A TAXONOMIC REVISION OF THE HELICONIJ IN MIDDLE AMERICA

By ROBERT ROY SMITH

A DISSERTATION PBESENTED TO THE GRADUATE COUNCIL OF THE DIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA 1968 Copyright by Robert Roy Smith 1968 ACKNOWLEDGMENTS

I wish to express deep appreciation to Dr. Daniel B.

Ward for his encouragement, advice and helpful criticisms

while directing this work. Acknowledgments are also ex-

tended to the curators and staffs of the herbaria listed

In the Introduction, for the loans and photographs of many of the pertinent types.

Special thanks are extended to Mr. John Beckner for

suggesting this problem, and for his continual Interest

throughout the study. I am grateful to Dr. Ward, Dr.

Klmbrough, Dr. Ford, Dr. Laessle, and Mr. Beckner for reading and evaluating the manuscript.

Appreciation Is extended to Mr. Steve Johnson for

technical assistance concerning the plates used In the systematic treatment.

A special thanks must be extended to my wife, Julia, for the devotion and understanding shown during the two years It took to complete this study.

Ill TABLE OP CONTENTS

Page ACKNOWLEDGMENTS IH

LIST OF TABLES •, . . vll

LIST OF FIGURES vlli

LIST OF MAPS Ix

LIST OF PLATES xl

INTRODUCTION 1

HISTORY 3

ECONOMIC VALUE 6

SURVEY OF THE ORDER SCITAMINEAE 8

MORPHOLOGY 23

POLLINATION 30

CYTOLOGICAL ASPECTS OF 32 PHYLOGENY 3^ SYSTEMATIC TREATMENT 40

HELICONIA ' i^Q

Key to Sections 43

I. Section TAENIOSTROBUS 44

Key to the Series 4^

Key to the In Series Imbrlcatae 46 !• Hellconla marlae i^-p 2. Hellconfa reticulata 53 3» Hellconla Imbrlcata 58 Series Eplscopales 61

xx Page II. Section HELICONIA 64

Key to the Species In Section HELICONIA 65 4. Helicon la carlbaea 66 5« Hellconla bourgaeana . », , , 73 6. Hellconla wagnerlana » « « « « « » 83 7. Hellconla blhal ....T 91

III. Section PENDUIAE 98

Key to Species In Section PENDUIAE 99 8. Hellconla marglnata IO3 9. Hellconla colgantea IO7 10. Hellconla nutans 112 11. Hellconla allenll II7 12. Hellconla vellerlgera 122 13« Hellconla magnlf lea . 127 l4. Hellconla curtlspatha I3I 15« Hellconla xanthotrlcha I36 16. Hellconla paulll ...T7T Ikl

17. Hellconla longa . Ik6 18. Hellconla coll Ins lana I54

19» Hellconla catheta . I65

IV. Section STENOCHIAMYS 172

Key to the Series STENOCHIAMYS I72

Key to the Species of Series Cannoldeae I73 20. Hellconla pslttacorum I75 21. Hellconla aurantlaca 181 22. Hellconla leucoflora 188 23« Hellconla hlrsuta 24. Hellconla vaginalis 202 25. Hellconla trlnldatTs 213 26. Hellconla swart z lana 218 Series Dlstantes 223 Key to the Species of Series Dlstantes , 224 27. Hellconla llbrata 227 28. Hellconla lankesterl 234 29» HellconTa mollnaeana 239 30. Hellconla adflexa777 244 31 • HellconTa metalll'ca 249 32. Hellconla osaensls . 255 33« Hellconla llnnaeana 26O 34. Hellconla schledeana 265 35 • HellconTa yeracruzll 274 36. Hellconla latlspatha . , 278 37« Hellconla tortuosa 291 38. HellconTa Irrasa .T 299 Page

HYBRIDIZATION 305

1 . Heliconla X dresslerlana 306

2. Heliconla X glllyana . 309 3 • Heliconla X mooreana . .•«.•..••.• 311 h, Heliconla X procera 313 5. Heliconla X woodsonlana 316 6. Heliconla X zacapaea . 318 APPENDIX

I. New Names and Combinations 321

II. Doubtful and Rejected Names 322

III. Lists of Species by Country 325 IV. A Tentative List of South American Species That Do Not Occur in 328

V. Presentation of Form for Field Study 331

LIST OF REFERENCES 338

BIOGRAPHICAL SKETCH 3Vf

1 LIST OF TABLES

Table Page

I. Summary of Suborder Categories In Scitamlneae. . . . 1?

II. Chromosome Counts for Hellconla 33

vll LIST OP FIGURES

Figure Page

1. Key to the Families In the Order Scltamlneae, . . . 10

2. Floral Diagrams of Families In the Order Scltamlneae 21

3. General Phylogenetlc Tree for the Species of Hellconla 39

I

vlll LIST OP MPS

Map Page

1. of Distribution Helicon la mariae , ^ 52

2. Distribution of Helicon la reticulata 57

3. Distribution of Heliconia Imbricata 63

Distribution of Heliconia caribaea 75

5. Distribution of Heliconia bourgaeana 82

6. Distribution of Heliconia wagnerlana 90

7. Distribution of Heliconia blhai 97

8. Distribution of Heliconia marglnata 106

9. Distribution of Heliconia colgantea Ill

10. Distribution of Heliconia nutans 116

11. Distribution of Heliconia all en 11 121

12. Distribution of Heliconia vellerigera 126

13. Distribution of Heliconia magnif lea I30

14. Distribution of Heliconia curtlspatha I35

15. Distribution of Heliconia xanthotricha 1^*0

16. Distribution of Heliconia paulil 11*5

17. Distribution of Heliconia longa I53

18. Distribution of Heliconia coll ins iana 164

19. Distribution of Heliconia catheta I7I

20. Distribution of Heliconia pslttacorum 180

21. Distribution of Heliconia aurantlaca I87

. it Map Page

22. Distribution of Heliconla leucoflora I93

23. Distribution of Heliconla hlrsuta 201

2^*. Distribution of Heliconla vaginalis 212

25. Distribution of Heliconla trlnldatls 21?

26. Distribution of Heliconla swart zlana 222

27. Distribution of Heliconla libra ta 233

28. Distribution of Heliconla lankesteri 238

29 • Distribution of Heliconla mollnaeana 2'*^3

30. Distribution of Heliconla adflexa 2kQ

31. Distribution of Heliconla metallica 254

32. Distribution of Heliconla osaensis 259

33 • Distribution of Heliconla llnnaeana 26k

3t*. Distribution of Heliconla schiedeana 273

35« Distribution of Heliconla 11 277

36. Distribution of Heliconla latispatha . . « 290

37. Distribution of Heliconla tortuosa 298

38. Distribution of Heliconla Irrasa JOk LIST OP PLA.TES

PLATE Page

1. HELICONIA MRIAE Hook, f 49

2. HELICONIA RETICULATA (Griggs) Wlnkl 55

3. HELICONIA IMBRICATA (0. Ktze.) Baker 60

4. HELICONIA CARIBAEA Lam 69

5. HELICONIA BOURGAEANA 0. G. Peters 78

6. HELICONIA WAGNERIANA 0. G. Peters 85

7. HELICONIA BIHAI (L.) L 93

8. HELICONIA I4ARGINATA (Griggs) Plttler 10^+

9. HELICONIA COLGANTEA R. R. Smith, sp. nov 109

10. HELICONIA NUTANS Woodson 11^4-

11. HELICONIA ALLENII Standi. & L. 0. Wms 119

12. HELICONIA VELLERIGERA Poepp 12^1

13. HELICