short,

simple, more or less villous; leaves opposite, petiolate, the blades very thin, those

of a pair subequal, coarsely dentate, crenate, or serrate; inflorescences axillary or

terminal, pedunculate, racemose, few-flowered, the flowers bracteate, the bracts

opposite; calyx tube short-turbinate, the 5 lobes free, elongated, spreading, green,

entire; corolla small, white, sometimes marked with rose or lavender, tubular or

funnelform, erect in the calyx, ecalcarate, the tube not ventricose, little ampliate

upward, the limb nearly regular or obscurely bilabiate; stamens 4, and a small

staminode usually present, the filaments inserted on the base of the corolla tube;

anthers first coherent in pairs by their tips, later becoming separated, the thecae

distinct, suborbicular, dehiscent above by a slit; ovary half-inferior, the style

slender, the stigma shallowly bilobate; glands of the disc 2-5, linear; capsule

obovoid, membranaceous, convex or short-conic at the apex, bivalvate; seeds

numerous, minute.

About 40 species, all in tropical America, with only one in

Guatemala.

Diastema rupestre Brandegee, Univ. Calif. Publ. Bot. 6: 65.

1914.

Damp or wet, mixed, mountain forest, sometimes on cliffs,

700-1,500 m.; San Marcos; Suchitepe"quez. Mexico (Chiapas);

Nicaragua.

Erect herbs, commonly 10-25 cm. tall, the stems succulent, simple, sometimes

reddish, sparsely white-villous or glabrate; leaves on long, slender petioles, the

blades broadly elliptic or ovate-elliptic, mostly 5-10 cm. long, obtuse to acuminate,

rounded and oblique at the base, coarsely crenate-dentate, sparsely villous on both

surfaces with weak, multiseptate hairs; racemes mostly 4-6 cm. long, sparsely

short-villous, the 5-10 pedicels slender, usually 2-3 times longer than the calyx,

the bracts linear or linear-lanceolate, 3-4 mm. long, green, ciliate; calyx tube at

an thesis little more than 1 mm. long, in fruit about 3 mm., glabrate, the lobes

oblong-linear, green, about 3 mm. long, acute, ciliate; corolla white, 10-15 mm.

long, the tube tinged outside with lavender, glabrous outside, about 2.5 mm. broad

in the throat, the lobes rounded, 2-3 mm. long; style included, glabrous; stamens

included; glands of the disc 5, linear-clavate; capsule obovoid, bivalvate.

FIG. 52. Diastema rupestre. A, habit, X K; B, multiseptate hairs, greatly

enlarged; G, calyx opened to show pistil and linear disk glands, X 5; D, corolla

opened to show stamens and staminode, X 5; E, calyx with one lobe removed,

showing opened capsule and seeds, X 5, with two seeds greatly enlarged.

273

274 FIELDIANA: BOTANY, VOLUME 24

This species is very similar to the Costa Rican D.cristatum Hanst.,

which has larger corollas, 15-22 mm. long and about 4 mm. broad.

DRYMONIA Martius

Shrubby or suffrutescent, often scandent, epiphytic or terrestrial plants, the

stems more or less quadrangular; leaves opposite, those of a pair equal or subequal,

petiolate, the blades usually membranaceous, rarely coriaceous, and more or less

pubescent; flowers axillary, solitary or several; bracts large and foliaceous or

small and inconspicuous, often caducous; calyx segments 5, broad, free or partially

connate, green or colored, often dentate or fimbriate; corolla more or less funnel-

form, oblique to horizontal in the calyx, more or less saccate to calcarate at the

base, ampliate upward, the throat broad, the limb spreading, obliquely bilabiate,

the 5 lobes rounded, often fimbriate; stamens 4, didynamous, included, a small

staminode sometimes present, the filaments inserted on or near the base of the

corolla tube, short-connate at the base, the anthers narrowly oblong, sagittate

at the base, thecae parallel, united by their inner faces with the pores uppermost

until the pollen is shed, then the filaments coiling and the anthers separating; disc

reduced to one posterior gland; ovary superior, the style elongating, included,

stigma stomatomorphic or bilobate; fruit capsular, broadly ovoid to globose, first

fleshy, becoming coriaceous, purplish, red, or orange inside, bivalvate; seeds

obliquely ellipsoid.

Species 25 to 30, in tropical America, with six in Guatemala.

Stems, petioles, and leaf blades glabrous D. psila.

Stems, petioles, and leaf blades more or less pubescent.

Leaves commonly more than 15 cm. long, sometimes as much as 30-40 cm. long

or more.

Bracts small and inconspicuous; calyx segments deeply sinuate-dentate or

fimbriate-dentate, mucronate; corolla 2.5-3 cm. long. . . .D. macrophylla.

Bracts large and foliaceous; calyx segments entire, obtuse, not mucronate;

corolla 6-7 cm. long D. macrantha.

Leaves commonly less than 15 cm. long, mostly 4-14 cm. long.

Margins of leaf blades conspicuously sinuate-dentate; calyx segments com-

monly sinuate-denticulate (rarely merely sinuate) D. serrulata.

Margins of leaf blades entire or nearly so; calyx segments entire.

Young stems long-hirsute; lower leaf surfaces red or purple; lateral veins

4-5 (6) pairs D. oinochrophylla.

Young stems short-pilosulous; lower leaf surfaces green; lateral veins

7-8 pairs D. guatemalensis.

Drymonia guatemalensis (Morton) D. Gibson, Phytologia 23:

336. 1972. Alloplectus guatemalensis Morton, Contr. U. S. Nat.

Herb. 29: 37. 1944.

Known only from the type, San Marcos, south-facing slopes of

Volcan de Tajumulco, Steyermark 37^03, 1,400-1,700 m.

GIBSON: FLORA OF GUATEMALA 275

Shrubby epiphyte, the stems simple or sparsely branched, densely short-

pilosulous when young; leaves on petioles 1-3.5 cm. long, the blades broadly

oblanceolate, those of a pair more or less unequal, mostly 6-15 cm. long, 3-5 cm.

broad, short-acuminate, obliquely cuneate at the base, entire or nearly so, sparsely

and minutely strigillose on both surfaces, the lateral veins 7-8 pairs; bracts

caducous; pedicels 1-3 in the leaf axils, shorter than the calyx, densely short-

pilosulous; calyx pale green and more or less suffused with rose-red, sparsely

strigillose, the segments subequal, ovate to lance-ovate, erect, 1.5-2.5 cm. long,

rather abruptly long-attenuate, subcordate at base, entire; corolla creamy- white,

3-4 cm. long, sparsely short-pilose or almost glabrous outside, oblique in the calyx,

the tube saccate at the base, ampliate above, the limb about 2 cm. broad, the lobes

subequal, broad, suberose and somewhat undulate; anthers oblong, about 5 mm.

long, sagittate at the base, thecae parallel, dehiscent only at the basal end; ovary

densely tomentose; stigma bilobate; disc gland solitary; fruits unknown.

Drymonia macrantha (Donn.-Sm.) D. Gibson, Phytologia 23:

336. 1972. Alloplectus macranthus Donn.-Sm. Bot. Gaz. 31: 117.

1901.

Damp, mixed or pine forest, 350-1,400 m.; Alta Verapaz (type

from Cubilgiiitz, Tuerckheim 764.2) ; Solola.

Small, epiphytic or terrestrial shrubs with thick branches, minutely strigillose

on the stems, leaves, and inflorescences; leaves short-petiolate, the blades thin,

oblong-elliptic to broadly ovate-elliptic, very large, mostly 20-45 cm. long and

10-20 cm. wide, acute or short-acuminate, cuneate at the base and decurrent on the

petiole, the lateral veins 7-10 pairs, the margins subentire or crenate-serrate;

flowers often numerous, the pedicels clustered in the upper leaf axils; bracts large

and foliaceous; calyx first green, becoming yellowish or orange, 2-3.5 cm. long,

the segments short-connate, subequal, oblong, obtuse, entire; corolla white outside,

yellow within, puberulent outside, 6-7 cm. long, the tube gradually ampliate, the

limb 3-4 cm. broad, the lobes unequal, rounded, more or less fimbriate or the

anterior one fimbriate and the others nearly entire; anthers oblong, appearing

sagittate at the base, about 5 mm. long, dehiscent basally ; ovary densely pubescent;

stigma stomatomorphic; fruit broadly ovoid or subglobose, about 17 mm. long;

seeds red, about 1 mm. long.

Drymonia macrophylla (Oerst.) H. E. Moore, Baileya 3: 109,

/. 1+3. 1955. Caloplectus macrophyllus Oerst. Centralam. Gesner.:45.

1858. Drymonia mucronulosa Hanst. Linnaea 34: 363. 1865- 66 (type

specimen in Herb. Wendland said to have been collected in Guate-

mala by "Sell"). Alloplectus macrophyllus Hemsl. Biol. Centr. Am.

2 : 484. 1882. D. ocroleuca Standl. Field Mus. Bot. 12 : 413. 1936 (type

from British Honduras, Schipp S-901), non Poepp. & Endl. 1845.

D, belizensis Standl. Trop. Woods 52: 29. 1937.

On trees in damp or wet forest, 50-400 m.; Alta Verapaz; Izabal;

El Quiche". British Honduras; Costa Rica.

FIG. 53. Drymonia oinchrophylla. A, habit, X H» B» calyx opened, with

corolla partly removed to show pistil and disk gland, X IK; C, ovary and disk

gland, X 3; D, corolla opened to show stamens, X IK; E, dorsal view of anther,

X 3K; F, anthers after pollen has been shed, X 3.

276

GIBSON: FLORA OF GUATEMALA 277

Shrubby or suffrutescent epiphytes, the stems thick, puberulent or glabrate;

leaves short-petiolate, those of a pair subequal or unequal, the blades elliptic,

oblong-ovate, or oblanceolate, sometimes falcate, very shortly acuminate or acute,

cuneate or obtuse at the base, more or less strigose-puberulent on both surfaces,

more densely so beneath on costae and veins, mostly 15-35 cm. long, 5-14 cm. wide,

lateral veins 5-7(8) pairs, the margins denticulate or crenate; bracts small and

inconspicuous, lanceolate, acuminate, sometimes abruptly so; pedicels 1-3,

pubescent, shorter than the calyx; calyx segments thin, foliaceous, papery when

dried, puberulent to pilosulous, subequal, 10-15 mm. long, broadly ovate to ovate-

oblong, mucronate, the mucro 2-4 mm. long, the margins fimbriate-dentate;

corolla cream-colored or pale yellow with lavender or maroon markings, 2.5-3 cm.

long, the tube short-calcerate at the base, gradually ampliate upward, densely

puberulent outside, the lobes rounded, at least the anterior ones minutely fimbriate;

anthers 3-4 mm. long, sagittate at base, dehiscent at basal end only; style minutely

pubescent; stigma stomatomorphic; disc gland solitary; fruits not seen.

Drymonia oinochrophylla (Donn.-Sm.) D. Gibson, Phytologia

23: 336. 1972. Alloplectus oinochrophyllus Donn.-Sm. Bot. Gaz. 54:

239. 1912.

On trees in wet forest, 750-1,700 m.; Alta Verapaz (type from

Pansamala, Tuerckheim 1080); Huehuetenango.

Usually shrubby epiphytes, often branched, usually scandent, the young

stems villous-hirsute with long, spreading, multiseptate reddish hairs; leaves on

hirsute petioles mostly 0.5-2 cm. long, the blades subcoriaceous, those of a pair

subequal to unequal, ovate to ovate-elliptic or oblong-elliptic, mostly 5-12 cm.

long and 2.5-5 cm. wide, abruptly short-acuminate, obliquely cuneate at the base,

entire or nearly so, sparsely strigose to glabrate on the upper surface, red-purple

beneath and often villous along the costae, the lateral veins 4-5 (6) pairs; pedicels

1-3 in the leaf axils, pilose, 0.5-1 cm. long; bracts lanceolate, reddish, 0.5-1 cm.

long; calyx green or more or less suffused with red, sparsely villous, the segments

subequal, 2.5-3 cm. long, lance-ovate, long-attenuate, subcordate to cordate at the

base, entire; corolla white or cream-colored, 3.5-4 cm. long, sparsely pilose outside,

oblique in the calyx, the tube saccate at the base, ampliate above, the limb oblique,

the lobes suborbicular, about 1 cm. long, more or less crenulate; anthers oblong,

sagittate at the base, about 5 mm. long, the thecae parallel, dehiscent at the basal

end only; ovary densely tomentose; stigma bilobate; disc gland solitary, obscurely

bilobate; fruits unknown.

Drymonia psila D. Gibson, Phytologia 23: 335. 1972.

Damp forest, 1,200-1,300 m., Alta Verapaz; Izabal (type from

Cerro San Gil, Steyermark 41980).

Shrubby, sometimes scandent epiphytes, glabrous throughout, the stems often

red or purplish; leaves on petioles 1-3.5 cm. long, the blades coriaceous, broadly

elliptic to oblanceolate, mostly 9-25 cm. long, short-acuminate, cuneate and often

oblique at the base, glabrous on both surfaces, paler beneath, margins entire,

lateral veins 5-7 pairs; pedicels 1-3; bracts small, linear-lanceolate, caducous;

FIG. 54. Drymonia psila. A, leafy stem, X K; B, two views of one calyx,

the upper one opened to show pistil with disk gland, X 2; C, corolla opened to

show stamens (left) and view of base of corolla tube (right), X 2; D, detail of an-

thers, X 3; E, ovary with disk gland, X 3.

278

GIBSON: FLORA OF GUATEMALA 279

calyx glabrous, the 5 segments unequal, one considerably smaller than the others,

oblong-ovate to lanceolate, acute, accrescent, 1-2 cm. Jong; corolla white or cream-

colored, sometimes marked with purple, glabrous, 3-3.5 cm. long, the tube ampliate

above, the limb oblique, the lobes more or less rounded, crenate, the anterior one

more or less fimbriate; anthers about 3 mm. long, appearing sagittate at the base,

united face to face; ovary glabrous; disc gland solitary; stigma stomatomorphic;

fruits unknown.

Drymonia serrulata (Jacq.) Mart, ex DC. Prodr. 7: 543. 1839.

Besleria serrulata Jacq. Hort. Schonbr. 3: 21, t. 290. 1798. Besleria

spectabilis HBK. Nova Gen. & Sp. 2: 400. 1817. Drymonia specta-

bilis Mart, ex DC. Prodr. 7: 543. 1839. D. bicolor Lindl. Bot. Reg.

24: t. 4. 1838. D. jacquinii G. Don, Gen. Syst. 4: 653. 1838. D.

chiapensis Brandegee, Univ. Calif. Pub. Bot. 6: 64. 1914.

On trees in wet forest, 300-1,100 m.; Alta Verapaz; Pete"n; Santa

Rosa; Sol old; Suchitepe"quez. Southern Mexico; British Honduras

to Panama; Colombia.

Woody or suffrutescent, often scandent epiphytes, the stems simple or branch-

ing, reddish, sparsely strigose, puberulent toward the apex; leaves petiolate, those

of a pair equal or subequal, the blades usually oblong to elliptic, sometimes obovate,

mostly 8-12 (15) cm. long, 3-7 cm. wide, acuminate to subacute, cuneate to

rounded at the base, sparsely strigillose on both surfaces or puberulent beneath,

the margins conspicuously sinuate-dentate, the lateral veins 6-8 pairs; pedicels

solitary or 3 together, pubescent or puberulous, mostly 1-1.5 cm. long; calyx

segments ovate to lanceolate, 2-4(5) cm. long, accrescent, commonly sinuate-

denticulate, rarely merely sinuate, acuminate, rounded or subcordate at the base,

sparsely strigillose or puberulent; corolla finely puberulent outside, yellowish

white or sometimes flushed or variously marked with pink, lavender, or maroon,

3.5-5 cm. long, the tube ampliate above, the lobes rounded, all more or less erose-

fimbriate or the larger anterior one conspicuously so and the dorsal lobes sometimes

nearly entire; anthers oblong, 4-6 mm. long, sagittate at the base, united face to

face, dehiscent only at the base; stigma bilobate; capsule globose to broadly ovoid,

puberulous; seeds very numerous, fusiform, shining, about 1 mm. long.

Called "viejito" in Chiapas.

KOHLERIA Regel

Reference: C. V. Morton, The Genus Kohleria in Mexico [Gesneri-

aceae], Baileya 15: 61-78. 1967.

Terrestrial herbs, shrubs, or subshrubs, abundantly pubescent with usually

multiseptate hairs; leaves opposite or whorled, petiolate, those of a pair subequal

or unequal, the blades usually thick, the margins serrate or crenate; inflorescences

axillary, the flowers pedicellate, solitary or several in the axils of reduced leaves

and the inflorescence then appearing terminal and often racemiform, or the pedicels

fasciculate or 2-4 or more on a common peduncle, the pedicels then subtended by

FIG. 55. Drymonia serrulata. A, habit, X 1A; B, corolla, natural position,

X 1A", C, corolla dissected to show stamens, X 1; D, calyx and pistil, X 1.

280

GIBSON: FLORA OF GUATEMALA 281

a pair of bracts and the inflorescence subumbellate; calyx tube more or less turbi-

nate, the 5 lobes free, about equalling or considerably longer than the tube,

pubescent; corolla tubular, erect or somewhat oblique in the calyx, usually ven-

tricose near the middle toward the throat, hirsute with usually reddish hairs, the

limb narrow, more or less bilabiate or nearly regular, the 5 lobes short, often vari-

ously spotted or striped within; fertile stamens 4, the filaments inserted on the base

of the corolla tube, a small staminode present; anthers initially coherent, later

coherent or free, the thecae distinct, fully dehiscent; disc glands usually 5, distinct

or not, sometimes the 2 posterior ones connate, or all of them united to form a

thin, entire, crenate, or very shallowly lobate annulus ; ovary more than two-thirds

inferior, the placentae ovuliferous on both surfaces; style pubescent or pilosulous,

elongating after the anthers have shed their pollen, the stigma stomatomorphic or

bilobate; fruit capsular, conic or rostrate at the apex, bivalvate; seeds small,

very numerous.

Sixty or more species, in tropical America, with eight in Gua-

temala.

Leaves in area of inflorescence much reduced, often bractlike; flowers axillary but

the total inflorescence appearing terminal and often racemiform; common

peduncle usually obsolete.

Leaves often in whorls of 3 or 4; pedicels commonly 2-5 in leaf axils; corolla

lobes red throughout or only 2-3 lobes spotted; disc glands free or the 2

posterior ones connate.

Indument of pedicels and calyx reddish or brownish; calyx lobes ovate to

oblong-ovate, often recurving, 3-5 mm. long; corolla with 2-3 lobes more

or less spotted K. spicata.

Indument of pedicels and calyx white or whitish; calyx lobes triangular, erect,

about 2 mm. long; all corolla lobes unspotted K. longifolia.

Leaves opposite, not in whorls; pedicels always solitary in leaf axils; corolla lobes

all conspicuously spotted; disc glands united to the middle to form a

ring K. lanata.

Leaves in area of inflorescence little if at all reduced; inflorescences conspicuously

axillary and often appearing umbelliform, the pedicels 1-4 on a common

peduncle.

Pedicels usually solitary, the peduncle marked only by the presence of bracts

at the base of the pedicel.

Leaf blades glabrous or more or less pilose above; common peduncle 1.5-3.5

cm. long; indument of lower leaf surfaces, pedicels and calyces appressed.

K. fruticosa.

Leaf blades scabrous above, often rugose to bullate; common peduncle

0.2-1 cm. long; indument of lower leaf surfaces, pedicels and calyces

spreading K. rupicola.

Pedicels 2-4 on a conspicuous common peduncle.

Leaf blades mostly 12-28 cm. long; bracts subtending the pedicels obovate,

leaflike; calyx lobes broadly ovate to obovate K. skutchii.

Leaf blades mostly 6-15 cm. long; bracts subtending the pedicels linear or

very narrowly oblanceolate; calyx lobes not as above.

Calyx lobes 2-6 mm. long, broadly or narrowly triangular and acute, or

sometimes narrowing abruptly to a subulate apex; corolla commonly

2-2.5 cm. long K. deppeana.

Calyx lobes 6-10 mm. long, linear to linear-lanceolate, attenuate or narrow-

ing gradually to a subulate apex; corolla commonly 3-4 cm. long.

282 FIELDIANA: BOTANY, VOLUME 24

Indument of inflorescence spreading; corolla 3.5-4 cm. long. .K. elegans.

Indument of inflorescence appressed; corolla about 3 cm. long.

K. elegans var. pedunculata.

Kohleria deppeana (Schlecht. & Cham.) Fritsch in Engler &

Prantl, Pflanzenf. IV. 3b: 174. 1894. Gesneria deppeana Schlecht. &

Cham. Linnaea 5: 110. 1830. (?) Columnea umbellata Bertol. Fl.

Guat. 426. 1840 (type from San Lucar, i.e. San Lucas, probably in

Sacatepe"quez, Velasquez). Moussonia costaricensis Klotzsch ex Oer-

sted, Centralamer. Gesner. 33. 1858; Hanstein, Linnaea 34: 286.

1865. Isoloma deppeanum Hemsl. Biol. Centr. Amer. Bot. 2: 478.

1882. Gesneria lasiantha Zucc. Abh. Math. Phys. Kl. Acad. Wiss. 1:

300. 1832. K. deppeana var. lasiantha Fritsch, Bot. Jahrb. 50: 425.

1913. Chompipe, Susana sylvestre (Escuintla) , khos-corrusumal, sel-

dia (Guatemala); kak-rok-kiche (Chichipin, San Marcos).

Damp or wet thickets, mixed forest or pine forest, sometimes on

clay bluffs or sandy banks near rivers and streams, rarely in marshes,

usually 1,200-2,700 m., but sometimes as low as 500 m.; Alta

Verapaz; Chimaltenango; Chiquimula; Escuintla; Guatemala; Hue-

huetenango; El Progreso; Quezaltenango ; El Quiche"; Sacatepe"quez ;

San Marcos. Mexico; Honduras; Costa Rica.

Usually erect shrubs or suffruticose herbs to 2 m. tall, the stems sometimes

weak and reclining on other plants or pendent from banks or cliffs, the branches

densely villous-tomentose with spreading or subappressed, often reddish hairs;

leaves on usually short, rarely somewhat elongated petioles, the blades of a pair

more or less unequal, oblong-lanceolate to oblong-ovate, mostly 5-17 cm. long,

acuminate, obtuse or acute at the base and often oblique, the margins crenate or

serrate, densely and softly pilose to velutinous on both surfaces with spreading or

appressed, multiseptate hairs, paler beneath, the costae and veins often reddish

to maroon; peduncles mostly 3-8 cm. long, villous, 3-4-flowered, the flowers on

villous or pilosulous pedicels 1-3 cm. long, the pedicels subtended by a pair of

linear or narrowly oblanceolate bracts; calyx densely short-villous or pilosulous

with spreading or appressed hairs, the hypanthium turbinate, 2-3 mm. long at

anthesis, the lobes commonly broadly or narrowly triangular and acute, sometimes

recurved 2-4 mm. long, but frequently becoming abruptly subulate at the apex,

then 3-6 mm. long; corolla 2-2.5 cm. long (rarely to 3 cm.), bright red or orange-

red outside and densely pilosulous with reddish hairs, yellow inside, the lobes

marked with red within; stamens short-exserted ; style pubescent; capsule 6-9 mm.

long, densely pubescent.

A highly variable, widely distributed, and often misunderstood

species, frequently confused in herbaria with K. elegans (Dene.) Loes.

which has linear to linear-lanceolate calyx lobes and a larger corolla

(3-4 cm. long).

FIG. 56. Kohleria deppeana. A, habit, X l/i; B, flower at anthesis, X 2; C,

detail of corolla lobes, X 234; D, corolla opened to show stamens and staminode,

X 2; E, detail of anthers (initially coherent), X 2; F, calyx and style, with sepa-

rate view of top of the almost wholly inferior ovary, X 2; G, calyx enclosing cap-

sule, X 3 (with two multiseptate hairs, greatly enlarged); H, calyx and capsule

opened centrally (ovary is unilocular), X 3.

283

284 FIELDIANA: BOTANY, VOLUME 24

Specimens of Tuerckheim 10, cited by Fritsch (1913) as K. dep-

peana var. lasiantha, differ from the typical form only in their

appressed indument, but Skutch 1857, annotated by Morton as var.

lasiantha, has calyx lobes subulate at the apex and with relatively

wide sinuses. Numerous collections (i.e., Standley 59727, 63019, and

64643, Molina, Burger, & Wallental6184, Steyermark 33282, Williams,

Molina, & Williams 25141, Heyde &Lux 4443, and Salas 1401) with

very short, whitish hairs on pedicels and calyces, have calyx lobes

similar to Skutch 1857, certainly more obviously subulate than

triangular, usually 3-6 mm. long, and appear to fit the description

of Moussonia costaricensis Klotsch ex Oersted. Fritsch placed this

species in synonomy with K. elegans, explaining that after comparing

the Berlin specimen of M. costaricensis with the plate of M. elegans,

he felt justified in uniting them, although he was aware of their

differences in indument color and length. His discussion does not

mention differences in length of calyx lobes. Morton (1967) listed M.

costaricensis as a possible synonym of K. elegans, indicating his un-

certainty by his use of a question mark and his statement that he had

not seen the former. Hanstein (1865) recognized M. costaricensis as a

separate species and placed it immediately after M. deppeana, with

the remark, "Omino praecedenti similis," distinguishing it from M.

deppeana by its whitish hairs, by its calyx lobes a little longer than

the tube, and by its narrower corollas. He also described the corollas

of both M. deppeana and M. costaricensis as about 1 in. (2.5 cm.)

long, and accepted Decaisne's description of the corollas of M. elegans

as "sesquipollicaris" (3.7 cm. long). Although I have not seen the

Berlin specimen of M. costaricensis, it seems to me that Hanstein

was perhaps closer to the truth than Fritsch, that M. costaricensis is

more closely related to K. deppeana than to K. elegans and may

indeed be synonymous with the var. lasiantha. It is also quite

possible that all three may be parts of an aggressive complex.

It is interesting that one collection, Heyde & Lux 6210, cited by

Fritsch as K. elegans, may have been a mixed collection. Two

specimens of that number, F.M. 265011 and U.S. 1336322, have the

small calyx lobes, 2-4 mm. long, typical of K. deppeana, but a third

sheet, U.S. 258885, consisting of three specimens, is mixed. The

center and right-hand specimens have calyx lobes only 2-3 mm. long,

but the specimen on the left has flower buds with calyx lobes 6, 7,

and 8 mm. long, as in K. elegans. The other Guatemalan collection

cited by Fritsch as K. elegans, J. D. Smith 1867, has short white

GIBSON: FLORA OF GUATEMALA 285

indument, longer calyx lobes, 3-5 mm. long, and corollas only 2.5

cm. long, matching the description of M. costaricensis, and I have

therefore placed it in K. deppeana.

Hanstein, Fritsch, and Morton all placed "Gesneria elongata Mart.

& Gal. (Bull. Acad. Brux. 9(2) : 32. 1842), non HBK." in synonomy

with K. deppeana. It is omitted here because Martens and Galeotti

(I.e.: 33) pointed out that the calyx lobes of the specimen they

considered to be G. deppeana, Galeotti 1906}, were linear-lanceolate,

5-6 lines long (11.5-13.8 mm.) and quite different from the ovate

lobes, only 2 lines long, of the specimens they determined to be

G. elongata "H.B. et. Dec.," Galeotti 1903 and 1918.

Kohleria elegans (Dene.) Loesener, Bull. Herb. Boiss. 7: 574.

1899. Moussonia elegans Dene, ex Planch. Fl. Serres 5: t. 489. 1849

(type cultivated in Europe from material said to have been originally

received from Guatemala). Isoloma elegans Hemsl. Biol. Centr.

Amer. Bot. 2: 478. 1882. K. collina Brandegee, Univ. Calif. Publ.

Bot. 6: 66. 1914 (type from Cerro del Boqueron, Chiapas, Mexico).

Damp or wet thickets or forest, often on slopes and cliffs, 1,000-

2,300 m.; Alta Verapaz; Baja Verapaz; El Quiche"; San Marcos.

Western and southern Mexico; Costa Rica (?).

Shrubs or suffruticose herbs 1-3 m. tall, the stems and branches densely

villous with short, spreading, brownish or reddish to dark red, multiseptate hairs;

leaves short-petiolate, the blades of a pair unequal to subequal, ovate to ovate-

oblong or elliptic, mostly 5-15 cm. long, acuminate to long-acuminate, usually

rounded to subcordate at the base, rarely acute, densely villous, especially beneath,

with spreading, multiseptate hairs, the margins serrate or crenate; peduncles

axillary, 3-10 cm. long, the pedicels 1-4, mostly 1.5-6 cm. long, villous with usually

reddish to purplish (rarely whitish) hairs about 1 mm. long, subtended by a pair

of linear bracts; calyx densely villous with spreading, usually reddish hairs, the

tube turbinate, the lobes usually linear or linear-lanceolate, rarely lanceolate,

attenuate or gradually narrowing to a subulate apex, 6-10 mm. long at anthesis,

sometimes longer in fruit, twice as long as the tube or longer; corolla usually deep

red or bright red, sometimes orange-red, commonly 3-4 cm. long (in ours), about

1 cm. broad in the throat, usually densely red-villous outside, the lobes more or

less rounded with margins shallowly and unevenly denticulate, inconspicuously

spotted or striped within; stamens short-exserted; style pubescent, the stigma

stomatomorphic; capsule about 1.5 cm. long.

This species appears to have been long misunderstood and rather

widely confused with K. deppeana (Schlecht. & Cham.) Fritsch, and

the two may represent extremes of an aggressive complex. The calyx

lobes of the typical form of K. deppeana are commonly triangular

and only 2-4 mm. long; these sometimes become abruptly subulate

at the apex and are then 3-6 mm. long. The corollas of K. deppeana

286 FIELDIANA: BOTANY, VOLUME 24

are 2-2.5 cm. long. The illustration of Moussonia elegans Dene, is

of a plant with linear calyx lobes apparently 6-10 mm. long, and the

corolla is described as "sesquipollicaris" (3.7 cm. long). While some

corollas of several collections of Mexican plants determined by

Morton to be K. elegans are only 2.5 cm. long, mature corollas on all

of the Guatemalan specimens are 3-4 cm. long.

Kohleria elegans var. pedunculata (Brandegee) Morton,

Baileya 15: 75. 1967. K. pedunculata Brandegee, Univ. Calif. Publ.

Bot. 6: 67. 1914 (type from Cerro del Boqueron, Chiapas, Mexico,

Pur pus 6664-

Damp or wet thickets or forest, 2,000-2,500 m.; Quezaltenango.

Chiapas, Mexico.

Differs from K. elegans in its appressed indument and shorter

corollas, 2.8-3 cm. long. One isotype, F.M. 415527, is a mixed

collection; the specimen on the right side of the sheet is part of the

type collection, but the specimen on the left is K. fruticosa

Brandegee.

Kohleria fruticosa Brandegee, Univ. Calif. Publ. Bot. 6: 67.

1914 (type from Cerro del Boqueron, Chiapas, Mexico, Purpus

7007).

Damp thickets and forest, 2,300-2,800 m.; Quezaltenango; San

Marcos. Mexico.

Branching shrubs 1-2 m. tall, the stems villous with appressed or ascending

hairs; leaves opposite, short-petiolate, those of a pair subequal to unequal, the

blades oblong to oblong-lanceolate or sometimes obovate, mostly 5-8 cm. long,

acuminate, the base nearly rounded to cuneate, glabrous or more or less pilose

above, appressed-villous below, especially so on costae and veins, the margins

serrate; inflorescences axillary, the peduncles slender, 1.5-3.5 cm. long; pedicels

solitary, 1.5-2.5 cm. long, subtended by a pair of linear bracts 3-5 mm. long;

calyx lobes narrowly linear-lanceolate, 10-14 mm. long, tapering to an almost

filiform apex, more or less appressed-pilose; corolla red, about 3 cm. long, the tube

gradually ampliate, finely appressed-pilose outside or nearly glabrous, the lobes

more or less minutely pectinate-denticulate; disc glands indistinct; ovary sparsely

pubescent; style glabrous or nearly so, the stigma stomatomorphic; capsule

pubescent.

Mature leaves of the Mexican plants are glabrous above with

the young leaves pilose, while all leaves of the Guatemalan plants

are more or less pilose.

GIBSON: FLORA OF GUATEMALA 287

Kohleria lanata Lemaire, Illustr. Hortic. 8: L 287. 1861; Moore,

Gentes Herb. 8 : 379. 1954. Gesneria maculata Mocino & Sesse" ex DC.

Prodr. 7: 532. 1838, non Martius (1829).

Not reported from Guatemala but may be expected there.

Mexico (Chiapas and Guerrero), 850-1,350 m.

Herbs, commonly 10-40 cm. tall, villous to pilose throughout, the stems usually

densely villous with whitish to pale yellowish hairs; leaves opposite, those of a

pair subequal, on petioles 0.5-5 cm. long, the blades ovate to oblong-ovate,

acuminate, mostly 4-13 cm. long and 3-9 cm. wide, the margins crenate to serrate,

sparsely or densely pilose above, more or less villous beneath, often densely so and

appearing velvety; pedicels solitary in the axils of reduced leaves, 0.5-2.5 cm. long,

densely villous, the common peduncle obsolete; calyx densely villous, the lobes

oblong, acute or obtuse, mostly 6-9 mm. long, broadly spreading; corolla red to

orange-red, normally about 3 cm. long (sometimes only 2 cm.), the tube densely

villous with usually light (sometimes reddish) hairs, the 5 lobes spreading, con-

spicuously purple-spotted within; disc with 5 glands united to the middle into a

ring; ovary villous, the style villous to pubescent, the stigma bilobate; fruits not

seen.

Kohleria longifolia (Lindl.) Hanst. Linnaea 29: 524. 1858.

Gesneria longifolia Lindl. in Edwards Bot. Reg. 27: misc. 92, 1841

(based on a plant cultivated in England from material collected in

Guatemala by Hartweg) ; op. cit. 28, t. 4.0. 1842. Isoloma longifolium

Dene. Rev. Hort. 465. 1848. K. linkiana Hanst. Linnaea 26: 160,

t.l,}.18. 1854; op. cit. 29: 522, 565. 1858. Gesneria linkiana Kunth

& Bouche", Ind. Sem. Hort. Berol. 13. 1848 (based on cultivated

material said to be of Guatemalan origin). /. linkianum Hemsl.

Biol. Centr. Am. Bot. 2: 478. 1882. K. chiapensis Brandegee, Univ.

Calif. Publ. Bot. 6: 66. 1914 (type from Chiapas, Mexico).

Usually in damp or wet thickets or forest, sometimes on open or

brushy slopes and banks, 300-2,000 m.; Chimaltenango; Escuintla;

Quezaltenango; El Quiche"; Retalhuleu; Sacatepe"quez ; San Marcos;

Santa Rosa; Solola; Suchitepe"quez. Southern Mexico; El Salvador;

Nicaragua; Costa Rica (fide Standley).

Erect, stout, herbaceous or suffruticose plants to one meter tall, simple or

sparsely branched, the stems densely pilose with appressed or ascending white

hairs; leaves opposite or ternate, short-petiolate, the blades lance-oblong, mostly

8-20 cm. long, acute or acuminate, acute or attenuate to the base, villous-sericeous

with white hairs, densely so beneath, the margins crenate or serrate; inflorescences

appearing racemiform, the area of inflorescence mostly 10-25 cm. long, the flowers

1-many on slender or stout pedicels in the axils of reduced leaves; calyx densely

white-sericeous or strigose, the tube short and broad, the lobes triangular, acute,

about 2 mm. long; corolla brick red, 1.5-2 cm. long, the tube a little ventricose

above, densely red-pilose outside, spotted with darker red in the throat, the lobes

288 FIELDIANA: BOTANY, VOLUME 24

unspotted, 2-4 mm. long, spreading or reflexed; filaments glabrous; disc glands 5,

free; style densely pilosulous, the stigma bilobate; capsule about 8 mm. long, the

free portion conic, densely strigose.

This species appears to be closely related to K. spicata (HBK.)

Hanst., which occurs in much of the same area and in similar locations.

Kohleria rupicola Standl. & L. Wms., Ceiba 3: 62. 1952.

Damp banks, in pine woods or mixed forest, 2,000-2,500 m.;

Jalapa; Zacapa. Honduras (type from Dept. Intibuca, Standley

25168).

Shrubs or suffruticose herbs to 2 m. tall, sparsely branched, the stems and

branches densely villous with multiseptate hairs; leaves opposite, those of a pair

mostly 3-12 cm. long, 1.5-5 cm. wide, acuminate, more or less rounded at the base,

scabrous and more or less papillose-rugose to bullate on the upper surface, usually

densely villous with spreading hairs below, the margins crenate, lateral veins

7-9 pairs; peduncles axillary, short, 0.2-1 cm. long, often inconspicuous; pedicels

solitary or geminate, 1.5-4 cm. long, densely villous, subtended by a pair of linear

bracts 5-7 mm. long; calyx lobes linear to narrowly linear-lanceolate, attenuate to

the apex, variable in length, 5-17 mm. long, densely villous; corolla red, about

2.5 cm. long, the tube white strigose outside, the lobes about 5 mm. long, rounded,

entire or obscurely and irregularly denticulate; disc annular, nearly entire or very

shallowly and irregularly lobate; ovary villous; style pubescent, the stigma

stomatomorphic; capsule densely villous.

The leaves of the plants from Jalapa are not as rugose nor as

hairy as those from Zacapa and Honduras. It is also interesting

that on the type sheet, there are three specimens, two of which have

calyx lobes 5-7 mm. long, and one (on the right) has calyx lobes 13

mm. long. Those of a second Honduran collection, Molina & Molina

244.35, range from 7-12 mm. in length. The specimen from Zacapa,

Steyermark 42^76, has short calyx lobes, 5-7 mm. long; all specimens

from the two collections from Jalapa, Steyermark 32710 and Williams

13177, have calyx lobes 12-17 mm. long.

Kohleria skutchii Morton and Gibson, Phytologia 23: 336.

1972.

Damp or wet mountain forest, 2,400 m., Quezaltenango (type

from Fuentes Georginas, near Zunil, Skutch 906). Mexico (Chiapas).

Slender shrubs 2-3 m. tall, the stems and branches velutinous with multi-

septate, reddish hairs; leaves opposite, those of a pair subequal, on velutinous

petioles 3-7 cm. long, the blades oblong-ovate to elliptic-oblong, mostly 12-28 cm.

long, 5-13 cm. wide, acuminate, oblique and rounded to subcordate at base,

densely velutinous to tomentose above, velutinous below, the margins serrate to

crenate, lateral veins 9-10 pairs; inflorescences axillary, the common peduncle

FIG. 57. Kohleria skutchiL A, flowering branch of plant, X 1A'> B, flower,

X 2; C, calyx with pistil, X 2; D, corolla opened to show stamens and staminode,

X 2; E, two ovaries, X 5; F, two views of anthers, X 5; G, detail of corolla lobes,

X2.

289

290 FIELDIANA: BOTANY, VOLUME 24

elongating, 7-9 cm. long, velutinous, the pedicels usually 4, sometimes 3, velutinous,

3-4 cm. long, subtended by a pair of leaflike bracts 10-15 mm. long; calyx densely

velutinous, the lobes broadly ovate to obovate, acute or obtuse, 4-5 mm. long;

corolla erect in the calyx, 3.5-4 cm. long, the tube orange-red, velutinous outside,

slightly dilated at the base but not saccate, more or less ventricose above, 10-15

mm. wide, the lobes yellowish, marked with red, 7-9 mm. long, suborbicular, the

margins more or less pectinate-denticulate; stamens short-exserted, the filaments

inserted on the base of the corolla tube, pubescent, dilated at base, the anthers

initially coherent in a square; ovary densely villous; style pubescent; stigma

stomatomorphic; disc annular, the glands much reduced, irregular and incon-

spicuous; capsule unknown.

One sterile specimen from San Marcos, Williams, Molina, &

Williams 26252, previously identified as K. skutchii, does have the

large leaves of the species, but the stems and petioles are long-villous

with spreading, whitish hairs and the indument of the leaf blades is

comparatively sparse.

Kohleria spicata (HBK.) Hanst. Linnaea 29: 520. 1858.

Gesneria spicata HBK. Nov. Gen. & Sp. 2: 393, t. 188. 1817. G.

spicata var. schiedeana DC. Prodr. 7: 531, 1839. G. schiedeana

Hook., Curtis's Bot. Mag. 71. t. 4152. 1845. Isoloma spicatum Dene.

Rev. Hort. 465. 1848. K. ignorata Regel, Bot. Zeit. 1851 : 893. 1851 ;

Gartenflora 1: 1, t. 1. 1852. K. wagneri Regel, Ind. Sem. Hort.

Turic. 1853; Gartenflora 3: 347, 1. 103. 1854. I. kramerianum Lehm.

Hamb. Gartenz. 10: 458. 1854. K. schiedeana Hanst. Linnaea 29:

518. 1858. K. tetragona Oerst. Vidensk. Selsk. 5: 101. 1858. 7.

schiedeanum Hemsl. Biol. Centr. Amer. Bot. 2: 479. 1882. 7.

tetragonam Hemsl. op. cit. 480. Khos-corrusumal (fide Aguilar), and

mazoroo (Guatemala).

Damp or wet thickets or pine forest; sometimes on open slopes

or roadside banks, sea level to 1,500 m.; Alta Verapaz; Chiquimula;

Guatemala; Izabal; Pet£n; El Quiche"; Zacapa. Southern Mexico to

Panama; northern South America.

Stout, erect, usually simple, herbaceous or suffruticose plants to 3 meters

tall, the stems villous-hirsute with spreading reddish or brownish hairs; leaves

opposite or ternate, petiolate, the blades elliptic to lance-elliptic or oblong-ovate,

mostly 7-20 cm. long, acute or acuminate, obtuse to cuneate at the base, hirtellous

to velutinous above, villous-tomentose and paler beneath, often with red veins,

the margins crenate-serrate; inflorescences appearing racemiform, 5-35 cm. long,

dense or interrupted, the flowers 1-4 in the axils of reduced bractlike leaves, the

pedicels usually stout, to 2.5 cm. long; calyx villous-hirsute with spreading,

brownish or reddish hairs, the tube 2-3 mm. long, the lobes green within, ovate to

oblong-ovate, often becoming recurved, acute or obtuse, 3-4(5) mm. long; corolla

red to orange-red, 1.2-2 cm. long, red-villous outside, the tube ventricose, more or

GIBSON: FLORA OF GUATEMALA 291

less curved, the throat slightly contracted, the lobes spreading or reflexed, slightly

unequal, 2 or 3 more or less spotted; filaments glabrous, dilated at base, the anthers

about 2 mm. long; style pilosulous; stigma shallowly bilobate; disc glands 5 and

free, or the 2 posterior ones more or less connate; capsule about 8 mm. long,

densely villous-hirsute.

Morton (Baileya 15: 77. 1967) separates K. schiedeana from

K. spicata, stating that the typical form of K. spicata occurs from

Costa Rica to South America, that its inflorescences are much more

elongated than those of the Mexican plants and that the corolla

lobes "are more generally red and with large red blotches toward

the center but without a definite central red line (or lines) and with no

red lines or rows of dots extending down into the tube on the ventral

side." However, as areas of inflorescence of Guatemalan plants vary

from 5 to 35 cm. in length, and as color markings within the corollas

are rarely discernible in herbarium specimens, I am unable to

distinguish two species. As noted by Morton, an extreme form with

slender, elongated pedicels occurs in Mexico and Guatemala.

MONOPYLE Moritz

Slender perennial herbs, erect or subscandent, the stems sparsely branched or

simple, often reddish; leaves of a pair connected by a stipular line, mostly very

unequal, the smaller ones often stipuliform, the blades thin, serrate or crenate;

inflorescences terminal, more or less paniculate, few-many-flowered, the flowers

pedicellate; calyx lobes 5, equal or nearly so, entire; corolla open-campanulate,

narrowed at the base or sometimes a little saccate, minutely pubescent outside,

the limb broad, the lobes spreading, subequal; stamens 4, with a staminode

usually present, the filaments inserted on the base of the corolla tube, incurved

above; anthers coherent, bithecous, thecae slightly divergent; ovary almost wholly

inferior, the placentae ovuliferous on both sides; style short and thick, glabrous;

stigma stomatomorphic; disc absent; fruit capsular, linear-oblong, enclosed in the

calyx, opening by 2 longitudinal slits.

About eight species, ranging from Guatemala to Bolivia. Only

one is known from Guatemala, but another, M. maxonii Morton, is

found in southern Central America.

Monopyle puberula Morton, Field Mus. Bot. 18: 1184. 1938.

Damp or wet forest, usually on limestone, 500-1,500 m.; Alta

Verapaz. Costa Rica.

Plants erect, 30-80 cm. tall, the stems reddish, simple or sparsely branched,

sparsely pilosulous with short, whitish, uncinate hairs; leaves of a pair very unequal,

the larger ones on pubescent petioles 5-14 mm. long, the blades oblong or lance-

oblong, mostly 5-13 cm. long and 1.5-5 cm. wide, acuminate, very oblique and

LJ

FIG. 58. Monopyle puberula. A, habit, X 1A; B, flower, X 2^; C, corolla

opened to show stamens and staminode, X 2>£; D, calyx and pistil, X 2^; E,

two views of fruiting calyx, one opened to show seeds within capsule, X 2^.

292

GIBSON: FLORA OF GUATEMALA 293

obtuse at the base, the margins finely crenate-serrate or serrate, green above and

pilosulous, reddish beneath, puberulent on the costae and veins, minutely strigil-

lose between the veins; smaller leaves mostly 0.5-1 cm. long, subsessile or short-

petiolate, mostly ovate to rounded, or sometimes stipuliform; inflorescences 5-15

cm. long, subumbellate, pedunculate, the bracts commonly small and inconspic-

uous; pedicels usually 2-3 together, unequal, 5-15 mm. long; calyx tube cylindric,

in anthesis 3-4.5 mm. long, about 2 mm. broad, attenuate to the base but in fruit

becoming more or less saccate at the base, densely pilosulous with uncinate,

eglandular hairs, the lobes linear-oblong, 5-6.5 mm. long, green or tinged with

rose; corolla 20-28 mm. long, the tube white and the limb lavender, the tube

appressed-pilosulous outside, abruptly ampliate above, the limb about 2.8 cm.

broad, the lobes subequal, rounded, entire, eciliate, glabrous within; stamens

included, the anthers about 1 mm. long; capsule linear-oblong, 12-15 mm. long,

enclosed in the calyx.

It should be noted that, although according to the original

description the petioles of this species are commonly not more than

5 mm. long, the collection from Chicoy, at about 600 m., Tuerckheim

7929, annotated and cited by Morton as M. puberula, has numerous

petioles 7-14 mm. long, one to 18 mm., and that the leaves are nearly

glabrous beneath, as in the Costa Rican M. maxonii Morton. Some

calyx lobes on this collection are linear or linear-oblong but a few, on

younger flowers, are ovate-oblong to lanceolate, and somewhat

recurved as in M. maxonii. However, the hairs on the petioles are

not as long as those on the typical M. maxonii.

NAPEANTHUS Gardner

Reference: A. J. M. Leeuwenberg, Revision of Napeanthus, in the

Gesneriaceae of Guiana, Acta Bot. Neerl. 7: 340-354. 1958.

Terrestrial, usually perennial, herbaceous or sometimes suffruticose plants,

the stems very short; leaves crowded at the apex, appearing rosulate, sessile or

subsessile, the blades (in ours) oblanceolate to oblong-spathulate, acute or obtuse,

narrowed to the base, the margins dentate or serrate or almost entire; inflorescences

axillary, cymose or subumbellate or the pedicels rarely solitary, but usually

pedunculate and bracteate at the base of the pedicels; calyx campanulate, the 5

segments oblong or lanceolate, acuminate, entire, more or less venose, first connate

for one-third to one-half their length but later often becoming more deeply parted;

corolla campanulate (in ours) or subrotate, the tube short, the limb nearly regular

or somewhat bilabiate, the 5 lobes obtuse, subequal, spreading; stamens 4 with a

staminode usually present, or 5, included or exserted, the filaments inserted on the

base of the corolla; anthers free, the thecae confluent at the apex, fully dehiscent;

ovary superior, the style often curved, the stigma obscurely bilobate; placentae

ovuliferous on both sides; disc absent; capsule shorter than the calyx, bivalvate

(in ours) ; seeds ellipsoidal.

294 FIELDIANA: BOTANY, VOLUME 24

Twelve species, all in tropical America, from Guatemala south-

ward to Bolivia, with only one in Guatemala.

Napeanthus bracteatus Morton, Contr. U. S. Nat. Herb. 29:

39. 1944.

Dense, wet, mixed mountain forest, mostly on limestone, Alta

Verapaz (type from Secoyote" near Finca Sepacuite", Cook & Griggs

Plants 10-25 cm. tall, in age developing a woody, simple stem 5-10 cm. long;

leaves sessile, densely crowded at the apex of the stem, the blades thin, 7-27 cm.

long and 2-8 cm. wide, oblanceolate to oblong-spathulate, acute, attenuate almost

to the base and subcordate, more or less strigillose above, subtomentose beneath,

especially on the costae and the 10-15 pairs of lateral veins, the margins dentate

or denticulate; inflorescences usually numerous, the peduncles 5-10 cm. long,

sparsely pilose, the pedicels 2-5 cm. long, bibracteate below, the bracts foliaceous,

ovate or broadly lanceolate, acute, entire, 1-2 cm. long, sparsely pilose; calyx

7-10 mm. long, the segments equal or nearly so, lanceolate, acuminate, entire,

sparsely pilose outside, venose, ciliate; corolla white, campanulate, 10-12 mm.

long, the tube about 6 mm. long, minutely puberulent or almost glabrous, the limb

slightly bilabiate; stamens 4, didynamous, included; capsule one-half to one- third

as long as the calyx, glabrous.

Known only from Alta Verapaz, where it appears not to be

uncommon.

NIPHAEA Lindley

Low terrestrial herbs, the stems short, erect, simple (in ours) ; leaves petiolate,

crowded at the top of the stem (in ours), the blades thin, ovate or elliptic, serrate

or dentate; inflorescences axillary; pedicels 2-several in each leaf axil; bracts none;

calyx turbinate-campanulate, the tube adnate below to the ovary, the 5 lobes equal

or nearly so; corolla subrotate, the tube very short, the 5 lobes broad, the lateral

ones outside in bud; stamens 4 or 5, or 4 with a staminode, the filaments short,

erect, inserted on the base of the corolla, the anthers oblong, introrsely dehiscent,

thecae confluent at the apex; disc absent; ovary half-inferior, the style rather stout,

curved, the stigma stomatomorphic (in ours) but not much dilated; capsule

half-inferior, the free portion conic, bivalvate; seeds very numerous.

Of the four or five species in Central and South America, only

one is known from Guatemala.

Niphaea oblonga Lindl. Bot. Reg. 27: Misc. 80. 1841; 28: t. 5.

1842.

Based on plants cultivated in England, sent from Guatemala by

Hartweg. About 1,500 m.; Guatemala; Santa Rosa.

FIG. 59. Napeanthus bractealus. A, habit of plant, X K; B, pedicels arising

from bracts. (Illustrations of floral details could not be made, as no corollas were

found on any of the specimens examined, including the type material.)

295

296

FIELDIANA: BOTANY, VOLUME 24

FIG. 60. Niphaea oblonga. A, habit, X %; B, detail of indument, greatly

enlarged; C, flower, X 3J^; D, calyx with corolla in bud, X 3^.

Small, erect herbs, the stems slender, reddish, 3-15 cm. tall, villous; leaves

few, crowded at the top of the stem, on petioles 1-5 cm. long, the blades ovate,

obtuse or acute, cordate or rounded at the base, 4-10 cm. long, 2.5-7 cm. wide,

more or less hirsute, the margins coarsely serrate to biserrate; pedicels 2-several

in the upper leaf axils; calyx tube densely villous, the lobes lanceolate, acute,

green, entire, accrescent, 3.5-7 mm. long, sparsely villous; corolla white, more or

less villous outside, deeply lobate, the lobes broad, rounded, the limb 2-2.5 cm.

broad; stamens 4, the anthers 1.5-2 mm. long.

PHINAEA Bentham

Low, terrestrial, perennial herbs, the stems short, simple, erect or ascending;

leaves opposite, often clustered at the apex of the stem, petiolate, the blades thin,

soft, the margins dentate, serrate, or crenate; inflorescences axillary, the pedicels

filiform, 2 to many in the upper leaf axils; calyx turbinate-campanulate, the tube

adnate to the ovary, the 5 lobes spreading, equal or nearly so; corolla subrotate,

the tube very short, the limb spreading, the 5 lobes broad; stamens 4, with a

staminode usually present, the filaments inserted on the base of the corolla, the

anthers obovate to subglobose, thecae distinct, not confluent at the apex, dehiscent

above by a short slit; disc none; ovary more than half inferior, the style rather

stout, usually somewhat curved, the stigma only a little dilated; fruit capsular,

bivalvate; seeds numerous.

GIBSON: FLORA OF GUATEMALA 297

Nine species, ranging from Mexico to northern South America,

with two in Guatemala and British Honduras.

Leaves subsessile or on petioles usually less than 1 cm. long; corolla limb about

10 mm. in diameter P. repens.

Leaves on petioles 1-3 cm. long; corolla limb 6-7 mm. in diameter. .P. parviflora.

Phinaea parviflora (A. Brongn. & Bouche") Benth. ex Solereder,

Beih. Bot. Centralbl. 24, Abt. 2: 437. 1909. Niphaea parviflora A.

Brongn. & Bouche", Linnaea 25: 299. 1852.

Described from plants grown in Europe, said to have been col-

lected in Guatemala by Warscewicz. Mexico (Chiapas); British

Honduras.

Low herbs, the stems simple, to about 10 cm. long, reddish, pilose with white

hairs; leaves on pubescent to pilose petioles 1-3 cm. long, the blades subequal or

unequal, elliptical, obovate, or elliptic-ovate, 4-11 cm. long, 2-7 cm. wide, obtuse

to subacute, acute or cuneate at base, pubescent with short hairs on both surfaces

but often also pilose beneath on costae and veins, lateral veins 7-10 pairs, the

margins irregularly bidentate or crenate-dentate; pedicels few to numerous in the

leaf axils, mostly 3-6 cm. long, filiform, pubescent; calyx pilose at base, the

segments 2-3 mm. long, ovate-oblong to lance-oblong, acute or subac\ite, entire or

sometimes more or less denticulate near the apex, pubescent; corolla white, the

limb deeply lobate, about 7 mm. in diameter; capsule subglobose, included in the

calyx.

According to the original description, the plants are white-villous

and the calyx segments are ovate and acuminate. Our plants are

pubescent to pilose, with ovate-oblong to lance-oblong calyx seg-

ments. However, other characters, such as the unusually small

corolla, do match. To describe a new species at this time, without

having seen the original material of P. parviflora, seems unwise.

Phinaea repens (Donn.-Sm.) Solereder, Beih. Centralbl. 24,

Abt. 2: 435. 1909. Napeanthus repens Donn.-Sm. Bot. Gaz. 31:

118. 1901.

On rocks or tree trunks in wet, dense forest, 350-1,200 m.; Alta

Verapaz (type from Cubilgiiitz, Tuerckheim 7647) ; Chimaltenango.

Small herbs, the stems 2-6 cm. long, pilose with weak white hairs; leaves mostly

clustered at the end of the stem, subsessile or on petioles usually less than 1 cm.

long, rarely to 2 cm., the blades oblong-elliptic, mostly 4-12 cm. long, 2-5 cm.

wide, obtuse or acute, usually acute at the base, rarely nearly rounded, more or

less pilose on both surfaces, the lateral veins 6-10 pairs, the margins serrate-

dentate; pedicels few, filiform, 1-4 cm. long, pilosulous; calyx segments linear-

lanceolate to ovate-lanceolate, entire, about 2 mm. long at anthesis, pubescent;

corolla white, sparsely pubescent, the limb about 1 cm. in diameter, the lobes

298

FIELDIANA: BOTANY, VOLUME 24

FIG. 61. Phinaea repens. A, habit, X 1A; B, detail of indument, greatly

enlarged; C, flower with corolla opened to show stamens, staminode, and pistil,

X 5; D, view within corolla, showing filament attachment, X 4; E, calyx with

capsule, X 4, and inset of two seeds, greatly enlarged.

ovate, rounded, 3-4 mm. long; stamens glabrous, the filaments 1-2 mm. long; ovary

pubescent; capsule subglobose, included in the calyx.

RECHSTEINERIA Regel

Reference: Moore, H. E., A Proposal for the Conservation of the

Name Rechsteineria, Baileya 2: 24. 1954.

Terrestrial, herbaceous or somewhat suffruticose perennials from well-de-

veloped tubers, usually abundantly pubescent throughout, the stems erect, mostly

simple; leaves opposite or in whorls of 3 to 5, petiolate or sessile, the blades crenate

or dentate; inflorescences variable, axillary, the pedicels solitary, bracteate,

fasciculate or cymose, sometimes pedunculate and subumbellate, or often appear-

ing subracemose and terminal; calyx campanulate, nearly free from the ovary, the

tube adnate only at the base, the 5 lobes shorter than the tube (in ours); corolla

usually red or orange, tubular, erect, usually dilated to saccate at the base, only

a little contracted in the throat, the limb more or less bilabiate with the upper lip

erect and bilobate (in ours) or with 5 subequal short lobes; stamens 4, didynamous,

the filaments inserted on the base of the corolla tube, the anthers united by their

tips to form a square, circle, or a cross, thecae slightly divergent at base, fully

dehiscent; disc glands commonly 5, sometimes fewer, the 2 posterior ones larger,

sometimes connate, the others smaller or sometimes absent; ovary almost wholly

GIBSON: FLORA OF GUATEMALA 299

superior; style exserted, stigma stomatomorphic; fruit capsular, bivalvate; seeds

numerous, shining, fusiform.

About 75 species, most numerous in South America, but extend-

ing as far north as Mexico. The name Rechsteineria has been

conserved; the generic name Corytholoma has most commonly been

used for these plants in Central America, where only one species

occurs.

Rechsteineria warscewiczii (Bouche" & Hanst.) 0. Kuntze,

Rev. Gen. 2: 474. 1891. Gesnera warscewiczii Douche" & Hanst. Ind.

Sem. Hort. Berol. 1861; App. 9. 1861; Linnaea 34: 273. 1865-66.

Corytholoma warscewiczii Standley, Field Mus. Bot. 8: 43. 1930.

Damp or dry, open or brushy, sometimes rocky slopes, 300-2,300

m.; Chiquimula; Huehuetenango ; Jutiapa; El Quiche"; Santa Rosa;

Zacapa. Southern Mexico; Honduras; Canal Zone. Described from

cultivated plants believed to have been of Guatemalan origin.

Erect, herbaceous or somewhat suffruticose plants, the stems stout, simple

or sparsely branched, to 1 m. tall, densely villous with rather short, spreading

hairs; leaves opposite or whorled, a larger one often accompanied by small axillary

leaves, on short or elongated, stout petioles, the blades oblong-ovate to elliptic-

oblong or obovate-oblong, mostly 5-13 cm. long, obtuse or subacute, attenuate

to the base, rather thick, densely villous-hirsute on both surfaces, more densely

so beneath, the margins crenate; flowers axillary but the inflorescences appearing

terminal and racemiform, leafy below, the uppermost leaves reduced and bract-

like, the flowers on stout, erect, or ascending, bracteate pedicels equalling or shorter

than the calyx, broadly campanulate, 1-1.5 cm. long and broad, green or tinged

with red, densely short-villous, the lobes triangular, acute, shorter than the tube;

corolla orange-red or orange (rarely yellow, fide Steyermark), 3.5-4 cm. long,

densely short-villous, the tube somewhat dilated at the middle, slightly contracted

in the throat, the limb very oblique, bilabiate, the upper lip erect and bilobate,

about 1 cm. long, the lower lip very short; stamens exserted, anthers 1-1.5 mm.

long; disc glands usually 4, the posterior one large and bilobate, or 2 glands con-

nate, the others much smaller; capsule 12-15 mm. long, ovoid, acute, densely

hispidulous, little exceeding the somewhat accrescent calyx; seeds very numerous

minute, fusiform.

Handsome, showy plants, in general appearance much like some

species of Kohleria.

RHYNCHOGLOSSUM Blume

Reference: Burtt, B. L., Studies in the Gesneriaceae of the Old

World, XXIII: Rhynchoglossum and Klugia, in Notes Royal Bot.

Gard. Edinb. 24(2): 167-171. 1962.

FIG. 62. Rechsteineria warscewiczii. A, habit, X H; B, flower, X \y>\ C,

detail of anthers, dorsal view, X 2; D, dissection showing pistil and disk glands,

X 1^2- (Leaf pubescence not shown.)

300

GIBSON: FLORA OF GUATEMALA 301

Somewhat succulent terrestrial herbs, sometimes becoming fragile or brittle,

often rooting at the lowest nodes, the stems erect or ascending, simple or branched,

glabrous or pubescent; leaves alternate or one of the leaves of a pair small and

stipuliform, the blades thin, inequilateral, obliquely asymmetrical at the base;

flowers secund and lax in terminal racemes or the racemes finally opposite the

leaves, the bracts minute; flowers pedicellate, the pedicels becoming pendulous;

calyx tubular-campanulate, the tube symmetric or sometimes gibbous or calcarate

at the base, the 5 lobes shallow or parted to the middle; corolla tube cylindric,

symmetric at the base or somewhat gibbous, little ampliate above, almost closed

in the throat, the limb bilabiate, the posterior lip very short, erect-spreading, entire

or bilobate, the anterior lip large, convex, sinuate or broadly and shallowly trilo-

bate; perfect stamens 4 (in ours), didynamous, included, the filaments slightly

flattened, inserted above the middle of the corolla tube; anthers united in a

disclike form (in ours), the thecae divaricate but confluent at the apex; a staminode

sometimes present; disc carnose, cupular, surrounding the superior ovary; style

elongated, the stigma obliquely dilated, obscurely bilobate; placentae ovuliferous

on all sides; capsule included in the calyx, ovoid or elliptic, bivalvate.

A remarkably disjunct genus, with 13 species distributed in New

Guinea, Philippine Islands, Borneo, Malaya, China, India, Ceylon,

Colombia, Central America, and Mexico. Only one species is known

in Guatemala.

Rynchoglossum azureum (Schlecht.) Burtt, Notes Royal Bot.

Gard. Edinb. 24(2): 168. 1962. Klugia azurea Schlecht. Linnaea 8:

248. 1833.

Damp or wet, mixed, mountain forest, 1,200-2,500 m.; San

Marcos; Suchitepe"quez. Southern Mexico; Costa Rica.

Somewhat succulent herbs, erect or nearly so, commonly 30-50 cm. tall,

simple or sparsely branched, the stems stout, finely and often very inconspicuously

puberulent; leaves alternate, on petioles mostly 1-2.5 cm. long, the blades thin,

ovate, ovate-oblong, or elliptic, mostly 7-15 cm. long and 3-8 cm. wide, acuminate

or long-acuminate, very unequal at the base, rounded or semicordate on one side,

semicuneate on the other, glabrous above or nearly so, glandular-puncticulate

beneath, the margins undulate-dentate or subentire: racemes short-pedunculate,

usually many-flowered, equalling or shorter than the leaves, the pedicels and bracts

alternate; bracts linear, 2-3 mm. long; pedicels slender, 5-8 mm. long; calyx

purple or bluish, accrescent, 6-10 mm. long, a little saccate at the base, glabrous,

the lobes shallow, triangular, obtuse or subacute; corolla bright violet-blue, about

2.5 cm. long, not gibbous at the base, glabrous, the lobes denticulate; anthers

heart-shaped to reniform; capsule ovoid.

SOLENOPHORA Bentham

Terrestrial shrubs or suffruticose herbs, often large and coarse, pubescent or

glabrate, sparsely branched or simple; leaves opposite, petiolate, often very large,

FIG. 63. Rhynchoglossum azureum. A, habit, X 1A'> B» flower dissected to

show pistil and stamens, X 3^; C, detail of anthers, X ±1A; D» flower, X 21A;

E, calyx opened to show dehiscent capsule, X 3 1/2; seed, X 5.

302

GIBSON: FLORA OF GUATEMALA 303

those of a pair equal, subequal, or unequal, the margins of the blades dentate or

serrate; inflorescences usually pedunculate (the common peduncle rarely obsolete),

axillary, the flowers pedicellate, solitary or few; pedicels bibracteate; calyx tube

adnate below to the ovary, free above, much exceeding the ovary, the limb 5-

lobate, sometimes bilabiate; corolla (in ours) reddish orange, orange, or yellow,

the tube elongated, ampliate above, ecalcarate, the throat broad, the limb suberect,

the 5 lobes usually shallow, broad; stamens 4, included or exserted, the filaments

inserted at or near the base of the corolla tube, dilated at the base, the anthers

coherent to form a square, rectangle, or circle, the thecae parallel or a little diver-

gent, distinct, dehiscent throughout; a small staminode sometimes present; disc

usually reduced to one more or less bilobate gland, rarely 2, 3, or 5 glands present;

ovary wholly inferior, flat or scarcely convex at the apex, the style elongated,

thickened at the apex, the stigma dilated, stomatomorphic or shallowly bilobate ;

fruit capsular; seeds numerous, minute, more or less fusiform.

Sixteen or more species, in Mexico and Central America, with

10 in Guatemala.

Corollas 6-8 cm. long.

Calyx pilose, the limb at anthesis truncate or nearly so; corolla pilose. S. toucana.

Calyx glabrous or only sparsely hirsute, the limb never truncate; corolla glabrous

or nearly so.

Calyx in bud conspicuously urceolate, the limb narrowly contracted; at

anthesis calyx 3.5-4 cm. long S. obliqua.

Calyx in bud urn-shaped to tubular-campanula te; at anthesis calyx 2.5-3 cm.

long S. purpusii.

Corollas 2-5 cm. long.

Leaf blades mostly 15-30 cm. long, 8-15 cm. wide.

Common peduncle 1-2 cm. long; calyx 2-2.5 cm. long; corolla 2.5-3 cm. long,

spotted or striped inside with maroon S. maculata.

Common peduncle 5-10 cm. long; calyx 1-2 cm. long; corolla 4-5 cm. long,

not spotted nor marked inside S. pirana.

Leaf blades mostly 4-15 cm. long, 2.5-8 cm. wide.

Plants completely glabrous throughout S. erubescens.

Plants more or less villous, at least in part.

Common peduncle 2-8 cm. long S. tuerckheimiana.

Common peduncle usually less than 1 cm. long, sometimes obsolete.

Calyx lobes irregularly semiorbicular or shallowly ovate to very broadly

triangular, mostly 1-3 mm. long; corolla yellow, sometimes marked

with maroon lines, the limb 2-3 cm. wide.

Leaf blades very sparsely short-villous above, often rough to the

touch; calyx sparsely short-pubescent, 1.5-2 cm. long at anthesis;

corolla limb about 2 cm. wide, the lobes conspicuously fimbriate-

pectinate, the projections 1 mm. long or more S. wilsonii.

Leaf blades densely villous above, not rough to the touch; calyx

densely villous, 1-1.5 cm. long at anthesis; corolla limb about 3 cm.

wide, the lobes nearly entire or obscurely denticulate, the projec-

tions much less than 1 mm. long S. abietorum.

Calyx lobes almost regular and distinctly triangular, mostly 4-5 mm.

long; corolla orange, the limb to 1.5 cm. wide S. chiapasensis.

304 FIELDIANA: BOTANY, VOLUME 24

Solenophora abietorum Standl. & Steyerm. Field Mus. Bot.

23:236. 1947.

Damp or wet, mountain forest, sometimes with Abies, 1,500-

2,800 m.; Huehuetenango (type from Cerro Huitz, Sierra de los

Cuchumatanes, between Mimanhuitz and Yulhuitz, Steyermark

48643) collected also on Cerro Canana, near Canana.

Shrubs or suffruticose herbs, erect or decumbent, simple or sparsely branched,

50 cm. tall or more, the stems obtusely tetragonous, sparsely or densely villous

with long, lax, whitish, multiseptate hairs; leaves of a pair subequal, the slender

petioles 3-5.5 cm. long, the blades broadly ovate to obovate-elliptic, mostly

4-9 cm. long, 2.5-5.5 cm. wide, acute or short-acuminate, somewhat unequal at

the base and rounded or obtuse, serrate-dentate with somewhat appressed teeth,

green above and densely villous, often wine-colored beneath, densely and laxly

villous on the veins and costae, the lateral veins 6-9 pairs; flowers axillary, solitary,

the peduncle about 0.5 cm. long, pedicels 1-2.5 cm. long, densely villous, subtended

by a pair of linear bracts 7-10 mm. long; calyx 12-17 mm. long, broadly campanu-

late, green, sometimes marked with red, densely villous, the 5 lobes semiorbicular

to very broadly ovate, 2-4 mm. long, ciliate, obscurely denticulate; corolla yellow

with brown or purple lines within the throat, 3.5-4 cm. long, sparsely villous

outside, the tube ampliate above, about 1 cm. broad at the apex, the limb about

3 cm. broad, sparsely villous within the throat, almost regular, shallowly lobate,

the lobes obscurely denticulate; filaments glabrous; style villosulous; disc gland

solitary, bilobate, pubescent; capsule not seen.

Solenophora chiapasensis D. Gibson, Phytologia 23: 337. 1972.

Not reported from Guatemala, but may be expected there.

Chiapas, Mexico, 2,000-2,800 m. (type from Tenejapa, 2,775 m.,

Breedlove 15187).

Weak shrubs, subrepent or ascending to perhaps a meter tall, branching, the

older stems glabrate, the younger stems and branches more or less villous with very

short, reddish, multiseptate, spreading hairs; leaves of a pair subequal to unequal,

on petioles 1-4 cm. long, the blades elliptic to oblanceolate, mostly 3-9 cm. long,

1.5-4.5 cm. wide, acute or abruptly short-acuminate, cuneate or attenuate to the

base and often oblique, the margins serrate, sparsely pubescent to somewhat

scabrous above, glabrous below except on the costae and veins which are more or

less short- villous, often red or reddish, the lateral veins 7-8 pairs; inflorescences

axillary, the pedicels usually solitary, sometimes geminate, sparsely short-villous,

2-3.5 cm. long, subtended by a pair of small, inconspicuous, linear bracts, the

common peduncle obsolete or if present, inconspicuous, 3-8 mm. long; calyx rose-

red or reddish, tubular-campanulate, 8-15 mm. long, almost glabrous or thinly

villosulous, more densely so near the base, the lobes triangular, mostly 4-5 mm.

long, acute, denticulate; corolla orange, about 3 cm. long, sparsely villous or almost

glabrous, the limb 1-1.5 cm. wide, the lobes semiorbicular, 3-5 mm. long, the

margins fimbriate-pectinate. usually conspicuously so; stamens about equalling

the corolla or slightly exserted, the anthers 1.5-2 mm. long; disc gland solitary,

large, very shallowly and irregularly bifid or bilobate, more or less pubescent, at

GIBSON: FLORA OF GUATEMALA 305

least near the apex; style pubescent; stigma stomatomorphic; capsule about 8 mm.

long, enclosed in the accrescent calyx; seeds broadly fusiform.

Solenophora erubescens Donn.-Sm. Bot. Gaz. 16: 197. 1891.

Damp or wet, mixed mountain forest, sometimes in rocky places,

1,100-2,000 m.; Alta Verapaz (type from Pansamala, growing on

rocks about a waterfall, Tuerckheim 631}; Huehuetenango.

Plants herbaceous or suffrutescent, to 1 m. tall, usually simple, glabrous

throughout, the stems terete, often reddish, with conspicuous interpetiolar lines

between the bases of the petioles; leaves of a pair subequal, on petioles 1-5 cm.

long, the blades coriaceous when dried, broadly ovate to oblong-elliptic, mostly

4-8 cm. long, acute to very obtuse or almost rounded at the apex, acute or obtuse

at the base, the margins sinuate-serrate, paler beneath, sometimes dull red or

purple-red beneath; peduncles axillary, 0.5-1 cm. long, flowers 1-3, on pedicels

1-2 cm. long, bibracteate; calyx green or sometimes spotted with red, 13-15 mm.

long, the tube obconic, the lobes short, erect, ovate to broadly triangular, usually

obscurely dentate; corolla deep yellow, 2-2.5 (3) cm. long, glabrous outside, the

tube 3-5 mm. broad, the limb about 1.5 cm. broad, with maroon lines within, the

lobes broad, rounded, undulate, more or less crenate, one lobe sometimes obscurely

pectinate or denticulate; stamens included; capsule said to be ovoid, about 1 cm.

long, crimson, crowned with the persistent bilobate gland.

Solenophora maculata D. Gibson, Phytologia 23: 339. 1972.

Known only from the type, Steyermark 35992, collected between

Todos Santos and Finca El Porvenir, middle slopes of Volcan

Tajumulco, San Marcos, 1,300-2,500 m.

Shrubs or suffruticose herbs, simple or sparsely branched, the young stems

reddish, usually villous, later more sparsely villous or glabrate; leaves of a pair

subequal, on glabrous petioles 5-10 cm. long, the blades broadly elliptic or ovate-

elliptic, mostly 11-25 cm. long, 8-14 cm. wide, short-acuminate, oblique at the

base, sparsely short-villous above, glabrate beneath, the costae and veins rose-red,

the margins doubly serrate, the lateral veins 9-12 pairs; inflorescences axillary,

the common peduncle 1-2 cm. long, sparsely pubescent or glabrate, bracts oblong-

lanceolate or oblong-elliptic, 0.5-1 cm. long, pedicels glabrous, 2-3 cm. long; calyx

suffused with rose or maroon, sparsely villous near base, glabrous above, 2-2.5 cm.

long, the triangular lobes unequal, 2-4 mm. long, more or less denticulate; corolla

2.5-3 cm. long, deep yellow or orange-yellow, spotted or striped with maroon

within, the tube erect, ampliate above, about 1 cm. broad below the lobes, villous

outside, the lobes about 4 mm. long, irregularly undulate; stamens exserted, the

anthers about 2 mm. long; style exserted; stigma stomatomorphic; disc gland

solitary, more or less bilobate; capsule not seen.

Solenophora obliqua Denham and Gibson, Phytologia 23: 340.

1972. Zacate de vaca (San Marcos).

FIG. 64. Solenophora maculata. A, leafy stem with axillary inflorescences, X

%; B, flower, X 11A; C, detail of anthers, X 2J^; D, disk gland and base of style,

X3.

306

FIG. 65. Solenophora obliqua. A, section of flowering branch, X 1A; B, calyx,

X 1M; G, flower with calyx cut away and corolla opened at base to show disk

gland, staminode, and portions of style and filaments, X 1J4; D, two views of

fruiting calyx, the upper one opened to show persistent disk gland and seeds, X 1 ;

E, two seeds, greatly enlarged.

307

308 FIELDIANA: BOTANY, VOLUME 24

Damp or wet, mixed forest, usually on white sand slopes, often

in wet ravines, 1,300-2,700 m.; Chimaltenango; Quezaltenango

(type from western slopes of Volcan Zunil, Steyermark 35182}; San

Marcos; Suchitepe"quez.

Shrubs or weak trees to 6 m. tall, the stems and branches glabrate or sparsely,

bifariously pubescent with multiseptate hairs; leaves opposite, those of a pair

subequal, the petioles essentially glabrous, mostly 3-12 cm. long, the blades

oblong-ovate to elliptic-oblong, acuminate or acute, cuneate or rounded at the

base and usually very oblique, mostly 14-30 cm. long, 7-17 cm. wide, sparsely

pilose above, glabrous or glabrate beneath, the margins doubly serrate, the lateral

veins 10-14 pairs; inflorescences cymose, the common peduncle 1-2 cm. long or

sometimes the flowers solitary, the peduncle then much reduced and inconspicuous,

the pedicels 0.5-3 cm. long, bibracteate, the bracts leaflike, mostly 2-3 cm. long,

sometimes caducous in age; calyx glabrous, brownish or green with orange-brown

stripes, 3.5-4 cm. long, 3 to 4 times longer than the ovary, more or less bilabiate,

conspicuously urceolate in bud, the limb narrowly contracted, the 5 lobes irregular,

short, 3-8 mm. long, triangular to lanceolate, appearing irregularly denticulate

but the lobules actually rounded and obtuse, the tube in anthesis becoming deeply

cleft on one side for about two-thirds of its length, the calyx then appearing

narrowly tubular-campanulate; corolla 6.5-7.5 cm. long, glabrous or nearly so,

pale orange to reddish-orange outside, orange-yellow and sometimes marked with

maroon within, the tube gradually ampliate, the limb somewhat oblique, about

3 cm. wide, the lobes broadly suborbicular, spotted with purple or maroon within,

the margins more or less fimbriate-pectinate; stamens short-exserted, the filaments

glabrous, the anthers initially coherent, the thecae oblong; style densely pubescent;

stigma stomatomorphic; disc gland solitary and usually deeply bilobate, pubescent;

fruit far surpassed by the calyx tube, the capsule rupturing irregularly near the

middle when mature, containing many minute, reddish-brown, shining, ellipsoidal

to fusiform seeds.

Solenophora pirana Morton, Phytologia 1 : 149. 1935.

Damp or wet, mixed mountain forest, 2,300-3,800 m.; Chi-

maltenango (type collected near Chichavac, Skutch 680}; Que-

zaltenango; El Quiche"; San Marcos. Mexico.

Shrubby or suffruticose plants, 1-4 m. tall, the stems erect, stout, simple or

sparsely branched, the young stems sparsely villous with long, spreading, multi-

septate hairs; leaves of a pair subequal, on petioles 4-12 cm. long, these more or

less villous, the blades elliptic or ovate-elliptic, mostly 14-30 cm. long, 9-15 cm.

wide, acute or abruptly short-acuminate, oblique at the base and usually rounded

or subcordate, rarely broadly cuneate, sparsely villous above, usually short-

villous on the veins beneath but sometimes glabrate, the margins doubly serrate,

the lateral veins 9-12 pairs; inflorescences axillary, subumbellate or subracemose,

with 3-5 or more flowers, the peduncles commonly 5-10 cm. long, the bracts oblong

to elliptical, mostly 1-2 cm. long, the pedicels mostly 1-4 cm. long, more or less

villous or glabrate; calyx at anthesis campanulate, almost turbinate in age,

accrescent, 1-2.5 cm. long, villous below, nearly glabrous above, somewhat

oblique at the apex, shallowly lobate, the short lobes irregular, more or less ovate

GIBSON: FLORA OF GUATEMALA 309

to broadly triangular, denticulate; corolla yellow, unmarked within, 4-5 cm. long,

erect, the tube thinly villous, slightly ventricose above, 1-1.5 cm. broad, the limb

narrow, 1-1.5 cm. broad, the short lobes subequal, more or less rounded to nearly

truncate, the margins irregularly undulate; stamens exserted, the filaments

glabrous, the anthers 1.5-2 mm. long; style pubescent; disc glands 1, 2, 3, or 5,

at least the large posterior one pubescent.

Although S. pirana was originally described as having five disc

glands, and that is true of some dissections prepared from type

material, at least one later dissection proved to have only three

glands (one large posterior, more or less bilobate gland and two

smaller anterior ones) . Dissections were then prepared from material

taken from Skutch 31*6 (Chimaltenango), Standley 671*91 and 85936

(Quezaltenango), Proctor 25528 (El Quiche"), and Contreras 5197 (El

Quiche*). All had only one large, more or less bilobate gland, except

Contreras 5197, which had two, the posterior one large and bilobate,

the anterior one small and entire.

Solenophora purpusii Brandegee, Univ. Calif. Publ. Bot. 6:

65. 1914.

Wet, mixed mountain forest, often in thickets along streams,

2,300-2,700 m.; San Marcos; Suchitepe'quez. Mexico (type from

Cerro del Boqueron, Chiapas).

Large, coarse herbs or shrubs 1.5-4.5 m. tall, simple or sparsely branched, the

young stems thick, thinly villous with long, weak, multiseptate hairs; leaves

large, those of a pair unequal to subequal, on petioles 3-10 cm. long, the blades

thin, elliptic, broadly elliptic, or ovate-elliptic, mostly 10-30 cm. long and 7-15 cm.

wide, acute or short-acuminate, very oblique at the base and more or less cuneate

or rarely subcordate, the margins doubly dentate, sparsely short-villous on the

upper surface, slightly paler beneath and often almost glabrous, usually puberulent

or villosulous along the veins; peduncles 0.5-3 cm. long, few-flowered, the flowers

on pedicels 1-3 cm. long, these subtended by a pair of foliaceous bracts 1-2 cm.

long; calyx 2.5-3 cm. long, glabrous, green tinged with dark red, deeply cleft on

one side, the lobes short, irregular, subequal, denticulate, the tube about 1 cm.

broad, conspicuously venose; corolla 7-8 cm. long, deep yellow to orange-yellow,

the lobes spotted with maroon within, glabrous outside, the tube gradually ampliate

upward, about 2 cm. broad above, the lobes short, rounded, sinuate-denticulate;

style pubescent; stigma stomatomorphic; disc gland solitary, more or less bilobate.

Showy and handsome plants, producing an abundance of large,

brightly colored flowers.

Solenophora toucana Denham and Gibson, Phytologia 23: 341.

1972. Trompeta ortigosa (Guatemala).

Known in Guatemala from a single collection, Morales 91*7 from

Palencia, Guatemala, 1,480 m. El Salvador; Honduras.

310 FIELDIANA: BOTANY, VOLUME 24

Shrubs 2-4 m. tall, the stems and branches more or less pilose with multi-

septate hairs; leaves of a pair subequal, on pilose petioles mostly 5-12 cm. long,

the blades oblong-ovate to broadly oblong-elliptic, mostly 16-30 cm. long, acumi-

nate, obliquely cuneate to nearly rounded at base, sparsely or densely pubescent

to pilose above, pilose below on costae and veins, otherwise pubescent or glabrate,

the margins doubly serrate, the lateral veins about 10 pairs; inflorescences cymose,

the common peduncle seldom more than 5 mm. long, the pedicels 5-15 mm. long,

bibracteate, the bracts linear; calyx campanulate, 1.5-2.5 cm. long, more or less

pilose, densely so near the base, very shallowly lobate but appearing almost trun-

cate, the margins unevenly denticulate; corolla orange outside, yellow within,

5.5-7 cm. long, the tube gradually ampliate, pilose outside, the limb 3-3.5 cm.

in diameter, the lobes suborbicular, broadly rounded, spotted with red within, the

margins fimbriate-denticulate; stamens short-exserted, the filaments somewhat

pubescent, the anthers coherent; style exserted, densely pubescent; stigma sto-

matomorphic; disc gland solitary and usually deeply bilobate, pubescent; capsule

unknown.

Solenophora tuerckheimiana Dorm.-Sm. Bot. Gaz. 46: 114.

1908.

Wet, mixed, mountain forest, 1,600-2,800 m.; Alta Verapaz

(type from the region of Coban, Tuerckheim II. 2028} Huehuete-

nango (Cerro Canana, Sierra de los Cuchumatanes).

Herbs or shrubs 1-1.5 m. tall, simple or sparsely branched, the stems and

petioles often reddish, densely villous with very short, multiseptate, spreading

hairs, or merely furfuraceous-villosulous; leaves of a pair subequal to unequal, on

petioles 1-7 cm. long, the blades rather thick when dried, elliptic, obovate-elliptic,

or broadly obovate, mostly 8-15 cm. long and 4-7.5 cm. wide, acute or abruptly

short-acuminate, oblique at the base and usually more or less acute, sometimes

obtuse, rather densely or sparsely villosulous above with very short, thick hairs,

often wine-red beneath, short-villous on the veins, glabrous between the veins,

margins serrate-dentate, lateral veins 8-10 pairs; inflorescences few-flowered,

subumbelliform or cymose, the peduncles mostly 3-8 cm. long, the pedicels 1-2.5

cm. long, often reflexed, subtended by a pair of linear bracts; calyx rose-red, broadly

obconic-campanulate, 7-12 mm. long, thinly villosulous, the lobes very broadly

and irregularly suborbicular to broadly subtriangular or ovate, very short, mostly

1-2 mm. long, rarely to 3 mm., obtuse to almost rounded, denticulate; corolla

orange, 3-4 cm. long, thinly long-villous, the limb about 15 mm. broad, the lobes

semiorbicular, 5-7 mm. long, at least the largest one obscurely short-pectinate,

the others entire or pectinate-denticulate only near the base; stamens exserted,

the anthers 1.5-2 mm. long; disc gland shallowly bilobate but often appearing

entire due to indument; capsule said to be 1 cm. long or longer.

According to the original description, the calyx lobes are "... del-

toidei 4 mm. longi"; however, no calyx lobes on the type collection

or on later collections accurately match this description.

FIG. 66. Solenophora toucana. A, section of flowering branch, X J^; B,

flower with one corolla lobe removed to show anthers and style, and with part of

corolla tube removed to show disk gland and base of filaments and style, X 1/4;

G, calyx, X 1^; D, detail of calyx lobes, X 21A.

311

FIG. 67. Solenophora wilsoniL A, habit, X 1A; B, flower in bud, X 1J^; C,

corolla opened to show stamens and staminode, X lJ/£; D, dorsal and ventral

views of anthers, X 3J/£; E, calyx opened to show pistil and disk gland, X 11A'< F»

dehiscent capsule, X IK-

312

GIBSON: FLORA OF GUATEMALA 313

Solenophora wilsonii Standley, Field Mus. Bot. 17: 210. 1937.

Reina de las flores (Huehuetenango).

Dense, wet, mixed forest 300-2,000 m.; Alta Verapaz (type from

Chacirociha, Finca Seamay, Senahu, Hatch & Wilson 201}; Hue-

huetenango; Izabal.

Erect herbs or shrubs to 1 m. tall, usually simple, the young branches densely

covered with very short, thick hairs which although multiseptate, appear almost

papilliform without high magnification; leaves of a pair subequal, on pubescent

or short-villous petioles mostly 2-8 cm. long, the blades broadly elliptic, oblong-

elliptic, or obovate-elliptic, mostly 7-15 (18) cm. long and 3-8 cm. wide, short-

acuminate, oblique at the base and cuneate to very obtuse or almost rounded, the

margins coarsely dentate to obscurely appressed-serrate, very sparsely short-

villous above, often rough to the touch, glabrous beneath except on the veins and

costae which are covered with very short, thick, almost papilliform hairs, the

lateral veins 9-10 pairs; flowers axillary, solitary or collected in sessile cymose

inflorescences, the common peduncle only 2-5 mm. long or obsolete, the flowers on

short, more or less villosulous pedicels 2-7 mm. long, subtended by a pair of linear

bracts 7-10 mm. long; calyx broadly campanulate, 1.5-2 cm. long, green, sometimes

striped with rose or purple, almost 1 cm. broad, sparsely villosulous, the lobes

short, broad, subequal, shallowly ovate to broadly triangular, obtuse or acute,

more or less dentate; corolla yellow, 3.5-4 cm. long, sparsely long-villous, the limb

about 2 cm. broad, the lobes subequal, rounded, glabrous within, the margins

closely and conspicuously fimbriate-denticulate; stamens included, the anthers

about 1.5 mm. long; disc gland solitary, very shallowly bilobate; capsule unknown.

This species conforms in many ways with the description of the

Mexican S. obscura Hanst. except in corolla color, described by

Hanstein as "albidis." Our plants also closely resemble the type-

photograph of Liebman 9322 (F.M. Neg. 22715), except that the

calyx lobes of S. obscura appear nearly regular and distinctly tri-

angular. Mr. Dale Denham, of the Gesneriad Gardens, Boulder,

Colorado, who has seen the type specimen, has informed me (per-

sonal communication) that the bracts in the S. obscura inflorescences

are foliar, to 5 mm. wide; the bracts on our plants are only 1-2 mm.

wide.

Flora of Guatemala - Part X, Number 4

LENTIBULARIACEAE. Bladderwort Family.

DOROTHY NASH GIBSON

Reference: J. H. Barnhart, Segregation of genera in Lentibularia-

ceae, Mem. N. Y. Bot. Gard. 6: 39-64. 1916. Peter Taylor, Lenti-

bulariaceae, in Flora of Trinidad and Tobago 2: 288-300. 1955;

Lentibulariaceae in Maguire, Bot. Guayana Highlands, VII, Mem.

N. Y. Bot. Gard. 17: 205-228. 1967.

Annual or perennial, insectivorous herbs, growing in water or wet soil, some-

times epiphytic; leaves forming a basal rosette or on stolons or reduced to small

alternate cauline scales; scapes erect, the inflorescence racemose or the flowers

solitary; bracts at the base of the pedicels minute, the bracteoles minute or none,

rarely present at the base of the flower and larger; calyx segments 2, 4, or 5, open

in bud; corolla gamopetalous, the limb bilabiate, the tube short, the lower lip

usually spurred; stamens 2, the filaments inserted on the base of the corolla,

alternate with the lobes of the anterior lip, included; anthers usually transversely

constricted, more or less bithecous, dorsifixed, dehiscent by a common longitudinal

slit; disc none; ovary superior, ovoid or globose, unilocular, the ovules 2 to many

on a free-central placenta; stigma bilamellate, sessile or borne on a very short

style; fruit capsular, globose or ovoid, usually 2-4-valvate or circumscissile, rarely

indehiscent; seeds small, variously shaped, the testa striate, rugose, or reticulate.

Four genera with probably considerably fewer species than the

200 or more reported, widely distributed in tropical and temperate

regions of both hemispheres. Three genera are represented in

Central America.

Calyx segments 2 Utricularia.

Calyx segments 4 or 5.

Scapes ebracteate, bearing a single flower Pinguicula.

Scapes bracteate, usually bearing more than one flower Genlisea.

GENLISEA St. Hilaire

Small, terrestrial or subaquatic herbs; foliage leaves rosulate, arising from a

short perennial rhizome, the blades obovate or spathulate to filiform; trap leaves

descending, flask-shaped; scapes slender, simple, erect, bearing minute scales or

bracts; inflorescence racemose, the flowers usually few, each pedicel subtended by

315

316 FIELDIANA: BOTANY, VOLUME 24

a small bract, minutely bibracteolate at the base; calyx lobes 5, equal or subequal,

the calyx sometimes somewhat bilabiate; corolla bilabiate, the spur incurved, the

posterior lip erect, entire or emarginate, the anterior one larger, spreading, usually

trilobate, the lobes reflexed-spreading; stamens 2, the filaments stout, incurved or

straight, the anthers dorsifixed, thecae divaricate, confluent; capsule circumscissile;

seeds ovoid, reticulate.

Perhaps 15 or more species, mostly in South America, with three

in tropical Africa, and only one in Central America.

Genlisea filiformis St. Hilaire, Voyage Distr. Diam. 2: 430.

1833. G. luteo-viridis Wright in Sauvalle, Anal. Acad. Cien. Habana

6: 314. 1869; Fl. Cub. 90. 1873.

In swampy ground, at or near sea level, British Honduras (All

Pines, Schipp 607). Cuba; Brazil; Colombia; Venezuela; British

Guiana.

Small, slender plants, the leaves few or numerous, forming a basal rosette, the

blades obovate-spathulate to suborbicular, 3-5 mm. long, rounded or very obtuse

at the apex, narrowed at the base into a slender, winged petiole as long as or longer

than the blade; scape filiform, commonly 8-20 cm. tall, wiry, flexuous, bearing a

few long, gland-tipped hairs and 3-5 minute, ovate, acute bracts; flowers 2-4,

remote, slender-pedicellate; calyx lobes subobtuse, elliptic-oblong or lance-oblong,

ciliolate; corolla greenish yellow or lemon-yellow, about 4 mm. long, the spur thick,

very obtuse, only a little longer than the lips; seeds compressed, obovate, angulate.

This genus is apparently known in continental North America

by the single collection cited above.

PINGUICULA L.

References: Alfons Ernst, Revision der Gattung Pinguicula, Bot.

Jahrb. 80, Heft 2: 145-194. 1961. Rogers McVaugh and John T.

Mickel, Notes on Pinguicula, sect. Orcheosanthus, Brittonia 15:

134-140. 1963. S. Jost Casper, Gedanken zur Gliederung der

Gattung Pinguicula L. Bot. Jahrb. 82: 321-335. 1963; Once More:

The Orchid-Flowered Butterworts, Brittonia 18: 19-28. 1966; Mono-

graphic der Gattung Pinguicula L., Bibliotheca Botanica, Heft

127/128: 1-209. 1966.

Terrestrial herbs, growing in wet or very damp places; leaves radical, forming

a rosette, pale green, entire, broad, fleshy, and with an oily or fatty texture (hence

the English name "butterwort" sometimes given to the plants), the surface

viscidulous and with an almost crystalline appearance; scapes erect, naked, bearing

a solitary flower; calyx segments 4 or 5, sometimes appearing bilabiate, the posterior

lip trilobate, the anterior one emarginate, bifid, or bilobate; corolla bilabiate, the

lobes spreading, entire or emarginate, the 2 posterior ones slightly or much shorter

than the others; anthers transverse, confluent, monothecous; lamina of the stigma

FIG. 68. Genlisea filiformis. A, habit, X 1; B, flower, X 13; C, corolla dis-

sected to show pistil and stamens, X 15; D, section of placenta with seeds, greatly

enlarged.

317

318 FIELDIANA: BOTANY, VOLUME 24

very unequal, the posterior one larger, often fimbriate; capsule 2-4-valvate; seeds

oblong, rugulose.

Casper lists 46 species, widely distributed in temperate, cold, and

tropical regions, with three in Guatemala. Some species produce

both a "winter" and a "summer" rosette of leaves, these forms often

varying greatly in size and shape.

Corolla deep red-violet or purple, the spur commonly 15-30 (40) mm. long.

P. moranensis.

Corolla white, or pale lilac, or white marked with lilac, the spur less than 6 mm.

long.

Leaves 15-25 mm. long; corolla 11-12 mm. long, the spur 3.5-5 mm. long.

P. lilacina.

Leaves 5-10 mm. long; corolla scarcely 6 mm. long, the spur about 2 mm.

long P. crenatiloba.

Pinguicula crenatiloba A. DC. in DC. Prodr. 8: 30. 1844.

P. nana Mart, et Gal. ex Hemsl. Biol. Centr. Am. Bot. 2: 471.

1882, nomen.

Grassy slopes or more commonly on bare, wet clay banks, usually

in pine forest, 800-2,100 m.; Chiquimula; Huehuetenango; Zacapa.

Mexico; El Salvador; Honduras; Panama.

Leaves few, rosulate and flat on the ground, very pale green, thick and suc-

culent, broadly obovate, 5-10 mm. long, rounded at the apex, more or less pilose

on the upper surface, the margins often inrolled; scapes 1-several, slender, erect,

mostly 3-6 cm. long, minutely pilosulous; flowers solitary; calyx 1-2 mm. long,

usually minutely pilosulous, the segments ovate; corolla white, usually with a

yellow throat, rarely purplish white, the lips very unequal, the larger lip shallowly

trilobate, the spur slender, about 2 mm. long, often minutely pilosulous.

Very small and inconspicuous plants, difficult to see among the

grasses where they usually grow.

Pinguicula lilacina Schlecht. & Cham. Linnaea 5: 94. 1830.

P. obtusiloba A. DC. in DC. Prodr. 8: 30. 1844. P. scopulorum

Brandegee, Zoe 5: 257. 1908. P. divorum Standl. & Steyerm. Field

Mus. Bot. 23: 179. 1944 (type from Huehuetenango, Steyermark

50061). Flor de piedra (Huehuetenango).

Damp or wet slopes and banks, often in oak or oak-pine forest,

1,900-2,400 m.; Alta Verapaz; Huehuetenango. Mexico.

Leaves numerous, forming a dense rosette closely appressed to the soil, sessile,

the blades pale green and thin, broadly obovate to suborbicular, 15-25 mm. long,

broadly rounded at the apex, broadly cuneate at the base, the margins usually

inrolled, nearly glabrous or with scattered hairs above, obscurely puncticulate;

scapes 1-several, slender, filiform or nearly so, 5-10 cm. long, short-pilosulous or

GIBSON: FLORA OF GUATEMALA

319

FIG. 69. Pinguicula lilacina. A, habit, X/4; B, posterior view of calyx, X4;

G, flower, X 4; D, pistil and stamens, X 4; E, calyx and capsule, X 3, with seeds

greatly enlarged.

glandular-pilosulous near the top; calyx 2-3 mm. long, sparsely and minutely

glandular-pilosulous, the lobes unequal, ovate to oblong, obtuse or rounded at the

apex; corolla pale lilac or white, or white with lilac markings, 11-12 mm. long,

glabrous or very minutely glandular-pilosulous outside, the larger lip trilobate,

the smaller lip about half as long, bilobate; spur nearly straight, 3.5-5 mm. long;

capsule subglobose.

Pinguicula moranensis HBK. Nova Gen. & Sp. 2: 226. 1817;

Casper, Brittonia 18: 26. 1966. P. caudata Schlecht. Linnaea 7: 393.

1832; Hemsley, Biol. Cent. Am. Bot. 2: 470. 1882, pro parte; Ernst,

Bot. Jahrb. 80: 158. 1961, pro parte. P. flos-mulionis Morr., La

Beige Hort. 22: 371. 1872. P. sodalium Fourn., Bull. Soc. Bot. Fr.

20: 67. 1873. P. bakeriana Sander, Gard. Chron. n.s. 15: 541. 1881.

P. rosei Watson, Gard. Chron. ser. 3, 49: 82. 1911.

On shaded, steep slopes, banks, or cliffs, on earth or limestone,

often in pine or pine-oak forest, usually in places saturated with

water, 1,800-3,700 m.; Baja Verapaz; Chimaltenango; Chiquimula;

Huehuetenango; El Progreso; Quezaltenango; El Quiche"; San Marcos.

Central and southern Mexico; El Salvador; Honduras.

Plants with numerous leaves forming a dense rosette, more or less appressed

to the soil, the blades thick and fleshy, mostly 3-6 cm. long, glabrous or nearly

so, rounded at the apex, the winter leaves spathulate, the summer leaves obovate

320 FIELDIANA: BOTANY, VOLUME 24

to elliptic or suborbicular, cuneate or attenuate to the base and decurrent on short

petioles; scapes erect, fairly stout, 5-20 cm. tall, usually solitary, bearing a single

flower, the upper part of the scape and the calyx often minutely glandular-pilo-

sulous; corolla large and showy, deep reddish violet to purple, the limb about

2 cm. long, the spur slender, straight or somewhat curved, commonly 1.5-4 cm.

long, the larger lip deeply trilobate, the lobes broadly oblong to cuneate-obovate,

rounded at the apex.

Fairly common plants in some regions of Guatemala. The leaves

of the "winter rosette" vary greatly in size and form from those of

the "summer rosette."

UTRICULARIA L.

References: Peter Taylor, The genus Utricularia L. (Lentibularia-

ceae) in Africa (south of the Sahara) and Madagascar, Kew Bull.

18(1): 1-245. 1964; Lentibulariaceae, in Maguire and Collaborators,

The Botany of the Guayana Highlands— Part VII, Mem. N. Y. Bot.

Gard. 17(1) : 206-228. 1967.

Annual or perennial, terrestrial, epiphytic, or floating aquatic herbs lacking

true roots, the stems becoming variously modified to form vegetative parts func-

tioning as roots (rhizoids), stems (stolons), and leaves (erect, thalloid, or divided

into capillary segments), or carnivorous traps: leaves rosulate from the base of the

inflorescence or alternate, opposite, or verticillate on the stolons; traps borne on

one or more of the vegetative parts; inflorescence a simple or branched scape,

usually erect, glabrous, filiform, bracteate (the bracts sometimes sterile scales),

bearing one to several flowers disposed in a raceme; pedicels usually short, sub-

tended by a bract and often 2 bracteoles; calyx deeply bilabiate, the 2 segments

equal or subequal, persistent, sometimes accrescent; corolla bilabiate, the upper

lip usually erect, the lower lip usually longer and spreading, spurred at the base;

stamens 2, the filaments inserted at the base of the corolla, usually curved, the

anthers dorsifixed, with more or less confluent thecae; ovary unilocular, style

usually short, persistent, the stigma bilamellate, one lamella usually much larger

than the other; ovules numerous on a basal or free-central placenta; fruit capsular,

dehiscing by valves or pores or sometimes circumscissile or rarely indehiscent;

seeds usually many, variously shaped and sculptured.

Species variable, often polymorphic, perhaps 150 widely dis-

tributed in both hemispheres. In addition to the 13 listed here, a

few more are found in southern Central America.

Plants free-floating aquatics, commonly with elongating stolons, the leaves sub-

merged, bearing numerous conspicuous traps.

Corollas yellow.

Leaves to 2 cm. long, simple or very sparsely branched; bracts minute, usually

much less than 1 mm. long; scapes filiform, bearing 1-6 flowers; upper lip

of trap with 2 long, often branching appendages (often twice as long as

the trap) U. gibba.

GIBSON: FLORA OF GUATEMALA 321

Leaves to 15 cm. long, multiple-pinnate; bracts conspicuous, 2-4 mm. long;

scapes stout (1-3 mm. broad) bearing 5-20 flowers; upper lip of trap

naked or with 2 short, simple or sparsely branched appendages.

U. foliosa.

Corollas not yellow.

Leaves whorled; scapes commonly 5-15 cm. long; corolla pale purple to blue

(in ours) U. purpurea.

Leaves alternate; scapes commonly 2-6 (8) cm. long; corolla pink, reddish, or

rose (in ours) U. hydrocarpa.

Plants terrestrial or epiphytic, the stolons short, the few leaves usually borne on

or near the base of the scape and appearing rosulate, the traps usually minute

and inconspicuous, mostly borne on the stolons but sometimes arising from

the leaves.

Calyx lips deeply fimbriate-pectinate U. fimbriata.

Calyx lips not fimbriate-pectinate.

Plants epiphytic; calyx conspicuously large in proportion to the corolla, the

lips at anthesis 7-8 mm. long but in fruit often 10-12 mm. long and

8-9 mm. wide [7. jamesoniana.

Plants terrestrial; calyx not as above.

Bracts of the scape laciniate-dentate U. hispida.

Bracts of the scape entire.

Bracts peltate or cupular; bracteoles absent.

Bracts cupular; flowers always solitary, more or less resupinate;

corollas rose to purple U. resupinata.

Bracts peltate; flowers commonly 2-25 (rarely only one), not resupinate;

corollas yellow U. subulata.

Bracts neither peltate nor cupular; bracteoles present.

Pedicels 5-20 mm. long; calyx densely and minutely glandular-pubes-

cent; corollas white or sometimes flushed or marked with violet.

U. amethystina.

Pedicels 1-3 mm. long; calyx glabrous; corollas yellow or orange-

yellow.

Spur of corolla about 1 mm. long; filaments of stamens geniculate

about the middle U. guyanensis.

Spur of corolla 4-8 mm. long; filaments not geniculate.

Scapes 10-40 cm. long; calyx prominently venose; spur of corolla

5-8 mm. long U. juncea.

Scapes 5-12 cm. long; calyx not prominently venose; spur of

corolla 3-4 mm. long U. adpressa.

Utricularia adpressa Salzmarm in St. Hilaire & Girard, Ann.

Sci. Nat. II, 11 : 159. 1839. U. aureola Blake, Contr. Gray Herb. 52:

88. 1917 (type from British Honduras, Manatee Lagoon, Peck 235).

U. aureolimba Steyermark, Fieldiana:Botany 28: 535. 1953.

Wet sandy places, at or near sea level; British Honduras. Vene-

zuela; British and French Guiana; Brazil.

Terrestrial herbs, the stolons short; leaves unknown, probably fugacious;

scape erect, almost filiform, glabrous, 10-15 cm. long, bearing 2-8 flowers; bracts

ovate, obtuse or subacute, 0.7-1.2 mm. long, the bracteoles lanceolate, smaller;

322 FIELDIANA: BOTANY, VOLUME 24

pedicels 0.5-3 mm. long; calyx lips ovate to oblong, 2-3.5 mm. long; corolla yellow,

6-8 (12) mm. long, the upper lip elliptic-ovate to suborbicular, entire, the lower

one orbicular, entire, 3-4.5 mm. long; spur 3-4 mm. long; capsule subglobose,

about 2 mm. in diameter; seeds globose.

Utricularia amethystina St. Hilaire & Girard, Ann. Sci. Nat.

II. 11: 163. 1839. U. sinuata Benj. Linnaea 20: 491. 1847. U.

adenantha Standley, Field Mus. Bot. 23: 179. 1944 (type from

British Honduras, Schipp S-89). U. williamsii Steyermark, Ceiba

1:126. 1950.

Damp, grassy flats, 1,150-2,100 m., Zacapa. British Honduras

(in swampy, sandy places at sea level); Mexico; Honduras south

to Argentina.

Terrestrial herbs, the stolons slender, sparsely branched, bearing a few globose

traps; leaves few, petiolate, the blades obovate or spathulate, 2-4 mm. long (in

ours), rounded at the apex; scape erect, almost filiform, mostly 5-15 (25) cm. tall,

glabrous, bearing 2-6 flowers; bracts sessile, lanceolate or linear-lanceolate, acute,

about 0.6 mm. long; bracteoles similar to bracts but smaller; pedicels 3-20 mm.

long; calyx lips ovate, obtuse or subacute, minutely glandular-pubescent, often

densely so; corolla often minutely glandular-pubescent, usually white, sometimes

pink or mauve, quite variable in size and shape, in ours the limb 2-3 mm. broad,

the spur slender, more or less acute, shorter than or twice as long as the lower lip.

Utricularia fimbriata HBK. Nova Gen. & Sp. 2: 225. 1818;

P. Taylor, Kew Bull. 18: 72. 1964.

Wet, sandy soil, usually in pine forest, at or little above sea level;

British Honduras. Florida; Cuba; tropical South America; tropical

Africa.

Terrestrial herbs, the rhizoids and stolons few, branching; traps ovoid, ap-

pendaged, borne on rhizoids, stolons, and leaf petioles; leaves few or numerous,

linear-filiform, to 15 mm. long, commonly fugacious; scape 5-25 cm. long, simple,

filiform, wiry, glabrous, bearing 1-5 flowers and numerous peltate, pectinate scales;

bracts broadly ovate, 1-1.5 mm. long, deeply pectinate-dentate; bracteoles almost

twice as large as the bracts, deeply pectinate-dentate; pedicels about 1 mm. long

(in ours); calyx lips unequal, more or less broadly ovate, 2-4 mm. long, deeply

fimbriate-pectinate; corolla yellow to deep orange, mostly 5-9 mm. long, the upper

lip rounded, truncate, or emarginate, the lower lip 3-5 mm. long, entire or some-

times obscurely and irregularly lobate, the spur conical, usually 2-3 mm. wide,

almost parallel with and about as long as the lower lip; capsule globose, about

1.5 mm. in diameter.

Although the pedicels of all collections examined from British

Honduras were about 1 mm. long, one specimen from Venezuela

(Maguire, Wurdack & Bunting 36062-A, determined by Taylor) has

pedicels 2-10 mm. long.

FIG. 70. Utricularia foliosa. A, habit of plant, X ^; B, flower, X 5; C,

stamens and pistil, X 6; D, flower with upper lip of corolla raised to show spur, X

5; E, fruit and calyx, X 5; F, fruiting pedicel with young capsule enclosed in

calyx, and showing bract at base of pedicel, X 7; G, two views of stigma, X13;

H, section of leaf segments bearing traps, X 5.

323

324 FIELDIANA: BOTANY, VOLUME 24

Utricularia foliosa L. Sp. PL 18. 1753; P. Taylor, Kew Bull. 18:

174. 1964. U. oligosperma St. Hil. Voy. Distr. Diam. 2: 427. 1833.

U. mixta Barnh. Mem. Torr. Bot. Club 16: 111. 1920.

Floating in shallow water, margins of lakes and swamps, some-

times trailing over muddy shores where water has receded, 470-1,600

m.; Chiquimula; Izabal; Jalapa; Jutiapa. Florida; Mexico; British

Honduras (at or near sea level); Honduras; Costa Rica; Nicaragua;

Panama; West Indies; South America (to Argentina) ; tropical Africa.

Aquatic herbs, all parts except the inflorescences submerged, with robust,

branching stolons sometimes several meters long; leaves alternate on the stolon

branches, multiple-pinnate, to 15 cm. long, bearing conspicuous, broadly ovoid

traps 1-2 mm. long, upper lip of trap naked or with 2 short, simple or sparsely

branched appendages; scapes erect, simple, stout, 1-3(4) mm. in diameter, glabrous,

bearing 5-20 flowers; bracts broadly ovate, 2-4 mm. long, truncate or short-

acuminate; bracteoles absent; pedicels stout, 4-15 mm. long, usually recurved in

age; calyx about 4 mm. long, the lips broadly ovate, subequal; corolla yellow, 9-15

mm. long, the upper lip entire, the lower one entire or emarginate, the spur com-

monly 5-6 mm. long, shorter than or about equalling and nearly parallel with the

lower lip; capsule globose, indehiscent, 6-8 mm. in diameter, containing 4-12

winged seeds.

Utricularia gibba L. Sp. PL 18. 1753; P. Taylor, Kew Bull. 18:

197. 1964. U. obtusa Swartz, Prodr. Veg. Ind. Occ. 14. 1788. U.

pumila Walter, Fl. Carol. 64. 1788. U. fibrosa Walter, I.e. U.

parkeriana A. DC. in DC. Prodr. 8: 9. 1844. U. secunda Benj.

Linnaea 20: 308. 1847. U. emarginata Benj. I.e. 489.

Floating in shallow water, margins of lakes, pools, and swamps,

or sometimes trailing in mud where water has receded, often mixed

with other vegetation, near sea level to 2,000 m.; Alta Verapaz;

Huehuetenango ; Izabal; Jalapa; Jutiapa; Pete"n; Santa Rosa. East-

ern United States; Mexico; British Honduras; Honduras; El Salva-

dor; Costa Rica; West Indies; South America to Argentina; Africa

(from Nigeria to Transvaal, fide Taylor).

Aquatic herbs with all parts submerged except the inflorescences, the stolons

fasciculate at the base of the scape, filiform, branching, often forming dense mats;

leaves filiform, alternate on the stolons, simple or sparsely divided or forked,

bearing numerous ovoid traps 1-1.5 mm. long, the upper lip of each trap with 2

long, often branching appendages; scapes erect, filiform, glabrous, 2-15 (35) cm.

tall, bearing 1-6 flowers; bracts sessile, 0.5-1 mm. long, entire or obscurely dentate;

bracteoles absent; pedicels 2-15 (30) mm. long; calyx lips 1-3 mm. long, subequal,

broadly ovate, rounded, entire; corolla yellow, commonly 6-12 mm. long (in ours),

the lips nearly equal or the lower lip smaller, variable in contour and apices, but

the lower lip (in ours) usually rounded or truncate; spur variable, conical or

slender, shorter or much longer than the lower lip; capsule globose, 2-4 mm. in

diameter; seeds more or less winged.

GIBSON: FLORA OF GUATEMALA 325

Utricularia guyanensis A. DC. in DC. Prodr. 8: 11. 1844. U.

peckii Blake, Contr. Gray Herb. 52: 90. 1917 (type from British

Honduras, Peck 371). U. rubricaulis Tutin, Journ. Bot. 72: 312,

/. 5. 1934.

In wet, sandy soil, at or little above sea level, British Honduras.

Trinidad; British Guiana; Surinam.

Terrestrial herbs; rhizoids and stolons branched, bearing globose traps; leaves

linear, fugacious; scapes erect, simple, glabrous, 6-20 cm. tall, mostly 4-8 (12)-

flowered; bracts ovate, rounded or acute, about 1 mm. long; bracteoles smaller,

lanceolate, acute; pedicels about 1 mm. long; calyx lips subequal, 1-2 mm. long,

ovate, the upper one rounded or subacute, the lower one sometimes subtruncate;

corolla yellow or orange, 4-5 mm. long, both lips entire, the lower one about 3 mm.

long, the spur conical, saccate, about 1 mm. long; filaments of stamens geniculate

in the middle; style about as long as the ovary; capsule ovoid, 1.5-2.5 mm. long;

seeds more or less globose.

Utricularia hispida Lamarck, Encycl. 1: 50. 1791. U. angusti-

folia Benj. Linnaea 20: 311. 1847. U. macerrima Blake, Contr.

Gray Herb. 52: 89. 1917 (type from British Honduras, Peck 222}.

Damp or wet savannas, 150 m.; Alta Verapaz (near Cerro

Chinaja, Steyermark 45706). British Honduras (near sea level,

usually in pine forest) ; Nicaragua; Trinidad ; northern South America.

Terrestrial herbs, the capillary stolons bearing ovoid traps about 1 mm. long;

leaves few, fugacious, variable, the blades narrowly linear to obovate-spathulate,

said to attain 20 cm. in length; scapes erect, slender, glabrous, 15-50 (70) cm. tall;

bracts lance-ovate, acuminate, laciniate-dentate, 1-2 mm. long, the bracteoles

narrowly linear-lanceolate, shorter; pedicels usually 2-3, ascending, 2-15 mm. long;

calyx lips ovate, unequal, 2-4 mm. long; corolla purple, violet, blue, or white,

8-11 mm. long (in ours), the upper lip ovate to oblong, entire, the lower one more

or less orbicular, entire; the spur slender, acute, 6-7 mm. long, slightly longer than

the lower lip; capsule globose, about 2.5 mm. in diameter, the seeds numerous.

Utricularia hydrocarpa Vahl, Enum. PI. 1: 200. 1827; Taylor,

Mem. N. Y. Bot. Gard. 17: 226. 1967.

Floating in shallow water of swamps or pools, at or little above

sea level ; British Honduras (All Pines, Schipp S-90) . Honduras ; West

Indies; British Guiana; Venezuela.

Aquatic herbs, all parts submerged except the inflorescences, the stolons to

30 cm. long, mostly simple; leaves alternate on the stolons, to 3 cm. long, bipinnate,

the filiform segments bearing ovoid traps 1-3 mm. long; scapes 2-8 cm. long,

filiform, glabrous, bearing 2-5 flowers, bracts sessile, ovate, about 2 mm. long;

bracteoles absent; pedicels 2-13 mm. long, reflexed in fruit; calyx lips subequal,

ovate, 2-3 mm. long; corolla pink, reddish, or deep rose (in ours), 5-8 mm. long,

the lips orbicular, entire, the spur broadly conical, shorter than the lower lip;

capsule globose.

326 FIELDIANA: BOTANY, VOLUME 24

Utricularia jamesoniana Oliver, Journ. Linn. Soc. 4: 169. 1860.

U. verapazensis Morong ex Donn.-Sm. Bot. Gaz. 18: 209. 1893.

U. concinna N. E. Browne, Trans. Linn. Soc. II. 6: 55. 1901.

Known in Guatemala only from the type of U. verapazensis, Alta

Verapaz, near San Pedro Carcha, 900 m., Tuerckheim 1109. Costa

Rica; South America.

Epiphytic plants growing on tree trunks among ferns, mosses, and other small

plants, the filiform stolons bearing traps up to 1 mm. long; leaves persistent,

slender-petiolate, the blades to 1.5 cm. long (in ours), obovate-spathulate to

oblanceolate, obtuse, entire; scape slender, erect, 1-3-flowered, 3-15 cm. tall,

glabrous, bearing 1-several small subulate scales; bract at the base of the pedicel

linear-lanceolate, obtuse, sessile, about 5 mm. long, the 2 bracteoles narrower and

slightly shorter; pedicels capillary, 5-15 mm. long; calyx deeply bilobate, the lobes

subequal, herbaceous, 7-8 mm. long at anthesis, in fruit often 10-12 mm. long and

7-9 mm. wide, broadly cordate-ovate, very obtuse or rounded at the apex, entire;

corolla pale greenish purple, about 1 cm. long, the upper lip broadly rounded at

the apex, the lower lip much larger, trilobate, the middle lobe emarginate, the lips

slightly pubescent on and near the margins; spur upcurved, acute, longer than

the lower lip, minutely glandular-pubescent.

Utricularia juncea Vahl, Enum. PL 1: 202. 1804; P. Taylor,

Mem. N. Y. Bot. Gard. 17: 215. 1967. U. virgatula Barnh. Bull.

Torr. Bot. Club 34: 580. 1908. U. juncea f. minima Blake, Contr.

Gray Herb. 52: 89. 1917 (type from Toledo, British Honduras,

M. E. Peck 502a). U. juncea f. virgatula Fern. Rhodora 46 : 60. 1946.

In bogs or wet margins of swamps or ponds, 1,500-1,900 m.;

Jalapa. (In British Honduras, in wet, sandy or swampy places at

or near sea level, often in pine forest.) Eastern United States; British

Honduras; West Indies; northern South America.

Terrestrial herbs with slender, branching rhizoids and stolons; traps globose,

borne on rhizoids and leaves; leaves fugacious, narrowly linear, said to attain

20 mm. in length; scapes erect, simple, glabrous, commonly 10-40 cm. tall, bearing

1-6 (12) flowers; bracts ovate, acute, 1-2 mm. long, the bracteoles narrowly

lanceolate, shorter; pedicels scarcely exceeding the bracts; calyx often yellowish,

3-4 mm. long at anthesis, 5-7 mm. in fruit, the lips more or less plicate, at least

the upper one prominently venose, ovate, acute or acuminate, the upper one entire,

the lower one entire or bifid; corolla yellow, 10-12 (15) mm. long (in the small form

only 3-8 mm.), the upper lip nearly orbicular, truncate or nearly so, the lower lip

entire, the spur descending, 5-8 mm. long (2-3 mm. in the small form); capsule

ovoid, 2-3 mm. long, invested in the persistent calyx.

One collection from British Honduras, Schipp 633, with scapes

20-40 cm. long, has small flowers, about 8 mm. long.

Utricularia purpurea Walter, Fl. Carol. 64. 1788.

GIBSON: FLORA OF GUATEMALA 327

Floating in shallow water, lakes and swamps at or little above

sea level; British Honduras. Canada and eastern United States to

Florida; West Indies.

Aquatic herbs, all parts submerged except the inflorescences, the stolons

slender, sometimes as much as a meter long; leaves whorled on the stolons, long-

petiolate, the segments filiform, decompound, bearing numerous ovoid traps 1-3

mm. long; scapes commonly 5-15 (30) cm. long,, erect, slender, sometimes swollen

and spongy near the base, bearing (1) 2-5 flowers near the apex; bracts suborbicu-

lar, about 1 mm. long; bracteoles absent; pedicels ascending, 6-8 mm. long or as

much as 15 mm. long in fruit; calyx lips subequal, broadly ovate, usually rounded,

2-3 mm. long; corolla pale purple to bluish purple (in ours), 8-15 mm. broad, the

upper lip entire or somewhat undulate-crenate, the lower one 8-12 mm. long,

trilobate, with a yellow spot at the base; spur conical, shorter than the lower lip

and more or less appressed to it; capsule ovoid to subglobose, 2-3 mm. in diameter.

Utricularia resupinata B. D. Greene in Bigelow, Fl. Bost. ed.

3: 10. 1840. U. spruceana Benth. in Oliver, Journ. Linn. Soc. 4:

173. 1860.

In swampy or boggy ground, at or near sea level; British Hon-

duras (All Pines). Southeastern Canada to Florida; Venezuela

and Brazil.

Plants terrestrial, or sometimes rooting in the bottoms of shallow pools, with

slender horizontal, creeping stolons; leaves alternate, tri-parted, the middle lobe

linear, the lateral lobes capillary, rootlike, bearing minute traps, these apparently

without appendages; scape erect, almost capillary, to 20 cm. tall, bearing a single

flower, the pedicel elongating, subtended by a single, cupular bract 1-2 mm. long;

bracteoles absent; calyx lobes subequal, about 2 mm. long, appressed to the mature

capsule; corolla rose or purplish, 8-12 mm. long, the upper lip narrowly oblong-

spathulate, the lower lip spreading, entire; spur conic-cylmdric, obtuse, the tip

distant from the lower lip and turned upward ; capsule globose, 3-4 mm. in diameter,

containing numerous seeds.

Utricularia subulata L. Sp. PL 18. 1753; P. Taylor, Kew Bull.

18(1) : 81. 1964. U. subulata var. cleistogama A. Gray, Syn. Fl. 2(1) :

317. 1878. U. cleistogama Britton, Trans. N. Y. Acad. Sci. 9: 12.

1889. U. subulata L cleistogama Fernald, Rhodora 23: 291. 1921.

Wet meadows, swamps, bogs, often in pine or pine-oak forest,

near sea level to 1,500 m.; Alta Verapaz; Chiquimula; Izabal; Jalapa;

Jutiapa; Zacapa. Eastern and southern North America (Nova

Scotia to Florida and Texas) ; Mexico; British Honduras, El Salvador,

Honduras to Panama; West Indies; northern South America; Africa;

Asia.

Terrestrial plants with few stolons; leaves fugacious, petiolate, the blades

linear, 1-2 cm. long; scape erect, filiform, wiry, usually fractiflex, mostly 5-20 cm.

328 FTELDIANA: BOTANY, VOLUME 24

tall, simple or branching; bracts peltate, entire; bracteoles absent; flowers com-

monly 2-25, rarely only one, the pedicels mostly 2-6(10) mm. long, axillary in the

bracts; scales few, peltate; calyx lobes subequal, broadly ovate, at anthesis about

1 mm. long, somewhat longer in fruit; corolla yellow or orange-yellow (or said to be

sometimes whitish in cleistogamous flowers), 6-12 mm. long (cleistogamous flowers

about 2 mm.); the upper lip ovate, obtuse, the lower one trilobate; spur conic,

parallel with and about equalling or longer than the lower lip (in cleistogamous

flowers more or less saccate) ; capsule globose, 1-2 mm. in diameter; seeds numerous

more or less ovoid, striate.

Small plants and the cleistogamous form of U. subulata are often

confused with U. pusilla Vahl, said to be a shorter, stouter plant

with fertile inflorescence bracts alternating with sterile ones, shorter

pedicels, and with the spur twice as long as the lower lip. Although

Taylor states that U. pusilla occurs throughout Central America, I

have seen no material from Guatemala. Some of our plants do have

spurs almost twice as long as the lip, but in every case the bracts

are all fertile, the pedicel length is quite variable, and I cannot see

that these forms are any stouter than U. subulata.

ACANTHACEAE. Acanthus Family.

DOROTHY NASH GIBSON

References: G. Lindau, Acanthaceae, in Engler & Prantl, Pflan-

zenf. 4 (3b) : 274-354. 1895. C. E. B. Bremekamp, The Delimitation

of the Acanthaceae, Proc. Kon. Ned. Akad. v. Wetensch. Ser. C, 56:

533-546. 1953; Delimitation and Subdivision of the Acanthaceae,

Bull. Bot. Surv. India 7: 21-30. 1965. E. C. Leonard, The Acan-

thaceae of the Yucatan Peninsula, Carnegie Inst. Wash. 461: 193-

238. 1936; The Acanthaceae of Colombia, Contr. U. S. Nat. Herb.

31: 1-781. 1951-1958. Wm. F. Grant, A Cytogenetic Study in the

Acanthaceae, Brittonia 8: 121-149. 1955. Bhoj Raj, Pollen Mor-

phological Studies in the Acanthaceae, Grana Palynol. 3(1) : 3-108.

1961. Bruno Petriella, El Polen de las Acanthaceae Argentinas,

Rev. Mus. La Plata, Bot. 53 (11) : 51-68. 1968. R. W. Long, The

Genera of Acanthaceae in the Southeastern United States, Journ.

Am. Arb. 51(3) : 257-309. 1970.

Herbaceous or suffrutescent plants, rarely shrubs or small trees, scandent only

in Mendoncia and Thnnbergia, glabrous or pubescent, the pubescence almost

always of simple, multicellular hairs, often glandular-viscid, the leaves and other

parts often with conspicuous cystoliths, these appearing as fine lines on the leaf

surfaces and sometimes mistaken for strigose pubescence; leaves usually opposite,

petiolate, and entire, often somewhat undulate or undulate-dentate, rarely dentate

or lobate; stipules none; inflorescences axillary or terminal, often spicate, racemose,

GIBSON: FLORA OF GUATEMALA 329

or paniculate, sometimes cymose, the cymes sometimes fasciculate, the flowers

rarely solitary; flowers usually bracteate and bibracteolate; primary bracts or

bract-like floral leaves often broad and imbricate; calyx segments usually 5 or by

abortion of the posterior one only 4, free or more or less connate, imbricate, valvate,

or often separated long before anthesis, calyx truncate or 10-15-dentate in Thun-

bergia, annular or cupular in Mendoncia; corolla gamopetalous, often funnelform,

the tube usually well developed, cylindric or ventricose or slender-tubular below

and abruptly dilated into a funnelform throat, the limb usually more or less

bilabiate, sometimes the 5 lobes spreading and appearing nearly regular, but usually

the posterior lip erect or suberect, entire or bifid, rarely fimbriate, the anterior lip

more spreading, trifid or trilobate or rarely subentire, the lobes imbricate in bud or

contorted; perfect stamens 4 and didynamous, or 2, inserted on the tube of the

corolla, included or ascending below the posterior lip, rarely exserted; staminodes

often present; anthers usually dorsifixed, commonly with 2 distinct thecae, some-

times only one, dehiscent by a longitudinal slit, sometimes parallel and inserted

at the same height on the connective, or often unequal and affixed at different

heights on the connective, frequently calcarate at the base, one sometimes abortive

or vestigial; hypogynous disc various in form, often obscure; ovary superior, sessile

on the disc, bilocular; style simple, usually filiform, the apex usually bilobate, but

one lobe often minute or abortive; stigmas minute, terminal or short-decurrent

dorsally; ovules anatropous or shortly amphitropous, placentation usually axile

(parietal in Elytraria); fruit usually capsular, very rarely baccate or drupaceous,

the capsules usually clavate, sometimes oblong or nearly linear, terete or com-

pressed, usually contracted below into a stout stipe, elastically dehiscent from the

apex to the base, the valves becoming recurved; seeds arising at maturity from

the thickened hard funicles, these (called retinacula) incurved-ascending; seeds as

many as the ovules or by abortion fewer, usually lenticular or discoid, sometimes

nearly globose, the testa often rugose or muricate, sometimes minutely glochidiate,

commonly mucilaginous when wetted; endosperm usually none; embryo com-

pressed.

The Acanthaceae are a large pantropical, rarely temperate,

family with about 200 genera, many of which are monotypic. Most

species are found in dense, wet forest but a few genera are charac-

teristic of open places and some are common weeds of waste and

cultivated ground. Many have large and handsome flowers but the

flowers are often produced only in small numbers and the corollas

frequently fall if the plants are handled.

I agree with Bremekamp and Raj that the Mendoncioideae,

Nelsonioideae, and Thunbergioideae should properly be raised to

family rank, but they are included here, following traditional usage

as a matter of convenience.

The taxonomy is still unsatisfactory, perhaps because classifica-

tion of the family has been based primarily on the disposition of the

thecae on the connective, and on pollen characters.

330 FIELDIANA: BOTANY, VOLUME 24

Plants of the genus Acanthus, native of the Mediterranean region,

are planted in the Jardin Botanico of Guatemala and perhaps

elsewhere. They differ from all Central American members of this

family in having sinuate-pinnatifid leaves, the teeth of the leaves

often with spinose tips.

A few other genera besides the 36 treated here have been dis-

covered in southern Central America.

Fruits drupaceous; plants always scandent Mendoncia.

Fruits capsular; plants not scandent (except in Thunbergia, an introduced genus).

Calyx spathelike and bilobate at anthesis Spathacanthus.

Calyx neither spathelike nor bilobate at anthesis.

Plants often scandent; calyx truncate or 10-15-dentate Thunbergia.

Plants not scandent; calyx 3-5-lobate.

Calyx segments 3: corolla oblique, gibbous-campanulate in throat.

Louteridium.

Calyx segments 4 or 5; corolla not as above.

Fertile stamens 4 (in ours).

Anthers monothecous.

Cystpliths absent; corollas red or yellow; filaments of stamens

inserted near base of corolla tube Aphelandra.

Cystoliths present; corollas purple, pink, or white; filaments of

stamens inserted near or above middle of corolla tube.

Stenandrium.

Anthers bithecous.

Calyx segments 4; bracts of inflorescence conspicuously secund.

Lophostachys.

Calyx segments 5; bracts of inflorescence not secund.

Plants small trees or large shrubs; corolla tube ampliate from

almost the base, or abruptly dilated shortly above the base.

Bravaisia.

Plants usually herbaceous, rarely suffrutescent; corolla tube not

as above.

Corolla conspicuously bilabiate; capsule bilocular from base.

Inflorescence spicate, terminal or axillary; thecae discrete.

Teliostachya.

Inflorescence fasciculate in leaf axils; thecae parallel.

Hygrophila.

Corolla appearing nearly regular, the limb spreading (in ours);

capsule bilocular from a point above the base, the basal

portion solid for at least 2 mm., either contracted or

stipiform.

Flowers in dense spikes bearing large, broad, imbricate bracts.

Leaf margins entire; placentae breaking away in mature

capsule Blechum.

Leaf margins crenate; placentae not breaking away in

mature capsule Hemigraphis.

Flowers not in dense spikes, or if spicate, not bearing large,

broad bracts.

GIBSON: FLORA OF GUATEMALA 331

Calyx segments connate for one-half to one-quarter their

length; thecae calcarate at base Dyschoriste.

Calyx segments parted nearly to base; thecae muticous at

base Ruellia.

Fertile stamens 2 (in ours).

Anthers commonly monothecous, or if 2 thecae, one always much

smaller than the other, the smaller one usually abortive, vestigial,

often sterile.

Corolla less than 5 mm. long; staminodes present Buceragenia.

Corolla more than 5 mm. long; staminodes absent.

Inflorescence short and dense, headlike to spikelike, with bracts,

bracteoles, and calyx segments rigid, filiform or oblanceolate

and cuspidate; calyx segments 4 Chaetothylax.

Inflorescence not as above; calyx segments 5.

Inflorescence an elongated, spikelike thyrse or narrow panicle;

corolla tube gradually ampliate from base Razisea.

Inflorescence a broad, often lax panicle; corolla tube usually

more or less saccate near base.

Anterior corolla lip shallowly trifid and appearing almost

truncate Hansteinia.

Anterior corolla lip trilobate, more or less spreading or erect.

Habracanthus.

Anthers bithecous, the thecae about the same size.

Bracts subtending the flowers united almost to the apex to form an

involucre.

Involucre formed by outermost bracts, these very large, 4.5-6.5

cm. long, bluish purple, acute to acuminate; corolla 7-7.5 cm.

long Neohallia.

Involucre formed by innermost bracts, these less than 1.5 cm.

long, green, mucronate; corolla 1-1.5 cm. long Henrya.

Bracts subtending the flowers not united to form an involucre, or

merely partially connate at base only, as in Dicliptera.

Calyx cupular, the tube equalling or longer'than the lobes.

Trybliocalyx.

Calyx not as above.

Leaves alternate or subopposite; cystoliths absent; seeds 12-20,

borne on papilliform retinacula Elytraria.

Leaves opposite; cystoliths present; seeds 8 or fewer, usually 4

or 2, borne on hook-like retinacula.

Outer bracts of inflorescence conspicuously 4-ranked.

Tetramerium.

Outer bracts of inflorescence not conspicuously 4-ranked.

Corolla appearing nearly regular, the lobes free and more

or less spreading.

Corolla lobes 5.

Stamens included ; thecae muticous or acute, one below

the other on the connective; corolla tube usually

straight and narrow; native plants.

Pseuderanthemum.

Stamens exserted; thecae conspicuously calcarate, at-

tached at apex of the connective; corolla tube

332 FIELDIANA: BOTANY, VOLUME 24

long-cylindric, often somewhat ventricose above;

cultivated plants Sanchezia.

Corolla lobes 4.

Bracts of inflorescence broad, foliaceous, imbricate,

venose Averia,

Bracts of inflorescence very narrow, neither imbricate

nor venose.

Flowers in lax spikes or racemes, these often forming

large, terminal panicles; calyx segments 5;

corolla tube (in ours) shorter than the lobes.

Carlowrightia.

Flowers usually 1-3 in a leaf axil; calyx segments 4

(in ours); corolla tube elongated, very narrow,

much longer than the lobes Siphonoglossa.

Corolla somewhat bilabiate to conspicuously ringent.

Staminodes present.

Bracts small, usually inconspicuous; calyx segments 5,

entire, uniformly narrow Odontonema.

Bracts large and conspicuous; calyx segments 4,

spinose-ciliate, the posterior and anterior segments

broad, the lateral ones narrow Barleria.

Staminodes absent.

Corolla divided nearly to the base; thecae both

attached at apex of connective Megalostoma.

Corolla not divided nearly to base; thecae more or less

unequally inserted on the connective.

Stems usually 6-angled; bracts subtending the calyx

partially united at base; calyx usually hyaline;

placentae separating elastically from their walls

in mature capsule Dicliptera.

Stems not 6-angled; bracts not united; calyx not

hyaline; placentae not separating from walls in

mature capsule.

Leaves of each pair unequal (in ours), one distinct-

ly smaller than the other; pollen multiporate,

the sexine composed of densely spaced, poly-

gonal insulae Poikilacanthus.

Leaves of each pair equal or nearly so; pollen not

as above.

Bracts and bracteoles filiform-caudate; pollen

spheroidal or subspheroidal, 3-colpate, trema

area lacking rows of insulae.

Streblacanthus.

Bracts and bracteoles not filiform-caudate; pol-

len ellipsoidal to prolate, usually 2-porate,

sometimes 3-4-porate, trema area always

provided with one or more rows of insulae.

Justida.

APHELANDRA R. Brown

Erect shrubs or herbs; leaves opposite, entire or sometimes sinuate-lobate or

spinose-dentate; inflorescences terminal, spicate, simple or branched, the flowers

GIBSON: FLORA OF GUATEMALA 333

large, solitary and sessile within green or colored, entire or dentate bracts; bracts

imbricate, usually longer than the calyx, ovate to lanceolate, coriaceous, herbace-

ous, or membranaceous; bracteoles usually similar to the calyx segments but often

smaller; calyx segments 5, usually narrow, subequal, or the posterior one larger;

corolla yellow, orange, or red, the tube straight or incurved, sometimes ampliate

above, the limb bilabiate, the posterior lip innermost in bud, erect, concave,

subentire or shallowly bilobate and spreading, the anterior lip outermost in bud,

reflexed-spreading, the lateral lobes united with the middle one to form a trilobate

lip, or small and arising from the sides of the lower lip; stamens 4, the filaments

inserted near the base of the corolla tube, short-exserted or shorter than the lip;

anthers connivent in pairs, monothecous, dorsifixed, sometimes minutely bearded

at the apex, muticous; pollen 3-colpate, prolate to perprolate; disc inconspicuous;

style obtuse at the apex or shallowly bilobate; ovules 2 in each locule; capsule

oblong, shortly contracted at the base; seeds 4 or by abortion fewer, compressed,

suborbicular.

About 75 species have been reported, widely dispersed in tropical

America, with six in Guatemala. A seventh, A. micans Moritz, a

species ranging from Costa Rica to northern South America, was

reported from Guatemala by Leonard in the Flora of Costa Rica,

but we have seen no collection from Guatemala.

Bracts of the inflorescence entire.

Bracts glabrous A. speciosa.

Bracts more or less pubescent.

Spikes very dense, short, mostly 2-8 cm. long; bracts pubescent with mostly

appressed, eglandular, or only minutely and inconspicuously glandular

hairs A. schiedeana.

Spikes lax, the rachis usually visible between the lower flowers, mostly 10-30

cm. long; bracts densely viscid-pilose with more or less spreading hairs.

A. schiedeana var. giganti flora.

Bracts of the inflorescence dentate.

Plants herbaceous or suffrutescent, usually simple; bracts pectinate-dentate;

corolla orange or orange-red.

Leaves linear or linear-lanceolate, mostly 1.5-3 cm. wide A. repanda.

Leaves lance-oblong to broadly elliptic, mostly 3.5-8 cm. wide. .A. aurantiaca.

Plants branched shrubs; bracts with 1, 2, or several, large, acute teeth; corolla

usually bright red.

Inflorescences elongated, mostly 6-25 cm. long; bracts rigid, never spreading,

bearing a few conspicuous glands A. deppeana.

Inflorescences short, mostly 2-4 cm. long; at least the lower bracts spreading

or recurved at the apex, all bracts without glands A. heydeana.

Aphelandra aurantiaca (Scheidw.) Lindl. Bot. Reg. 31: t. 12.

1845. Hemisandra aurantiaca Scheidw. Bull. Acad. Brux. 9(1): 22.

1842. A. acutifolia Nees in DC. Prodr. 11 : 299. 1847. Flor de rosario

(Alta Verapaz).

Damp or wet, mixed forest, sea level to 1,300 m., usually at

350 m. or less; Alta Verapaz; Baja Verapaz; Izabal; Huehuetenango ;

FIG. 71. Aphelandra aurantiaca. A, habit, ^ natural size; B, calyx opened

to show ovary, X 3}^; G, flower complete with bract and bracteole, X 2; D,

corolla opened to show stamens and pistil, X 2; E, connivent anthers, X 4; F,

opened capsule, X 3^; G, seed, X 4.

334

GIBSON: FLORA OF GUATEMALA 335

Peten. Southern Mexico; British Honduras, along the Atlantic

coast to Panama; northern South America.

Erect herbs, sometimes suffrutescent, usually unbranched, sometimes a meter

tall but usually half as high, glabrous or nearly so except in the inflorescence;

leaves short-petiolate, the blades lance-oblong to elliptic, mostly 10-30 cm. long

and 3.5-8 cm. wide, acute or acuminate, acute at the base, entire; inflorescences

terminal, the spikes solitary, mostly 6-23 cm. long, very densely flowered; bracts

green, appressed, ovate-lanceolate, acuminate, pectinate-dentate, finely puberulent

or glabrate, 2-3 cm. long and to 1 cm. wide; bracteoles and calyx segments lanceo-

late, puberulent, ciliate, the calyx 8-12 mm. long; corolla yellow-orange to reddish-

orange, puberulent outside, 5-6 cm. long, the upper lip erect, acute, entire, the

lower lip trilobate, the lateral lobes half as long as the middle one; capsule 12-15

mm. long, puberulent, containing 2-4 sparsely pubescent seeds.

These plants vary greatly in size, some of them blooming when

only 15 cm. tall.

Aphelandra deppeana Schlecht. & Cham. Linnaea 5: 96. 1830.

A. pectinata Willd. ex Nees in DC. Prodr. 11: 298. 1847. A. haenke-

ana Nees, I.e. Flor de San Julian (Guatemala) ; sic-chd (Quecchi, Alta

Verapaz); chacanal (Maya, British Honduras).

Wet to dry thickets or forest, often in rocky places, sometimes in

pine and oak forest, sea level to 1,800 m.; Alta Verapaz; Chimalte-

nango; Chiquimula; Escuintla; Guatemala; Huehuetenango ; Izabal;

Jalapa; Jutiapa; Pete"n; Quezaltenango ; El Quiche"; Retalhuleu;

Sacatepe" quez ; Santa Rosa ; Solola ; Suchitepe1 quez ; Zacapa. Southern

Mexico; British Honduras to El Salvador and Panama; West Indies;

northern South America.

Erect, branched shrubs 1-3 m. tall, the branches ochraceous, densely pilose

when young; leaves short-petiolate, the blades ovate-elliptic to lance-oblong,

mostly 6-15 cm. long and 3-7 cm. wide, sometimes larger, acuminate or long-

acuminate, rather abruptly contracted below and long-decurrent almost to the

base of the petiole, entire or nearly so, often scaberulous above, usually densely

and softly pubescent beneath; spikes terminal, solitary or clustered, usually

elongated; bracts green, lance-ovate, 8-15 mm. long, 4-6 mm. wide, acuminate,

with 2-several slender, appressed teeth or laciniations above, densely pubescent to

glabrate, ciliate, usually bearing several large glands; calyx 6-10 mm. long,

pubescent; corolla usually bright red, rarely yellow, 3.5-4 cm. long, pubescent,

the upper lip narrow, erect, bilobate, the lower lip spreading, entire or trilobate,

the lateral lobes very small; capsule 10-15 mm. long, obtuse, glabrous, containing

4 seeds.

Very common shrubs throughout the drier lowlands of Central

America, one of the most abundant plants of dry thickets. The

form with yellow corollas has been collected only once (Escuintla,

Standley 64,076) ; it differs in no other respect from the typical form.

336 FIELDIANA: BOTANY, VOLUME 24

Aphelandra heydeana Donn.-Sm. Bot. Gaz. 18: 210, t. 23.

1893. Flor de San Julidn (Guatemala).

Damp or dry, mixed forest or thickets, 1,000-1,600 m.; Guate-

mala; Santa Rosa (type from Chupadero, Heyde & Lux 4037).

Mexico (Chiapas).

Slender shrubs, 1.5-3 m. tall, the branches slender, pale, appressed-pilose when

young; leaves on petioles 3-10 cm. long, the blades ovate to ovate-lanceolate or

elliptic, mostly 8-20 cm. long and 4-7 cm. wide, acuminate or long-acuminate,

abruptly contracted at the base and narrowly long-decurrent almost to the base of

the petiole, glabrous or nearly so, paler beneath, the margins entire; inflorescences

terminal, the spikes usually solitary, mostly 2-4 cm. long; bracts green, foliaceous,

mostly 15-18 mm. long, obovate-oblong or lanceolate, long-attenuate, venose,

thinly pilose, at least the lower ones spreading or recurved at the apex, the margins

with 1-2 large, acute teeth; sepals 10-14 mm. long, narrowly lanceolate-acuminate;

corolla usually bright red, rarely yellow and red, 5.5-7 cm. long, glandular-pubes-

cent, the posterior lip erect, oblong, about 2 cm. long, entire, the anterior lip

trilobate, the middle lobe broadly obovate-oblong, about 12 mm. wide, much larger

than the lateral lobes; anthers about 6 mm. long; capsule about 1.5 cm. long,

glabrous or nearly so.

Aphelandra repanda Nees in DC. Prodr. 11: 728. 1847. A.

aurantiaca var. stenophylla Standl. Field Mus. Bot. 4: 324. 1929

(type from Lancetilla Valley, Honduras).

Wet, mixed lowland forest, at or little above sea level; Izabal.

British Honduras; Honduras; Costa Rica; western South America.

Herbaceous or suffrutescent plants, simple, to 60 cm. tall, glabrous or nearly

so except for the inflorescence; leaves short-petiolate, the blades linear-lanceolate,

mostly 10-25 cm. long and 1.5-3 cm. wide, long-acuminate, attenuate to the base,

slightly paler beneath, often lustrous above, the margins more or less undulate;

inflorescences terminal, the spikes solitary, sessile, very densely flowered, 3-15 cm.

long; bracts ovate-oblong, 2-3 cm. long, acuminate, pectinate-dentate, green,

minutely puberulent, usually ciliolate; calyx segments 8-12 mm. long, minutely

puberulent; corolla orange-red, 3.5-5 cm. long, the upper lip erect, entire, the lower

lip trilobate, the lobes subequal, the middle lobe oval, the lateral ones oblong-

lanceolate; capsule about 1 cm. long, minutely puberulent.

Although these plants are usually found in wet, lowland forest,

they have been collected at 1,300 m. in Costa Rica.

Aphelandra schiedeana Schlecht. & Cham. Linnaea 5: 95. 1830.

Damp, mixed mountain forest or thickets, 1,000-2,800 m.;

Chimaltenango; Escuintla; Quezaltenango; El Quiche"; Sacatepe"quez ;

San Marcos. Southern Mexico; El Salvador.

Rather slender shrubs, 1-3.5 m. tall, sometimes weak and reclining, the

branches appressed-pilose or glabrate; leaves on slender petioles, the blades

GIBSON: FLORA OF GUATEMALA 337

narrowly lanceolate to oblong-lanceolate, mostly 8-20 cm. long, narrowly long-

acuminate, long-attenuate to the base, sparsely short-pilose above or glabrate,

more or less short-pilose beneath, chiefly along the costa, or almost wholly glabrous;

inflorescences terminal, the spikes short and densely flowered, 2-8 cm. long; bracts

ovate to broadly elliptical, 2-3 cm. long, obtuse or acute, rarely acuminate, or the

lowermost ones rarely more or less cuspidate, usually sparsely or densely appressed-

pilose with glandular and eglandular hairs, rarely almost glabrous, usually red in

part or throughout, thin, entire; calyx segments linear-lanceolate, pale, 10-15 mm.

long, pilosulous or sometimes glandular-pubescent near the apex; corolla usually

bright red, rarely yellow, pubescent outside, 5-7 cm. long, the posterior lip about

2 cm. long, acute, the lower lip deeply trilobate, the segments lanceolate or the

middle one cuneate-oblanceolate to obovate; anthers about 4 mm. long; capsule

about 2 cm. long, oblong-ellipsoid, obtuse or subacute, lustrous, puberulent.

A. schiedeana var. gigantiflora (Lindau) D. Gibson, Fieldiana:

Botany 34: 57. 1973. A. gigantiflora Lindau, Bull. Herb. Boiss. 3

369. 1895 (type collected by Warscewicz in Costa Rica or Guatemala,

probably the latter). A. padillana Standley, Journ. Wash. Acad.

Sci. 14: 244. 1924 (type from El Salvador). A. gigantiflora f. lutea

Standl. & Steyerm. Field Mus. Bot. 23: 237. 1947 (type from Escuint-

la, Standley 89560).

Damp forest and thickets, 700-1,200 m.; Escuintla; Sacatep^-

quez; Chimaltenango. El Salvador; Costa Rica (?).

Differs from A. schiedeana only in its elongated inflorescences

(10-30 cm. long), more laxly flowered on lower portion, and the more

conspicuously glandular pubescence of rachis and bracts.

Aphelandra speciosa Brandegee, Univ. Calif. Publ. Bot. 6:

196. 1915. Flor de mayo (San Marcos) ; flor de la Santa Cruz de Mayo

(Quezaltenango).

Dense, wet, mixed mountain forest, 900-2,000 m.; Quezaltenango;

San Marcos; Suchitepe"quez. Mexico (Chiapas).

Shrubs 1-4 5 m. tall, usually sparsely branched, glabrous throughout or nearly

so; leaves on petioles 2-15 cm. long, the blades ovate to elliptic-oblong, mostly

15-30 cm. long and 5-12 cm. wide, acuminate or abruptly long-acuminate, acute

or attenuate to the base, paler beneath; inflorescences terminal, the spikes usually

solitary, mostly 10-25 cm. long; bracts deep red, ovate or oblong-ovate, commonly

3-4 cm. long, sometimes to 5 cm. long, 1.5-2.8 cm. wide, obtuse and often cuspidu-

late, entire; calyx segments lanceolate, 7-14 mm. long; corolla 4-6 cm. long, bright

red, the tube 2.5-3.5 cm. long, the posterior lip entire, the anterior one trilobate

to the base, the lateral lobes much smaller than the middle one; capsule about 2 cm.

long; seeds ovate, minutely tuber culate.

This beautiful plant is certainly the showiest of all the local

species.

m

FIG. 72. Aflm'a longipes. A, habit, X J^; B, flower with bracteoles, X 6; G,

calyx and bracteoles, X 6; D, opened capsule, X 9; E, papillate surface of seed,

greatly enlarged; F, capsule opened to show retinacula, X 9.

338

GIBSON: FLORA OF GUATEMALA 339

AVERIA Leonard

Erect or ascending, perennial herbs or low shrubs with pale stems, branching,

viscid-pubescent; leaves often deciduous, petiolate, the blades broad, entire;

inflorescences terminal or lateral, spicate; the bracts broad, foliaceous, trinerved to

venose, imbricate when young, later spreading and becoming more lax; bracteoles

narrowly lanceolate; calyx deeply cleft, the 5 segments subulate; corolla tube

slender, shorter than or about equalling the lobes, scarcely ampliate above, the

limb appearing almost regular, the 4 lobes rounded at the apex, spreading; stamens

2, anthers bithecous, thecae parallel, subequal, muticous at the base; pollen 3-

colporate, prolate; capsule broad, stipitate, containing 2-4 muriculate or papillate

seeds.

The genus consists of a single, variable species. It was named for

Mr. Sewell Avery of Chicago, who contributed funds for one of the

Field Museum botanical expeditions to Guatemala.

Averia longipes (Standley) Leonard, Journ. Wash. Acad. Sci.

30: 503. 1940. Drejerella longipes Standley, Field Mus. Bot. 8: 47.

1930 (type from Chichen Itza, Yucatan). A. serrata Leonard, Journ.

Wash. Acad. Sci. 30: 502. /. 1. 1940. A. melanosperma Leonard,

Ceibal: 110. 1950.

Dry thickets or brushy, rocky slopes, sea level to 1,300 m.; El

Progreso; Zacapa; Retalhuleu. Mexico (Chiapas and Yucatan);

Honduras; El Salvador.

Erect or ascending perennials, 20-60 cm. tall, rather densely branched,

herbaceous or suffrutescent, the branches pale, glandular-pilose or in age glabrate;

leaves petiolate, the blades lance-ovate, ovate, or broadly ovate, mostly 0.8-4 cm.

long, 0.4-2 cm. wide, obtuse or subacute, truncate or subcordate at the base,

remotely repand-dentate or entire, densely glandular-pubescent on both surfaces;

inflorescences numerous, the spikes mostly 1-3 cm. long; bracts subtending the

inflorescences foliaceous, petiolate, the blades rhombic or broadly ovate, commonly

4-15 mm. long, 3-10 mm. wide, sometimes larger, acute or obtuse, usually apiculate,

entire or with 1-3 inconspicuous teeth on each side; floral bracts similar but

smaller, nearly sessile or abruptly narrowed at the base into a short petiole,

glandular-pilose; bracteoles linear-lanceolate, 6-7 mm. long, densely glandular-

pilose; calyx segments subulate, 3-4 mm. long, glandular-pilose; corolla cream-

colored, 8-14 mm. long, glabrous outside; anthers purplish; capsule 3-4 mm. long,

glabrous; seeds first white, usually becoming brown or black in age.

The uppermost, younger floral bracts are often lanceolate, the

lower ones broadly ovate; a plant may have all bracts entire, nearly

all dentate, or some bracts entire and a few dentate.

BARLERIA L.

Herbs or shrubs, glabrous or pubescent; leaves entire; flowers often large and

showy, variously arranged but often in terminal, bracteate, densely-flowered

340 FIELDIANA: BOTANY, VOLUME 24

spikes, the bracts entire or serrate, the bracteoles linear; calyx deeply 4-lobate,

the posterior and anterior segments largest, the lateral ones innermost, narrow;

corolla tube usually slender, more or less ampliate above, straight or incurved, the

limb spreading, the 5 lobes imbricate, rounded or ovate, about equal or the anterior,

inner one shorter; fertile stamens 2 (in ours), the 2 posterior ones reduced to

staminodes, the filaments inserted near or a little above the base of the corolla

tube, the anthers oblong-sagittate, dorsifixed, the thecae equal, parallel, muticous;

pollen 3-porate, spheroidal; disc inconspicuous; style somewhat thickened at the

apex, obtuse, entire, or slightly complanate; ovules 2 in each locule; capsule ovoid

or oblong, little contracted at the base, subterete, sometimes contracted above

into a beak, seminiferous below the middle; seeds 4 or by abortion 2, flat, ovate or

orbicular, supported on usually acute retinacula.

Species 60 or more, mostly in Asia and Africa, only a few Ameri-

can. Only one is known from North America.

Barleria micans Nees in Benth. Bot. Voy. Sulph. 146. 1844.

B. discolor Nees, I.e. Barleriopsis glandulosa Oerst. Vid. Medd.

Kjoebenhavn 1854: 134. 1855.

Damp or dry thickets or lowland forest, often in rocky places,

135-1,300 m.; Escuintla; Huehuetenango ; Santa Rosa. Western

and southern Mexico; British Honduras to El Salvador and Panama;

Colombia.

Suffruticose herbs or shrubs, erect, stout, to 1.5 m. tall, simple or branching,

the stems strigose; leaves short-petiolate, the blades ovate or lanceolate, mostly

8-25 cm. long, 2-8 cm. wide, attenuate-acuminate, attenuate to the base and

decurrent on the petiole almost to the base, sparsely strigose or hirsute; inflores-

cences terminal, the spikes very densely flowered, sessile or nearly so, 3-10 cm.

long, 2-3 cm. broad; bracts lance-ovate or broadly ovate, 1-2.5 cm. long, 0.5-0.8

cm. wide, acute to acuminate, entire or nearly so, strigose, prominently hirsute-

ciliate, green, often turning purplish when dried; bracteoles linear, mostly 1.5-2 cm.

long; calyx segments very unequal, the margins of the anterior and posterior ones

more or less dentate, especially from the middle to the apex, spinose-ciliate, the

spines sometimes to 4 mm. long, the anterior segment broadly lanceolate to ovate-

oblong, mostly 1.5-2.5 (3) cm. long, 1-1.4 cm. wide, bidentate at the apex, the

posterior one slightly longer and more narrow, acuminate, cuspidate, the lateral

segments narrowly lanceolate or linear-acuminate, cuspidate, shorter than the

other two; corolla bright yellow when fresh, turning bluish-purple when dried,

4-6 cm. long, the tube slender, the lobes oblanceolate, spreading; fertile stamens

2, exserted, the anthers 4.5-6.5 (8) mm. long; capsule glabrous, about 1.5 cm. long,

containing 2-4 seeds, 4-5 mm. in diameter, minutely apiculate.

Corollas of the Guatemalan plants, as well as those of most

specimens from other countries, are 4-5 cm. long with anthers

4.5-6.5 mm. long. One collection from Jalisco, Mexico (Pringle 4587)

has corollas 6 cm. long, with anthers to 8 mm. long.

FIG. 73. Barleria micans. A, habit, X Vt\ B, corolla opened to show stamens

and staminodes (pistil removed), X 2; C, corolla lobes in bud, X 2; D, flower

with bracts and bracteoles, X 1; E, capsule, X 2; F, seed X 3; G, bracts and

bracteoles, X 2.

341

342 FIELDIANA: BOTANY, VOLUME 24

BLECHUM P. Browne

Perennial herbs, erect or most often decumbent or procumbent, pilose or

glabrate; leaves petiolate, the blades thin, entire; inflorescences densely spicate,

terminal or axillary, the flowers commonly sessile or short-pedicellate in the axils

of foliaceous bracts; bracts green, imbricate, usually ovate to suborbicular, acute

to short-acuminate, ciliate; bracteoles narrowly lanceolate to elliptic; calyx seg-

ments 5, linear-acuminate, subequal, ciliolate; corolla usually lavender, purple, or

white, the tube slender, straight or curved, little ampliate above, the limb spread-

ing, the 5 lobes rounded, contorted in bud, subequal; stamens 4, didynamous,

inserted above the middle of the corolla tube, the filaments connate at the base by

a membrane, the anthers oblong, dorsifixed, the thecae parallel, equal, muticous;

pollen 3-colporate, syncolpate, spheroidal; style subulate at the apex, the posterior

lobe usually minute; ovules few to 6 in each locule; capsule contracted at the base,

nearly ovate or suborbicular, compressed parallel to the septum; seeds flat,

suborbicular, the retinacula acute; placentae breaking away in the mature capsule.

Perhaps six or eight species, in tropical America, one sometimes

naturalized in the Old World tropics. Two occur in Guatemala and

three others have been reported from southern Central America.

The plant from British Honduras, H. H. Bartlett 11355, which

Leonard named Blechum cordatum (Carnegie Inst. Wash. Publ. 461:

200, /. 2. 1936), is Hemigraphis alternate, (Burm. f.) T. Anders, a

native of Malesia and no doubt an escape. It is occasionally planted

in gardens in Mexico and in Honduras.

Bracts densely puberulent or strigillose; corolla 1-2 cm. long, the throat 2-3 mm.

broad B. brownei.

Bracts essentially glabrous; corolla 3-3.5 cm. long, the throat about 1 cm. broad.

B. grandiflorum.

Blechum brownei Juss. Ann. Mus. Hist. Nat. Paris 9: 270.

1807. Ruellia blechum L. Syst. ed. 10: 1120. 1759. Barleria

pyramidata Lam. Encycl. 1: 380. 1785. Blechum pyramidatum

Urban, Repert. Sp. Nov. 15: 323. 1918. B. brownei f. puberulum

Leonard, Journ. Wash. Acad. Sci. 32: 184. 1942. Corrimiento

(Pete"n).

Damp or wet thickets or mixed forest, pastures, waste ground,

sometimes in sand along streams, sometimes in pine forest, sea level

to 1,400 m.; Alta Verapaz; Escuintla; Izabal; Pete"n; Quezaltenango;

Retalhuleu; Santa Rosa; Suchitepe'quez. Florida; Mexico; British

Honduras to El Salvador and Panama; West Indies; South America;

naturalized in the Old World tropics.

Perennial plants, usually decumbent or procumbent and rooting at the lower

nodes, sometimes suberect, usually less than a meter tall, the stems puberulent

or glabrate; leaves slender-petiolate, the blades ovate or lance-ovate, mostly 2-7

FIG. 74. Blechum brownei. A, habit, X l/i; B, flower, showing calyx and

bracts, X 3; G, corolla opened to show stamens, X 4; D, calyx opened to show

ovary with style, X 5; E, capsule, X 2J^; F, placenta with retinacula and seeds,

X 2K; G, leaf, X 5.

343

344 FIELDIANA: BOTANY, VOLUME 24

cm. long, acute or obtuse, obtuse at the base or contracted and short-decurrent on

the petiole, glabrate or sparsely pilose; inflorescences 3-6 cm. long, the bracts

ovate, mostly 1-2 cm. long, obtuse or subacute, venose, densely puberulent or

strigillose, ciliate; calyx 4-6 mm. long, the segments linear-acuminate, long-ciliate;

corolla lavender or white, 1-2 cm. long, pubescent outside, the slender tube little

dilated in the throat, the lobes short, rounded; capsule broadly oblong, 6-7 mm.

long, puberulent.

The Maya name of Yucatan is recorded as "acabxiu."

These are unattractive plants, one of the most abundant weeds

of waste ground throughout the Central American lowlands.

Blechum grandiflorum Oerst. Vid. Medd. Kjoebenhavn 1854:

168. 1855.

Damp or wet, often rocky thickets or in mixed forest, 600-1,300

m.; Alta Verapaz; Baja Verapaz. Mexico (Chiapas); Nicaragua.

Erect plants to about a meter tall, branching, the branches bifariously puberu-

lent or almost glabrous; leaves slender-petiolate, the blades ovate-lanceolate,

lanceolate, or lance-oblong, mostly 6-13 cm. long, long-acuminate, acute or obtuse

at the base and abruptly contracted, short-decurrent on the petiole, almost

glabrous, with dense, minute cystoliths on both surfaces, the margins entire or

somewhat undulate; inflorescences terminal and axillary, the spikes densely

flowered, mostly 3-6 cm. long and as much as 4 cm. broad; bracts broadly ovate to

suborbicular, 1.3-2.5 cm. long, acute or shortly cuspidate-acuminate, venose,

usually glabrous, ciliate; bracteoles oblong-lanceolate, usually acuminate; calyx

4-7 mm. long, the segments linear- acuminate, ciliate; corolla lavender to rose-

purple, 3-3.5 cm. long, puberulent outside, the tube much inflated in the throat

and about 1 cm. broad, the lobes very short, rounded or subretuse; capsule about

1 cm. long, puberulent.

BRAVAISIA DeCandolle

Trees or shrubs, glabrous or pubescent; leaves opposite, sometimes anisophyl-

lous, petiolate, the blades ovate, oblong-lanceolate, or elliptic, the margins entire;

inflorescences terminal, cymose, sometimes branching and often appearing thyrsi-

form, paniculate, or corymbose; bracts foliaceous; bracteoles ovate, oblong, or

obovate-spathulate to narrowly pandurate, shorter than or exceeding the calyx;

calyx lobes 5, separated almost to the base, the segments oblong; corolla tube

abruptly dilated into a campanulate throat, the limb erect-spreading, the 5 lobes

rounded or emarginate, contorted, subequal; stamens 4, didynamous, inserted

near the base of the corolla tube, included or exserted, the filaments distinct or

more or less united by a membrane at the base, the anthers dorsifixed, the thecae

parallel, nearly equal, calcarate at the base; pollen 2-porate, bilateral, prominently

banded; disc annular; style linear at the apex, revolute, the posterior lobe obsolete

or minute and dentiform; ovules 2-4 in each locule; capsule oblong, obovoid-

oblong, or ovoid, coriaceous, not compressed, little contracted at the base; seeds

2-8, lenticular, the retinacula often bidentate at the apex.

GIBSON: FLORA OF GUATEMALA 345

Three species, in tropical America. All three are found in Central

America with two in Guatemala. Bravaisia integerrima (Spreng.)

Standley is found in Mexico, El Salvador, and in southern Central

America, but we have seen no specimens from Guatemala; British

Honduras, or Honduras. It is a large tree, often 15-20 m. tall, with

broad leaves commonly 4-14 cm. wide, and the corollas are ampliate

from almost the base of the tube, the narrow portion of the tube

usually less than 1 mm. long.

Bracteoles longer than the calyx; corolla 1.5-2 cm. long B. tubiflora.

Bracteoles much shorter than the calyx; corolla 3.5-4.5 cm. long. . .B. grandiflora.

Bravaisia grandiflora Donn.-Sm. Bot. Gaz. 33: 255. 1902. B.

proximo, Blake, Contr. Gray Herb. 52: 96. 1917 (type from upper

Moho River, British Honduras, M. E. Peck 730). Boc-che (Quecchi,

Alta Verapaz).

Damp or wet, mixed forest near sea level to 1,000 m.; Alta

Verapaz (type from Sachicha, Tuerckheim 7924)', Pete"n. British

Honduras.

Shrubs or trees, sometimes to 9 m. tall; leaves short-petiolate, the blades

rather thick, elliptic or oblong-elliptic, mostly 6-18 cm. long, 3-6.5 cm. wide,

acuminate or caudate-acuminate, usually abruptly contracted at the base and

long-decurrent on the petiole, somewhat paler beneath, glabrous or sometimes

pilosulous on the petioles and costae; panicles of inflorescences sometimes about

as long as or longer than the leaves, usually many-flowered; bracts leaflike but

smaller, bracteoles ovate, oblong-ovate, or orbicular, 2-5 mm. long, mucronate at

the apex, ciliate, puberulent or almost glabrous; calyx often 3 times as long as the

bracteoles, the segments oblong, apiculate, 6-9 mm. long, puberulent and often

pilose; corolla pale lavender or pink, or the tube white and the limb with pale

lavender lobes, 3.5-4.5 cm. long, deeply lobate, the tube cylindric below, the limb

ampliate, the lobes oblong-obovate, pilose outside; filaments dilated and barbate

at the base, those of each pair distinct at base, the anthers 3.5-4 mm. long; ovary

oblong-ovoid; capsule unknown.

Bravaisia tubiflora Hemsl. in Hook. Icon. 16: t. 1516. 1886.

Hulaba and hulup (Maya, British Honduras).

Damp or wet forest, at or a little above sea level; British Hon-

duras. Mexico.

Shrubs or trees sometimes to 8 m. tall, the branches whitish, the young branch-

lets pilosulous; leaves thick, petiolate, the petioles often more or less pilose, the

blades ovate, oblong-lanceolate, or elliptic, 3-16 cm. long, 1.5-6.5 cm. wide,

short-acuminate to obtuse, acute at the base or abruptly contracted and more or

less decurrent on the petiole, glabrous or sometimes pilose on costae and veins

beneath, the short cystoliths numerous and usually conspicuous; cymes of inflores-

_

FIG. 75. Bravaisia grandiflora. A, habit, X J^; B, corolla opened to show sta-

mens and style, X 1%; C, anthers, X 5; D, detail of bracteoles and calyces, X 2^.

346

GIBSON: FLORA OF GUATEMALA 347

cences usually not more than 2 or 3 cm. long but sometimes occurring in great

numbers in the uppermost branches and appearing corymbose or paniculate;

bracteoles obovate-spathulate to narrowly pandurate, acute, at least the lower

portion ciliate, usually exceeding the calyx, mostly 8-12 mm. long; calyx lobes

oblong, 5-7 mm. long, obtuse or acute, not mucronate, subequal, more or less pu-

berulent or nearly glabrous, ciliate; corolla white or pale purplish, 1.5-2 cm. long,

the narrow portion of the tube 4-5 mm. long, pubescent outside, the lobes rounded,

5-8 mm. long, often puberulent, ciliate; stamens equalling or a little longer than the

corolla tube, the filaments of each pair connected at base by a membrane, some-

what dilated and barbate at base, the anthers 2.5-3 mm. long; capsule ovoid,

acute, 8-10 mm. long, glabrous, containing 4 shining, brown seeds.

BUCERAGENIA Greenman

Weak, soft-stemmed, erect herbs, the stems simple or sparsely branched;

leaves petiolate, the blades very thin, ovate to elliptic, entire; inflorescences

spicate, simple or branching and paniculate, the flowers solitary or fasciculate;

bracts and bracteoles minute; calyx segments 5, subequal, linear-subulate; corolla

white or purplish, small, the tube slender, scarcely dilated above, the limb shal-

lowly bilabiate, the posterior lip incurved, emarginate or bilobate, the anterior

lip erect or slightly spreading, trilobate; stamens 2, inserted at the middle of the

corolla tube, included; anthers usually monothecous, the lower theca usually

aborted or vestigial and sterile, if fertile, always considerably smaller, one or both

very minutely calcarate at the base; pollen spheroidal, 3-colporate; staminodes 2;

capsule clavate, narrowed below into a slender, solid stipe; seeds 4, flat, lenticular,

verrucose.

Three other species, all Mexican, have been reported.

Buceragenia glandulosa Leonard, Field Mus. Bot. 18: 1205.

1938.

Wooded swamps, 1,200-1,500 m.; Alta Verapaz. Mexico (Chia-

pas); Nicaragua; Costa Rica.

Slender, erect herbs about a meter tall, the stems glabrous or sparsely and

bifariously pilosulous; leaves on long, slender petioles, or the uppermost subsessile,

the blades ovate or oblong-ovate, mostly 7-15 cm. long and 3-7 cm. wide, acumi-

nate, acute to rounded at the base and abruptly and narrowly long-decurrent on

the petiole, glabrous, with numerous cystoliths; flowers sessile or nearly so, densely

fasciculate at the nodes of the spikes, the rachis glandular-pilosulous, the spikes

forming a large, open, terminal panicle; bracts lanceolate to triangular, about 2 mm.

long, the bracteoles similar but smaller; calyx lobes lanceolate, acuminate, 2.5-

3.5 mm. long, glabrous or nearly so; corolla 2-3 mm. long, glabrous, the tube

straight, the lobes rounded, about 0.5 mm. long; capsule 12-16 mm. long, glabrous;

seeds flat, verruculose, especially on the margins.

As stated by Leonard, this plant in general appearance resembles

P sender anthemum cuspidatum (Nees) Radlk. but may be distinguished

FIG. 76. Buceragenia glandulosa. A, habit, X K; B, corolla with calyx,

X 8^; C, corolla opened to show stamens and staminodes, X 11 Y?.', D, staminode

greatly enlarged; E, two anthers greatly enlarged, one theca aborted; F, calyx

(with bracts) partially opened to show pistil, X HjHj; G, fascicled inflorescences,

X 5; H, capsules, X 2; I, seed, X 3>£.

348

GIBSON: FLORA OF GUATEMALA 349

by its minute flowers which are densely fasciculate rather than

solitary.

CARLOWRIGHTIA Gray

Plants usually erect, herbaceous or somewhat woody, pubescent or glabrous;

leaves entire, broad or narrow; flowers small, in lax spikes or racemes, these often

forming large terminal panicles, the bracts and bracteoles small; calyx small, the

sepals 5, narrow; corolla tube short (in ours), slender, scarcely ampliate above, the

4 lobes almost equal, one shallowly bifid; stamens 2, the filaments inserted at or

above the middle of the corolla tube; anthers bithecous, nearly equal, muticous;

staminodes none; pollen 3-colporate, prolate to perprolate; stigma capitate, very

shallowly bilobate; capsule acute, contracted below into a short, solid stipe; seeds

4, flat, roughened; the retinacula long and thin.

Fifteen species have been reported, ranging from southwestern

United States to Central America. Only one species has been found

in Central America.

Carlowrightia costaricana Leonard, Field Mus. Bot. 18: 1206.

1938.

Damp or dry, often rocky thickets, 180-600 m.; Chiquimula;

Zacapa. Costa Rica, the type from Guanacaste.

Plants slender, erect, branched, to 75 cm. tall, the stems terete, the older ones

whitish, the young ones finely puberulent, soon glabrate; leaves membranaceous,

the lower ones on slender petioles to 1.5 cm. long, the upper ones subsessile, the

blades oblong-ovate, 2-10 cm. long and 1.5-4 cm. wide, long-acuminate to obtuse,

rounded to acute at the base, puberulent or glabrate; flowers borne in long, slender,

interrupted spikes, these usually forming an open panicle 10-20 cm. long, the

slender branches minutely puberulent or almost glabrous; bracts and bracteoles

subulate, to 3 mm. long; calyx about 1.5 mm. long, the sepals triangular, minutely

puberulent; corolla purple, about 7 mm. long, pubescent, the tube shorter than the

lobes; capsule 10-11 mm. long, acute, glabrous; seeds whitish, minutely roughened.

CHAETOTHYLAX Nees

Perennial herbs, sometimes suffrutescent below, pubescent or glabrous; leaves

petiolate, the blades entire; flowers sessile, usually in very dense, short, axillary

and terminal, subcapitate or spikelike inflorescences, sometimes solitary in the

upper leaf axils, the bracts and bracteoles usually narrow, rigid and filiform, or the

bracts sometimes broadly oblanceolate or elliptic and cuspidate; calyx segments 4,

linear to subulate, acute or acuminate, subequal; corolla tube slender, straight or

incurved, the limb bilabiate, the posterior lip innermost in bud, erect, narrow,

entire, the anterior lip spreading, trifid, the lobes subequal, the middle one outer-

most; stamens 2, anterior, the filaments inserted in or just below the throat of the

corolla, about equalling or slightly shorter than the posterior lip; anthers bithecous,

FIG. 77. Carlowrightia costaricana. A, habit, X V^, B, corolla opened to

show stamens, X 4>6; C, calyx and pistil, X 4^; D, fruiting branch, X 2^; E,

capsule with seeds, X 4)^.

350

FIG. 78. Chaetothylax rothschuhii. A, habit, X H; B, inflorescences,

C, capsule, X 6; D, corolla with calyx, X 6; E, corolla dissected to show one sta-

men and the style, X 6; F, limb of corolla prior to anthesis, X 6.

351

352 FIELDIANA: BOTANY, VOLUME 24

one theca much smaller than the other, the smaller one usually inserted much lower

on the connective, calcarate, often sterile, sometimes reduced to a small tooth;

pollen 2-porate, bilateral; staminodes none; disc annular or short-cupular; style

filiform, incurved at the apex, entire or minutely bidentate; ovules 2 in each locule;

capsule oblong, more or less contracted at the base; seeds 4 or by abortion fewer,

flat, minutely glochidiate, the retinacula acute or subobtuse.

Less than 10 species, ranging from southern Mexico into South

America. In addition to the two in Guatemala, two others have been

described from Nicaragua and Costa Rica.

Bracts filiform, pubescent C. rothschuhii.

Bracts oblanceolate, cuspidate, long-ciliate C. cuspidatus.

Chaetothylax cuspidatus D. Gibson, Fieldiana:Botany 34:

58. 1973.

Alta Verapaz, 650 m., valley of Santa Lucia, on road from Gualan

to Coban (type, Pittier 1792}. Honduras.

Herbaceous, erect or ascending plants 30-40 cm. tall, the stems bifariously

pubescent with white hairs; leaves short-petiolate, the blades lanceolate to ovate-

lanceolate, acuminate, entire, 2-6 cm. long, 1-2.5 cm. wide, glabrous or with

scattered pubescence mostly on costae and veins; inflorescences appearing sub-

capitate, 2-4 cm. long; bracts oblanceolate, 10-15 mm. long, cuspidate, long-

ciliate, the bracteoles linear-acuminate, rigid, 6-8 mm. long, ciliate; calyx segments

4, 7-9 mm. long, narrowly linear-acuminate, ciliate; corolla 2.5-3 cm. long, pubes-

cent outside, the tube 1.5-1.8 cm. long, the upper lip entire, the lower one trilobate;

stamens included, the filaments attached just below the throat of the corolla, the

anthers 1-1.5 mm. long, thecae superposed on a broad connective, the lower one

considerably smaller and calcarate; pollen 2-porate, bilateral; style filiform, the

stigma apparently entire; capsule unknown.

Chaetothylax rothschuhii Lindau, Bull. Herb. Boiss. 3: 492.

1895.

Dry or damp thickets, 250-500 m. ; Escuintla. Another collection

of this plant was made in Guatemala by Standley or Steyermark,

probably in the Oriente. The label, according to Dr. Standley's

notes, was lost by a careless mounter; therefore there is no data

concerning the local station. The species was described from Maz-

maz, Matagalpa, Nicaragua, and several collections have been made

in Honduras.

Erect or ascending herbs, the stems sometimes as much as 40 cm. long, simple

or sparsely branched, terete, hirsutulous above; leaves on very short petioles, the

blades rather thick and firm, lance-oblong or ovate-oblong, mostly 3-9 cm. long,

1-3 cm. wide, usually long-acuminate, sometimes acute, acute at the base, strigose

above along the costae, appressed-pilose beneath, the cystoliths obscure; inflores-

cences short and dense, many-flowered, headlike to spikelike, axillary and terminal,

GIBSON: FLORA OF GUATEMALA 353

sessile or nearly so, crowded at the ends of the branches; bracts and bracteoles

5-9 mm. long, filiform, usually rigid, pubescent; calyx segments filiform, 10-15 mm.

long; corolla white, puberulent above, the tube 9-11 mm. long, 1-1.5 mm. in

diameter, the posterior lip entire, 3-4 mm. long; capsule 6-7 mm. long, 2 mm.

broad, acute, minutely puberulent; seeds scarcely 1 mm. in diameter, very minutely

glochidiate.

DICLIPTERA Jussieu

Plants perennial, herbaceous or suffruticose, erect or ascending, the stems

hexagonal, pubescent or almost glabrous; leaves usually petiolate, the blades

entire; inflorescences cymose, bracteate, axillary or terminal, pedunculate, sessile

or subsessile, often crowded in the leaf axils, sometimes appearing spikelike;

flowers subtended by 2 or more pairs of bracts, the outer pair larger than the inner

ones; calyx usually dry or hyaline, the 5 segments linear or setaceous, subequal;

corolla commonly red or purple, the tube slender or ampliate above, the limb deeply

bilabiate, the lips narrow or ovate, the posterior one innermost in bud, erect, entire

or emarginate, the anterior lip usually broader, entire or very shallowly trilobate;

stamens 2, the filaments inserted on the corolla tube below the throat, almost

equalling the corolla in length, anthers bithecous, one theca attached lower than

the other on the connective, both muticous or the lower one rarely short-calcarate

at the base; pollen grains 3-colporate, perprolate to prolate; staminodes none; disc

cupular, truncate, oblique, or sinuate-dentate; style filiform, entire at the apex or

minutely bifid; ovules 2 in each locule of the ovary; capsule ovoid or suborbicular,

more or less compressed, subsessile or contracted and appearing stipiform, the

placentae separating elastically from their walls in the mature capsule; seeds 4 or

by abortion fewer, flat, suborbicular, smooth, muriculate or glochidiate, the

retinacula rather broad, acute.

There are about 80 species in the tropics or warm-temperate

regions of America, Africa, and Asia, with eight in Guatemala.

Various species have been misidentified as D. acuminata (Ruiz &

Pavon) Juss., but I have seen no Central American material that

matches either the original description and illustration (Fl. Peruv. 1:

10, t. 16, f. b, 1798, as Dianthera acuminata) or the Ruiz collection

from Peru. It would appear that the typical variety of this plant is

confined to South America and that the Mexican D. acuminata var.

spicata Nees may be incorrectly named.

Inflorescences on peduncles more than 2 cm. long (commonly 4-10 cm.), these

sometimes branching D. sciadephora.

Inflorescences not as above.

Corolla red or pale red.

Bracts subtending the cymes subulate; outermost pair of floral bracts 1-2.5

mm. wide; thecae nearly opposite D. assurgens.

Bracts subtending the cymes ovate to cylindric, obtuse and abruptly long-

acuminate; outermost pair of floral bracts 4-7 mm. wide; one theca well

below the other . . D. debilis.

354 FIELDIANA: BOTANY, VOLUME 24

Corolla purple, pink, or white flushed with pink.

Outermost pair of floral bracts abruptly spinose-aristate, the awns 2-3 mm.

long; usually enclosing only one flower D. unguiculata.

Outermost pair of floral bracts not spinose-aristate, but may be apiculate;

usually enclosing 3 or more flowers.

Outermost floral bracts oblong-spathulate, 4-4.5 mm. wide; corolla pink or

white with pink D. sumichrasti.

Outermost floral bracts ovate, obovate, or cuneate-obovate, 6-15 mm. wide;

corolla purple.

Outermost floral bracts densely pilose, usually spotted with magenta-

brown D. guttata.

Outermost floral bracts not as above.

Bracts cuneate-obovate, 8-12(15) mm. long, 4-7 mm. wide, ciliate

with long, mostly spreading hairs D. membranacea.

Bracts ovate to broadly ovate, 10-20 mm. long, 8-15 mm. wide,

ciliolate with minute, closely appressed hairs D. inutilis.

Dicliptera assurgens (L.) Juss. Ann. Mus. Paris 9: 269. 1807.

Justicia assurgens L. Syst. Nat. ed. 10. 2: 850. 1759. Dicliptera

sexangularis Juss. Ann. Mus. Paris 9: 269. 1807. Solenochasma

assurgens Jacq. Eclog. PL 2: 1. 1844. D. vahliana Nees in DC.

Prodr. 11: 489. 1847. D. mollis Nees, op. cii. 490. Yerba fluxion

(Pete"n) ; amor seco.

Damp thickets or forest, frequently along stream beds in sandy

or rocky places, sometimes in waste places, sea level to 900 m.; Alta

Verapaz; Pete"n; El Progreso; Retalhuleu; Solola; Suchitepe"quez.

Southern United States: Mexico; British Honduras to Panama;

West Indies; northern South America.

Erect, usually much branched herbs to 1.5 m. tall, the stems nearly glabrous

or thinly puberulous to pubescent; leaves petiolate, the blades ovate to lanceolate,

the upper ones 2-5 cm. long, 1-3 cm. wide, the lowermost ones to 9 cm. long, 3-4.5

cm. wide, usually acute or acuminate, sometimes obtuse and apiculate, rounded to

acute at the base, often abruptly decurrent on the petiole, glabrate or sparingly

hirtellous to pilose, especially on costae and veins; inflorescences lax and spikelike,

usually forming large, often leafy panicles, the flowers solitary or several in a

sessile or subsessile cymule subtended by a pair of subulate bracts 1.5-3 mm. long,

the floral bracts 6, the 2 outer ones linear-lanceolate or linear-oblong to oblong-

spathulate, unequal, 4-15 mm. long, 1-2.5 mm. wide, usually abruptly acute or

obtuse and apiculate, rarely acuminate to subulate, often somewhat scaberulous

and glandular-puberulent, the 4 inner bracts lanceolate, 2.5-5 mm. long, acumin-

ate, densely puberulent; calyx subhyaline, the subulate segments 2-4 mm. long;

corolla red, finely pubescent outside, to 3 cm. long, the narrow, subequal lips about

1.2 cm. long; filaments of stamens sparsely villous; anthers 2-2.5 mm. long, the

thecae nearly opposite; ovary and style glabrous; capsule 5-6 mm. long, pubescent,

containing 2 lenticular, minutely glochidiate seeds.

Only a few collections with the outer pair of floral bracts truly

subulate (matching the original description) were seen. Although

FIG. 79. Dicliptera assurgens. A, habit, X ^2', B, flower, X2; C, corolla

opened to show stamens and pistil, X 2; D, calyx with bracteoles and bracts, X 5;

E, section of fruiting branch showing mature capsule, X 2^; F, mature capsule

opened to show separation of placenta, X 23^; G, seed, X 5.

355

356 FIELDIANA: BOTANY, VOLUME 24

this form does occur throughout the range, the dominant form

appears to be that with the larger, more or less spathulate bracts

(D. vahliana). There is considerable intergradation of this character,

and both kinds of bracts may occur on the same plant (i.e. Pelly 24,

British Honduras).

Dicliptera debilis Leonard, Journ. Wash. Acad. Sci. 31: 99,

/. 4. 1941.

Known only from the type locality, dry, rocky brushy hills north

of Santa Rosa, Baja Verapaz, 1,500 m., the type being Standley

69823.

Weak, much-branched herbs to about 1 m. tall, the stems hexagonal, pilosulous

with short, recurved hairs; leaves petiolate, the blades ovate or lanceolate, 2-5 cm.

long, 1-2 cm. wide, acute or acuminate, acute and decurrent at the base, sparsely

pilose with subappressed white hairs beneath along the costae and veins, elsewhere

glabrous or nearly so; inflorescences cymose, sessile or subsessile, axillary, 3 or 4

flowers in each cyme; the pair of bracts subtending the cymes ovate to cylindric,

obtuse to truncate at the apex and abruptly long-acuminate, 3.5-5 mm. long, about

2 mm. wide at base; outer pair of floral bracts oblanceolate or cuneate-obovate,

one much larger than the other, as much as 12-15 mm. long and 4-7 mm. wide,

obtuse or rounded at the apex and apiculate, pilosulous or puberulent, many of

the hairs gland-tipped; inner bracts lanceolate, 5-7 mm. long, pubescent and

somewhat viscid; calyx 4-8 mm. long, puberulent, the segments subulate; corolla

pale red, 2.5-3 cm. long, minutely pubescent outside, the tube slender; filaments

of stamens pubescent near base; anthers 1-1.3 mm. long; ovary and style pubescent;

capsule compressed-ovoid, 6-9 mm. long, glandular-puberulent.

Dicliptera guttata Standl. & Leonard, Field Mus. Bot. 23:

239, /. 2. 1947.

Known only from the type, Zacapa, Sierra de las Minas, along

Rillito del Volcan de Monos, Volcan de Monos, 1,150-2,100 m.,

Steyermark 42396.

Plants suffrutescent, as much as a meter tall, the stems more or less hexagonal,

villosulous with short, weak, white hairs; leaves on slender petioles 1.5-5 cm. long,

the blades ovate or oblong-ovate, mostly 3-10 cm. long and 1-4 cm. wide, obtuse-

acuminate, narrowed at the base and decurrent on the petiole, hirsutulous on the

upper surface with appressed or subappressed hairs, rather coarsely hirsutulous

beneath, especially along the costae and veins; flowers arranged in crowded, com-

pound, terminal or axillary cymes as much as 2.5 cm. long and 3 cm. broad, these

subsessile or on pubescent peduncles to 1 cm. long; bracts subtending the cymes

5-6 mm. long, narrowly lanceolate and acuminate; the pair of larger floral bracts

obovate, 14-16 mm. long, 9-11 mm. wide, subobtuse, apiculate, densely pilose

and ciliate, usually conspicuously spotted with magenta-brown; inner bracts

subtending the flowers linear, acuminate, ciliate, 10-11 mm. long; calyx 8 mm.

GIBSON: FLORA OF GUATEMALA 357

long, the segments subulate, ciliate; corolla rose-purple, finely pubescent outside,

2.5 cm. long, the subequal lips 6-8 mm. long; filaments of stamens sparsely pubes-

cent; style glabrous or nearly so; capsule oval, compressed, 6 mm. long, 3 mm.

broad, obtuse and mucronate at the apex, sparsely hirtellous.

Dicliptera inutilis Leonard, Journ. Wash. Acad. Sci. 31: 101.

f. 5. 1941.

Damp thickets, 1,200 m. or less; Jutiapa (type from Los Llanitos,

near San Jose" Acatempa, Standley 60579); Retalhuleu (plains near

Champerico).

Erect branching herbs to about a meter tall, the branches hexagonal, sparsely

and minutely pubescent with more or less appressed hairs; leaves petiolate, the

blades ovate, 2-4 cm. long and 1-3 cm. wide, acuminate, obtuse or acute at the

base, appressed-pilosulous on the costae and veins, at least on the lower surface;

inflorescences cymose, terminal and axillary, sessile or very short-pedunculate

(usually less than 0.5 cm. long), each cyme composed of 3 cymules; a pair of

subulate bracts, 2-4 mm. long, subtending each cyme, each cymule subtended by

a pair of large, ovate to broadly ovate bracts, one larger than the other, as much

as 2 cm. long and 1.5 cm. wide, obtuse or narrowly rounded at the apex, often

apiculate, contracted at the subtruncate base into a short petiole, bright green,

minutely appressed-pubescent on the margins and sometimes so on the faces; 3 or

4 flowers within each of these large pairs of bracts, subtended by linear, acuminate

bracteoles 4-5 mm. long, puberulent; calyx segments subulate, 5-6 mm. long,

ciliate; corolla pale purple, to 2 cm. long, puberulent outside; filaments of the

stamens sparsely pubescent; ovary pubescent; capsule suborbicular to broadly

ovate, pubescent.

Dicliptera membranacea Leonard, Journ. Wash. Acad. Sci.

31: 102, /. 6. 1941.

Wet thickets or forest, 500-725 m.; Alta Verapaz; Escuintla (type

collected between Rio Jute and Rio Pantaleon, on the road between

Escuintla and Santa Lucia Cotz, Standley 63524}; San Marcos.

Mexico.

Erect, branching herbs to 1 m. tall, the branches hexagonal, glabrous or puberu-

lent at the nodes; leaves on petioles 1-6 cm. long, the blades ovate or oblong-ovate,

mostly 4-10 cm. long, 1.5-6 cm. wide, acute or acuminate, obtuse or acute and

abruptly decurrent at the base, essentially glabrous or sometimes more or less

pubescent beneath; inflorescences axillary or terminal, the flowers in sessile or

subsessile, dense, usually 4-flowered cymes, the cymes often crowded together and

appearing spikelike; bracts subtending the cymes ovate, foliaceous, to 12 mm. long

and 6 mm. wide, glabrous or minutely pubescent beneath; the outer pair of floral

bracts unequal, cuneate-obovate, 8-12 (15) mm. long and 3.5-6 (7) mm. wide,

costate, usually rounded and minutely apiculate or sometimes acute, ciliate; inner

bracts linear, acuminate, 7-12 mm. long, whitish, ciliate, the costae hirtellous;

calyx segments subulate, 4-6 mm. long, ciliate; corolla pale purple, puberulent

358 FIELDIANA: BOTANY, VOLUME 24

outside, to 2 cm. long; filaments of stamens villous near base, anthers to 1 mm.

long; ovary pubescent; capsule pubescent near apex.

Dicliptera sciadephora Donn.-Sm. Bot. Gaz. 23: 13. 1897. D.

vulcanica Leonard, Journ. Wash. Acad. Sci. 33: 71. 1943 (type

collected near Tajumulco, Volcan de Tajumulco, Steyermark 36861).

Damp thickets or forest, often along streams, sometimes in oak

forest, 1,350-2,400 m.; Alta Verapaz; Chimaltenango ; Huehue-

tenango (type from Jacaltenango, E. W. Nelson 3584); Pete"n; San

Marcos. Mexico (Chiapas); Nicaragua; Costa Rica.

Erect to subscandent, branching herbs, the stems more or less hexagonal,

sparsely pubescent to almost glabrous, the hairs somewhat appressed, spreading,

or retrorse; leaves of inflorescence subsessile to short-petiolate, those of lower stems

on petioles 1-3 cm. long, the blades ovate or oblong-ovate, mostly 2-10 cm. long,

1-5 cm. wide, usually acuminate, sometimes acute, obtuse or acute at the base

and abruptly decurrent, sparsely pilose or pilosulous, especially on the costae and

veins; inflorescences cymose, axillary, pedunculate, the peduncles mostly 4-10 cm.

long, sometimes shorter, each bearing mostly 3-5 pedicellate flowers, the pedicels

mostly 1-4 cm. long; each cyme subtended by a pair of subulate bracts 3-6 mm.

long; the outer pair of floral bracts linear or oblanceolate, acute to acuminate,

usually 8-15 mm. long, 2-4 mm. wide, puberulent, often pilose, sometimes with

gland-tipped hairs; the inner bracts lanceolate or subulate, 5-9 mm. long; calyx

7-10 mm. long, the segments lance-subulate, pubescent; corolla about 3 cm. long,

pale or dull red or reddish-purple, finely pubescent outside, the tube slender;

filaments of stamens more or less pubescent, especially near the base; ovary and

style pubescent; capsule 9-12 mm. long, about 5 mm. broad, puberulent, appearing

somewhat apiculate.

The inflorescences of a Chiapas collection, Breedlove 9049, are

more densely flowered and the indument is more glandular than any

Guatemalan collections I have seen. One Guatemalan collection,

Tuerckheim 2106, Alta Verapaz, is also atypical; its uniformly shorter

pedicels (0.5-1.5 cm. long) and some outer floral bracts as much as

18 mm. long and 6 mm. broad resemble those of the Costa Rican

plant, D. trifurca Oerst. However, the bracts of D. trifurca are

essentially glabrous and are rounded and apiculate at the apex.

Dicliptera sumichrasti Lindau, Bull. Herb. Boiss. 5: 665. 1897.

D. antidysenterica A. Molina, Ceiba 11: 65. 1965.

Wet forest, 350-1,000 m.; Alta Verapaz. Mexico; British

Honduras; Honduras.

Erect or ascending herbaceous or suffruticose plants to 1.5 m. tall, the stems

often branched, pilose or almost glabrous; leaves on petioles 1-8 cm. long, the

blades thin, mostly 3-14 cm. long, 2-7 cm. wide, ovate to lanceolate, acuminate

to long-acuminate, acute or obtuse at the base, pilose beneath, especially along

GIBSON: FLORA OF GUATEMALA 359

the costae and veins, or almost wholly glabrous; inflorescences pedunculate, the

peduncles usually 0.5-2 cm. long, cymose, sometimes appearing paniculate, the

flowers in dense clusters, 3-5 flowers subtended by a pair of lance-subulate bracts

4-8 mm. long, each flower subtended by 3-4 pairs of bracts, the outer pair oblong-

spathulate, long-ciliate, apiculate, the larger one 10-14 mm. long, 4-4.5 mm. wide,

the inner bracts lanceolate, pilose, 7-8 mm. long; calyx 3-4 mm. long, the segments

subequal, subulate; corolla pink or white flushed with pink, pubescent outside;

filaments of stamens pilose, the anthers about 1 mm. long; ovary and style pubes-

cent; capsule pubescent.

These plants have been confused with the South American D.

acuminata (Ruiz & Pavon) Juss. which also has ciliate bracts.

However, the inflorescences of D. acuminata are sessile, the outer

involucral pair of bracts are lanceolate to lance-oblong, acuminate,

only 2-3 mm. wide, and with a conspicuous apicula 1 mm. long or

more. The inner bracts are only 3-4 mm. long, and the calyx

segments 2-2.5 mm. long.

A species of Costa Rica and Panama, D. imbricata Leonard,

closely resembles D. sumichrasti, but differs in its smaller flowers

and shorter bracts. Its corollas are only 15-16 mm. long and the

two outer floral bracts are 7-9 mm. long.

Dicliptera unguiculata Nees in Benth. Bot. Voy. Sulphur 149.

1844. Hierba de cancer, panalitos (Guatemala).

Wet or dry thickets and fields, sometimes in rocky places or in

oak forest, common weeds in waste and cultivated ground, 900-1,800

m.; Chimaltenango; Guatemala; Jalapa; Quezaltenango ; Sacatepe*-

quez. Southern Mexico; Honduras and El Salvador to Costa Rica;

northwestern South America.

Erect or decumbent herbs, usually 30-60 cm. tall, branched, the stems pubes-

cent above; leaves slender-petiolate, the blades ovate, mostly 2-5 cm. long, 1-3 cm.

wide, sometimes to 8 cm. long and 5 cm. wide, acute or obtuse, obtuse or acute at

the base, densely pilosulous when young, sometimes glabrate in age; flowers

crowded in dense clusters, these forming dense, terminal or lateral spikelike

inflorescences 2-10 cm. long and about 2 cm. in diameter, each cymule usually

with only one flower and subtended by a pair of subulate bracts 4-7 mm. long;

the 2 outer floral bracts venose, usually suborbicular or ovate, sometimes spathu-

late, one larger than the other, usually 10-15 mm. long, 4-6 mm. wide, obtuse and

abruptly spinose-aristate at the apex, with an awn 2-3 mm. long, usually densely

hirsute with long slender hairs and long-ciliate, pale or bright green; inner bracts

linear-lanceolate, 4 mm. long, ciliate; calyx 3 mm. long, the segments narrowly

lanceolate, minutely ciliate; corolla rose-purple, to 1.5 cm. long, pubescent outside,

the tube slender, the lips subequal, 6-7 mm. long; f laments of stamens pubescent

or nearly glabrous; capsule clavate, 4 mm. long, hirsutulous; seeds flat, suborbicu-

lar, brownish, 1-1.5 mm. in diameter, minutely glochidiate.

360 FIELDIANA: BOTANY, VOLUME 24

In Costa Rica, where it is called "sornia," this plant has been

much used in treating amoebic dysentery.

DYSCHORISTE Nees

References: C. E. Kobuski, A monograph of the American species

of Dyschoriste, Ann. Mo. Bot. Card. 15: 9-90. 1928. R. W. Long,

Genera of Acanthaceae, Journ. Am. Arb. 51: 292-296. 1970.

Perennial herbs, sometimes suffruticose at base, prostrate to erect, usually

pubescent; leaves opposite, sessile or petiolate, entire or repand-crenate; inflores-

cences usually cymose, often appearing capitate, rarely racemose, axillary, the

flowers subtended by foliaceous bracts and bracteoles; calyx segments 5, usually

subulate-setaceous and ciliate, connate at anthesis for one-half to one-quarter their

length, rupturing easily in some species and often splitting to the base in fruit;

corolla tube incurved or straight, usually somewhat dilated at base, more or less

ampliate above, the limb spreading, oblique, 5-lobate, obscurely or distinctly

bilabiate; stamens 4, didynamous, the filaments attached below the throat of the

corolla; anthers bithecous, the thecae oblong, calcarate at the base, parallel or

slightly divergent; pollen 3-colporate, prolate, banded; ovary glabrous, with

usually 2 ovules in each locule; style filiform, pubescent; capsule ellipsoidal,

containing 2-4 flat, suborbicular seeds.

About 100 species have been reported from tropical and sub-

tropical America, Africa, and Asia, with three in Guatemala. Al-

though Kobusky recognized 40 American species, many are poorly

marked and the true number is probably fewer. Dyschoriste crinita

(Nees) Kuntze, a Brazilian species, was reported from Huehuetenango

and Quezaltenango by Loesner, but those plants are probably

referable to D. quadrangularis (Oerst.) Kuntze.

Corolla 2-2.8 cm. long D. ovata.

Corolla 1-1.5 cm. long.

Leaves mostly 3-10 cm. long, the blades ovate to oblong, usually acute to

acuminate at the apex; calyx segments at anthesis connate for about one-

half their length D. quadrangularis.

Leaves mostly 0.8-2 cm. long, the blades obovate to spathulate, obtuse or

rounded at the apex; calyx segments at anthesis connate for one-third or

one-fourth their length D. skutchii.

Dyschoriste ovata (Cav.) Kuntze, Rev. Gen. 2: 486. 1891.

Ruellia ovata Cav. Icon. 3 : 28. t . 254. 1794. Calophanes ovatus Benth.

in DC. Prodr. 11: 108. 1847. Clarencillo (Huehuetenango).

Damp or rather dry, open, grassy places, often in pine forest,

1,800-2,100 m.; Chimaltenango; Huehuetenango; reported by Loese-

ner from Quezaltenango. Southern Mexico.

Erect plants to 75 cm. tall, the stems simple or sparsely branched, obtusely

quadrangular or subterete, pilose, at least on the angles, with very short, spreading

FIG. 80. Dyschoriste ovata. A, habit, XJHj; B, corolla opened to show sta-

mens (flower in background with unopened corolla), X 3; G, anthers, X ll/z', D,

calyx opened to show pistil, X 3; E, calyces with immature capsules, X 2^; F,

two capsules, one opened to show seeds, X 23^; G, seed, X 10.

361

362 FIELDIANA: BOTANY, VOLUME 24

hairs; leaves short-petiolate, the blades narrowly ovate to obovate or oblong-

elliptic, mostly 3-4 cm. long, obtuse at apex, cuneate or attenuate to the base,

with scattered short spreading hairs on both surfaces along the costae and veins;

flowers glomerate in the leaf axils; bracts resembling the leaves but smaller and

narrower, about equalling or shorter than the calyces; calyx glabrous or sparsely

strigillose, densely covered with cystoliths, the segments 12-15 mm. long, subulate-

setaceous, ciliate, connate, at least at anthesis and usually in fruit, for one-quarter

to one-third their length; corolla lavender, 2-2.8 cm. long, puberulent outside, the

tube less than 1 cm. long, slender; capsule 10-12 mm. long, brown, containing

2-4 seeds.

Dyschoriste quadrangularis (Oerst.) Kuntze, Rev. Gen. 2: 486.

1891. Calophanes quadrangularis Oerst. Vid. Medd. Kjoebenhavn

1854: 120. 1855. Tucuquita de agua (Jutiapa).

Open grassy fields and hillsides, damp or dry, often rocky thickets,

pine-oak forest, or sometimes a weed in waste ground, 800-1,800 m.;

Chimaltenango ; Guatemala; Jalapa; Jutiapa; Sacatepe"quez; Santa

Rosa. Southern Mexico; El Salvador; Honduras; Nicaragua; Costa

Rica.

Plants decumbent, ascending, or erect, to 60 cm. tall, the stems simple or

sparsely branched, quadrangular and with very narrowly wing-margined angles,

densely short-pilose, at least on the angles; leaves short-petiolate, the blades

ovate to oblong, mostly 3-10 cm. long, usually acute or even acuminate, sometimes

with an obtuse tip, rather densely short-pilose or villosulous, or sometimes glabrate

except on the veins, nearly entire or more or less repand-crenate; flowers densely

cymose-glomerate in the leaf axils, numerous; bracts foliaceous, equalling or shorter

than the calyx; calyx 9-14 mm. long, densely covered with cystoliths and appearing

strigillose, the segments connate for about one-half their length, at least at anthesis

and sometimes in fruit, subulate setaceous, puberulent, ciliate; corolla commonly

pale purple or pinkish, rarely white, 7-12 (15) mm. long, pubescent outside;

capsule glabrous, 8-11 mm. long, containing 4 whitish-pubescent seeds.

Dyschoriste skutchii Leonard, Journ. Wash. Acad. Sci. 33:

70. 1943.

Open, rather dry, grassy places, often in pine forest, sometimes

in gravel along streams, 1,500-2,800 m.; Chimaltenango (type col-

lected near Tecpam, A. F. Skutch 4-74)', Huehuetenango; Jalapa; El

Quiche1. Mexico (Chiapas) ; Nicaragua.

Stems several, prostrate or ascending, to 35 cm. long, slender, simple or much

branched, sub terete, more or less puberulent with often recurved hairs; leaves

short-petiolate, the blades obovate or spathulate, mostly 0.8-2 cm. long, obtuse or

rounded at the apex, cuneate at the base, glabrous or sparsely hispidulous beneath

on the costae, often ciliate toward the base; flowers glomerate in the upper leaf

axils; bracts resembling the leaves but smaller; calyx 8-13 mm. long, glabrous or

GIBSON: FLORA OF GUATEMALA 363

sparsely hirtellous, the segments setaceous, hispidulous, ciliate, connate at anthesis

for about one-fourth their length; corolla lavender or purple, 11-14 mm. long,

puberulent outside, the narrow portion of the tube 4-5 mm. long, the limb 7-8 mm.

broad, the short lobes rounded; capsule brown, glabrous, about 10 mm. long,

containing 2-4 seeds.

These plants might be referred to any of several species described

from Mexico, but we hesitate to make such a reduction at this time

because the status of the Mexican species is so uncertain.

ELYTRARIA Michaux

Low perennial herbs, acaulescent or with short, slender, simple or sparsely

branched stems, the stems usually with clusters or rosettes of leaves at the apex;

leaves alternate or subopposite, petiolate, the blades membranaceous, attenuate to

the base, entire, often undulate; inflorescences spicate, the small flowers solitary

and sessile in the axils of small imbricate bracts; spikes slender, cylindric, simple

or branching; bracts rigid, oblong, elliptic, or lanceolate, imbricate, about equalling

the flowers; bracteoles narrow, scarious, about equalling the calyx; calyx lobes 4,

unequal in width, lanceolate or linear, acute to acuminate, scarious, the anterior

one sometimes bidentate at the apex; corolla white, blue, pink, lavender, or white

with purple, the tube slender, cylindric, the limb bilabiate, the lips narrow, the

posterior one outermost in bud, emarginate or bilobate, the anterior lip trilobate,

hoodlike at first; stamens 2, the short filaments inserted below the throat of the

corolla, scarcely exserted; anthers dorsifixed near the base, the thecae equal,

parallel, muticous or calcarate at the base; staminodes very minute or none;

pollen 3-colpate, prolate; disc inconspicuous; style very shallowly bilobate at the

apex; ovules 6-12 in each locule; capsule oblong-linear, more or less contracted and

acute or rostrate at the apex; seeds minute, ovoid or globose, minutely tuberculate-

rugose, borne on papilliform retinacula.

Species three or four, native in tropical America, one of them

introduced into the Old World tropics.

Considerable disagreement exists as to whether or not Ely tr aria,

together with other members of the Nelsonioidae, should be retained

in the Acanthaceae, as they differ from typical members of the

family principally in their well-developed endosperm, their minute

seeds, as many as 12 in each locule, absence of retinacula, absence

of cystoliths, and in their pollen morphology.

Bremekamp (Proc. Acad. Sci. Netherlands, Ser. C, 56: 533-546.

1953) transferred the Nelsonioidae to the Scrophulariaceae "in the

vicinity of the Rhinantheae as a tribe Nelsonieae." Johri and Singh

(Bot. Notiser 112: 225. 1959) concluded that the Nelsonioidae,

although separated from the rest of the Acanthaceae, are not truly

oriented toward the Scrophulariaceae. On the basis of pollen

morphology, Raj (Grana Palynologica 3: 89, 92-93, 101. 1961)

364 FIELDIANA: BOTANY, VOLUME 24

thought that the transfer of the tribe to the Scrophulariaceae was

justified. Burtt (Bull. Bot. Surv. India 7: 77-78. 1965) rejected

Bremekamp's transfer.

Perhaps creation of a separate family, first proposed by Lindley

in 1847, would be the best solution, despite the recurving capsule

valves that appear to have identified this group as acanthaceous;

certainly other families in the Tubiflorae share more than one type

of fruiting structure.

Plants acaulescent; bracts of inflorescence entire E. bromoides.

Plants usually caulescent; bracts of inflorescence each with a small tooth at apex

and with 2 hyaline teeth or wings on upper margin E. imbricata.

Elytraria bromoides Oerst. Nat. For. Kjoebenhavn Medd.

115. 1854.

Open grasslands, 200 m. or lower; Pete"n. Western Texas;

Mexico.

Plants small, acaulescent; leaves rosulate, short- petiolate, the blades spathu-

late, 3-15 cm. long and 1-2.5 cm. wide, obtuse or rounded at the apex, attenuate

to the base, undulate, sparsely pilose with long slender hairs or almost wholly

glabrous; peduncles very slender and wiry, 2-10 cm. long, solitary or several,

densely bracteate, the bracts lance-subulate, 4-6 mm. long, ciliate; flowering spikes

mostly 1-3 cm. long, the floral bracts closely imbricate, lanceolate, long-attenuate

to subulate, as much as 10 mm. long and 2 mm. wide, glabrous outside, ciliate;

calyx lobes lanceolate, the anterior one bilobate at the apex; corolla pale pink to

lavender, 5-7 mm. long; capsule glabrous, 4-5 mm. long; seeds 8 or more.

Elytraria imbricata (Vahl) Pers. Syn. PI. 1: 23. 1805. Justicia

imbricata Vahl, Ecol. Amer. 1. 1796. Verbena squamosa Jacq. PI.

Hort. Schoenbr. 1: 3, £. 5. 1797. Elytraria tridentata Vahl, Enum.

PL 1: 107. 1804. E. frondosa HBK. Nova Gen. & Sp. 2: 234. 1817.

E. scorpioides Roem. & Schult. Syst. Veg. Mant. 1: 128. 1822.

Tubiflora squamosa Kuntze, Rev. Gen. 2: 500. 1891. Elytraria

squamosa Lindau, Anal. Inst. Fisco-Geogr. Costa Rica 8: 299. 1895.

Flor de mosquito ( Jutiapa) .

Damp or dry thickets, open or shaded banks, ravines, roadsides,

waste places, 75-1,800 m.; Alta Verapaz; Baja Verapaz; Chimal-

tenango; Escuintla; Guatemala; Huehuetenango; Jutiapa; Pete"n; El

Quiche" ; Retalhuleu; Sacatepe"quez ; Santa Rosa; Suchitepe"quez;

Zacapa. Texas; Mexico; British Honduras to El Salvador and

Panama; West Indies; South America; introduced into Old World

tropics.

FIG. 81. Elytraria imbricata. A, habit, X ^2; B, flowering spike, showing

cauline bracts and bracts of inflorescence, X 4; C, bract, X 8; D, calyx, X 10;

E, corolla opened to show stamens and pistil, X 10; F, capsules, X 5; G, seeds,

X 15.

365

366 FIELDIANA: BOTANY, VOLUME 24

Perennials, usually with a slender stem 2-10 cm. long (rarely 15 or 20 cm. long),

naked, leaves and flowering spikes usually borne at the apex; leaves petiolate, ovate

to oblong or obovate, mostly 3-15 cm. long, commonly acute or acuminate, long-

attenuate to the base and narrowly long-decurrent on the petiole, sparsely ap-

pressed-pilose or almost glabrous, the margins often undulate; peduncles few or

numerous, 2-15 cm. long, slender, wiry, covered with imbricate, closely appressed,

coriaceous, ovate to subulate bracts; spikes of inflorescence mostly 2-6 cm. long;

bracts oblong or elliptic, sometimes lanceolate, 3-6 mm. long, mucronate, and

bearing near the apex 2 hyaline teeth or wings, these narrow or broad; bracteoles

subulate, about 3 mm. long, costate, the costae long-ciliate; corolla light blue to

pale purple, 3-6 mm. long; capsule oblong, glabrous, about 3 mm. long, containing

8, 12, or more seeds.

This is one of the most common weeds of waste ground in the

Central American lowlands. Dr. Standley has noted, in manuscript,

that such names as coquito, guacoco, cacahuillo, and culantrillo, used

in different regions, are given because the little plants topped with

clusters of spreading leaves are somewhat suggestive of coconut

palms.

HABRACANTHUS Nees

Plants erect or ascending, herbaceous or suffrutescent, usually glabrous or

nearly so; leaves petiolate, the blades entire, thin; inflorescences cymose, lax, the

cymes arranged in many-flowered, usually terminal thyrses; bracts small, linear,

the bracteoles minute or none; calyx segments 5, linear, subequal, acute; corolla

tube cylindric, straight or arcuate, the lower portion narrow, a little ampliate

above or abruptly enlarged above, the limb deeply bilabiate, the lips subequal,

the posterior lip innermost in bud, erect or incurved, narrow, entire, the anterior

one usually more or less spreading, shallowly trilobate at the apex; stamens 2,

anterior, exserted or equalling the corolla; anthers normally monothecous, oblong-

linear, dorsifixed at or below the middle, muticous; staminodes none; pollen 2-

porate, bilateral, subspheroidal, the sexine granular to verrucose; disc thickened

but inconspicuous; style filiform, obtuse and minutely bilobate at the apex; ovules

2 in each locule; capsule oblong, contracted and stipiform at the base; seeds 4 or by

abortion fewer, suborbicular, flat, more or less rugose.

Four species are found in Central America; three are in Guate-

mala and a fourth, H. silvaticus Nees, may be expected there and is

therefore included here. Thirty-four species have been reported

from northern South America.

Corollas red, 2.5-3 cm. long H, ruberrimus.

Corollas white, blue, or lavender, 1-2 cm. long.

Posterior corolla lip narrowly linear, 8-10 mm. long.

Leaves sparsely pubescent, the margins ciliate; anterior corolla lip 5-7 mm.

wide; anthers 3 mm. long H. latilabris.

GIBSON: FLORA OF GUATEMALA 367

Leaves essentially glabrous, the margins not ciliate; anterior corolla lip 2-4

mm. wide; anthers 1.5-2 mm. long H. silvaticus.

Posterior corolla lip narrowly ovate, 5-6 mm. long H. azureus.

Habracanthus azureus D. Gibson, Fieldiana: Botany 34: 59.

1973.

Wet mountain forest, 1,500-2,400 m.; Alta Verapaz; San Marcos

(type from west facing slopes of the Sierre Madre mountains near

Aldea Fraternidad, Williams, Molina and Williams 25808).

Herbaceous plants to 2 m. tall, simple or branching, the stems bifariously

pubescent to pilose; leaves on pubescent petioles, the blades thin, lance-oblong,

ovate-oblong, or elliptic, mostly 6-15 cm. long, 2.5-8 cm. wide, acuminate, usually

contracted at the base and more or less decurrent on the petioles, more or less

pubescent on both surfaces with segmented hairs, usually more densely so on

costae and veins, the margins ciliolate; inflorescences usually terminal, paniculate,

5-25 cm. long, much branched, the rachis glabrous, the flowers sessile or borne

on short, usually glabrous pedicels about 1 mm. long; bracts subulate, 1-2 mm.

long; calyx segments linear, 6-10 mm. long at an thesis, to 15 mm. in fruit, usually

glabrous, rarely glandular-pubescent; corolla about 2 cm. long, blue, blue-purple,

or white, glabrous, the narrow portion of the tube 3-4 mm. long, the expanded

throat about 6 mm. wide, the lips subequal, the posterior lip 5-6 mm. long, the

lobes of the anterior lip 1-1.5 mm. long; stamens long-exserted, the anthers about

3 mm. long; capsule 12-15 mm. long, glabrous, containing 4 verrucose seeds.

Both glabrous and glandular-pubescent calyces are sometimes

borne on the same inflorescence; when a calyx is pubescent, the

pedicel is usually pubescent as well. Aborted (or cleistogamous?)

flowers are present on all specimens examined.

Habracanthus latilabris D. Gibson, Fieldiana: Botany 34:

60. 1973.

Mixed forest, 1,640 m., El Quiche; Cerro Putul (Skutch 1836,

type).

Suffruticose, branching plants, about 1 m. tall, the stems more or less pubes-

cent; leaves short-petiolate, the blades entire, oblong-ovate or lanceolate, acumi-

nate, rounded and abruptly attenuate to the petiole or somewhat auriculate at the

base, mostly 6-8 cm. long, 2-4 cm. wide, sparsely pubescent with scattered hairs

on both surfaces, the cystoliths inconspicuous, margins ciliate; inflorescences

axillary and terminal, thyrsiform-paniculate, 7-15 cm. long, 3-6 cm. wide, glabrous,

the branches purplish, the pedicels commonly 5-7 mm. long; bracts and bracteoles

subulate, 0.5-1 mm. long; calyx 4-6 mm. long, the lobes linear-acuminate; corolla

white, 13-15 mm. long, the tube 1-2 mm. in diameter, about 4 mm. long, the

posterior lip narrowly linear, 8-9 mm. long, the anterior one 5-7 mm. wide, shallow-

ly trilobate at the apex; stamens exserted, the anthers about 3 mm. long; style

filiform, exserted; capsule unknown.

368 FIELDIANA: BOTANY, VOLUME 24

Habracanthus ruberrimus D. Gibson, Fieldiana: Botany 34:

62. 1973.

Mixed forest between Chama and Coban, 460 m., Alta Verapaz

( Harry Johnson 491 , type) .

Suffruticose, branching plants to 1.5 m. tall, the stems glabrous or bifariously

pubescent; leaves short-petiolate, the blades lanceolate to elliptic, acuminate,

acute or attenuate to the base, mostly 5-12 cm. long, 1.5-5 cm. wide, glabrous, or

the younger leaves sometimes with appressed pubescence on costae and veins;

inflorescences terminal, glabrous, commonly 5-10 cm. long, 1.5-5 cm. wide; bracts

and bracteoles subulate, 1-2 mm. long; calyx 3.5-5 mm. long, the lobes narrowly

linear-acuminate; corolla bright red, 2.5-3 cm. long, the tube about 1.5 mm. in

diameter below, 2-3 mm. above, lips 1.5-2 cm. long, the upper one narrowly

linear to oblong, the lower one 3-4 mm. wide, shallowly trilobate at the apex;

stamens exserted, anthers about 3 mm. long; style filiform, exserted; capsule

unknown.

Although most anthers are monothecous, a few are bithecous.

When this occurs, the lower theca is always much smaller, but not

always sterile.

Habracanthus silvaticus Nees in DC. Prodr. 11: 312. 1847.

Not in Guatemala but may be expected there. Mexico (Oaxaca

and Chiapas) ; Costa Rica, 1,000-1,800 m.

Erect but weak herbs, sometimes suffrutescent below, usually not more than a

meter tall, branched, the stems glabrous or puberulent; leaves on slender petioles,

the blades oblong-ovate to lanceolate, mostly 5-12 cm. long, 1.5-6 cm. wide,

acuminate with an obtuse tip, acute or attenuate to the base, glabrous, or some-

times pubescent beneath with small curved hairs on the costae and veins, both

surfaces usually with minute cystoliths; inflorescences mostly 5-10 cm. long and

4-5 cm. broad, lax, many-flowered, glabrous or nearly so, the branches whitish or

purplish; bracts subulate, the lowest 3-4 mm. long, the pedicels commonly 3-5

mm. long; calyx lobes usually pale, 3-5 mm. long; corolla 10-15 mm. long, white,

bluish, or lavender, the tube about 1 mm. in diameter below, 1.5 mm. broad in the

throat, the lips about 10 mm. long, the upper one narrowly linear to oblong, the

lower one 2-4 mm. wide, rounded at the apex and shallowly trilobate; stamens

exserted, the anthers 1.5-2 mm. long; capsule 10-15 mm. long; seeds whitish,

tuber culate-r ugose .

HANSTEINIA Oersted

Herbaceous or sometimes suflrutescent plants, erect or ascending, often weak

and brittle, glandular-pubescent or essentially glabrous; lower leaves petiolate,

the upper ones subtending the inflorescences usually sessile, the blades thin and

flaccid, entire; inflorescences usually terminal, lax, paniculate, the panicles pyra-

midal or thyrsiform; bracts small, the bracteoles minute or none; calyx accrescent,

the 5 segments linear or nearly so; corolla more or less saccate, the tube narrow

FIG. 82. Habracanthus ruberrimus. A, habit, X %', B, flower with calyx, style,

and stamens, X 23^; C, corolla tube opened to show ovary and filament attach-

ment, X 2,^; D, apex of style enlarged to show stigma, X 5; E, anthers, X 6.

369

370 FIELDIANA: BOTANY, VOLUME 24

at the base, abruptly expanded into an oblong or barrel-shaped throat, slightly

contracted below the limb, the lips short, the posterior one innermost in bud, erect,

entire or minutely fimbriate, the anterior lip very shallowly trifid and appearing

almost truncate; stamens 2, exserted, the filaments inserted near the base of the

corolla throat; anthers commonly monothecous (very rarely bithecous but then

the lower one always much smaller), linear, dorsifixed near the middle; staminodes

none; pollen 2-porate, bilateral, spheroidal, the sexine verrucose; disc short, thick;

style filiform, the stigma appearing entire but actually minutely bifid; ovules 2 in

each locule; capsule oblong, contracted and stipiform at the base; seeds 4 or by

abortion fewer, lenticular, compressed, foveolate-rugose, on long retinacula.

Five species, three in Mexico and Central America and two others

in South America; only two are known in Guatemala.

Peduncles and rachis of inflorescence usually densely glandular-pilose (rarely only

viscid); mature corollas 4-5 (6) mm. wide in throat H. monolopha.

Peduncles and rachis of inflorescence glabrous; mature corollas 7-10 mm. wide

in throat H. glabra.

Hansteinia glabra (Leonard) D. Gibson, Fieldiana: Botany 34:

62. 1973. Glockeria glabra Leonard ex Lundell, Contr. Univ. Mich.

Herb. 6:60, f. 4- 1941.

Quezaltenango, 2,666 m. (Vaght 301). Mexico (Chiapas, Matuda

S-212, type).

Herbaceous plants, the stems glabrous; leaves petiolate, the blades thin, ovate

to elliptic, mostly 6-12 cm. long, 3.5-6 cm. wide, acuminate, narrowed at the base

and decurrent on the petiole, glabrous or sparsely pubescent; inflorescences usually

terminal, thyrsiform, commonly 10-15 cm. long, the rachis glabrous; bracts

subulate, 1.5-3 mm. long; calyx 5-8 mm. long, glabrous, the segments linear;

corolla red and yellow, 2-2.5 mm. long, glabrous or nearly so; the narrow lower

part of the tube 3-4 mm. long, the expanded portion above 7-10 mm. wide, the

posterior lip erect, ovate, about 3.5 mm. long, the anterior one shallowly trilobate;

stamens exserted, the anthers 3-3.5 mm. long; ovary glabrous; capsule unknown.

Hansteinia monolopha (Donn.-Sm.) D. Gibson, Fieldiana:

Botany 34: 62. 1973. Glockeria, monolopha Donn.-Sm. Bot. Gaz. 27:

439. 1899. G. moralesii Standley, Field Mus. Bot. 8: 47. 1930 (type

from San Martin Jilotepeque, Chimaltenango, J. Morales R. 1235).

Damp or wet, mixed forest or thickets, 1,500-2,700 m.; Alta

Verapaz; Chimaltenango; Escuintla; Quezaltenango; Sacatepe"quez;

San Marcos. Mexico (Chiapas) ; Costa Rica.

Erect or ascending, weak, usually brittle herbs, 1-2.5 m. tall, usually densely

glandular-pilose on the stems, petioles, and inflorescences, viscid; lower leaves on

slender petioles, the blades thin, ovate or oblong-ovate, mostly 10-20 cm. long,

5-12 cm. wide, acuminate, usually abruptly contracted and decurrent at the base,

sparsely villous on both surfaces or glabra te; inflorescences paniculate, often as

much as 30 cm. long, much branched, the peduncles, rachis, and pedicels usually

FIG. 83. Hansteirtia monolopha. A, habit, X Y^', B, detail of inflorescence

showing flower, X 4, with inset (greatly enlarged) of glandular hairs of calyx; G,

corolla dissected to show stamens and pistil, X 4; D, calyx with capsule begin-

ning to open, X 3; E, calyx with opened capsule showing retinacula, X 3; F, seed,

X 10.

371

372 FIELDIANA: BOTANY, VOLUME 24

densely glandular-pilose (rarely only viscid); the flowers on slender pedicels; bracts

subulate, 2-3 mm. long; calyx usually densely glandular -pilose, the segments linear,

5-7 mm. long at anthesis, often to 10 mm. long in fruit; corolla red and yellow,

glabrous outside, the narrow portion of the tube about 1.5 mm. broad, 3-5 mm.

long, the throat 4-5 (6) mm. broad, 10-15 mm. long, the posterior lip about 3

mm. long; stamens exserted, often far beyond the lips, the anthers about 3 mm.

long; capsule 10-13 mm. long, glabrous.

HEMIGRAPHIS Nees

Erect or ascending, rarely creeping perennial herbs, more or less pubescent;

leaves opposite, the blades usually undulate or crenate, sometimes entire; in-

florescences terminal or axillary, spicate, with flowers in the axils of opposite,

imbricate bracts; calyx segments 5, subequal; corolla tube straight or slightly

curved, the limb spreading, the 5 lobes contorted in bud; stamens 4, didynamous,

the filaments connected at the base, inserted at or below the corolla throat, the

anthers bithecous, muticous; staminodes none; ovules 4-6 in each locule; stigma

bilobate, the lower lobe often inconspicuous; capsule fusiform, seminiferous from

the base, containing 6-12 flat, pubescent seeds.

The genus is Asiatic, occasionally planted in gardens in Mexico

and in Honduras, sometimes escaping.

Hemigraphis alternata (Burm. f.) T. Anders, Journ. Linn. Soc.

7: 114. 1864. Ruettia alternata Burm. f., Fl. Indica 135. 1768.

Ruellia colorata Blume, Bijdr. Fl. Nederl. Ind. 795. 1826. Hemi-

graphis color ata Hallier, Nova Acta Leopoldina, Abh. Deutsche

Akad. Naturforscher 70: 204. 1898. Blechum cordatum Leonard,

Carnegie Inst. Wash. Publ. 461: 200. 1936 (type from British

Honduras, Bartlett 11355}.

Ascending or creeping herbs, the stems pilosulous, rooting at the lower nodes;

leaves on petioles 1-5 cm. long, the blades mostly 3-6 cm. long, 2-5 cm. wide,

broadly ovate to ovate-oblong, acute or obtuse, cordate at base, densely puberulent,

sparingly pilosulous or glabrate, the margins crenate; inflorescences terminal,

spicate, 2-3 (10) cm. long, dense or lax; bracts ovate-lanceolate, obtuse, 0.8-1.5 cm.

long; bracteoles minute or absent; calyx 7-10 mm. long, the segments subulate,

ciliate; corolla white, about 1.5 cm. long; capsule not seen.

Native of Malaysia, and no doubt an escape in British Honduras,

the type locality of Blechum cordatum Leonard.

HENRYA Nees

Reference: George B. Happ, Henrya, Ann. Mo. Bot. Gard. 24:

541-568. 1937.

GIBSON: FLORA OF GUATEMALA

373

FIG. 84. Hemigraphis alternata. A, habit, X 1; B, flower with corolla sepa-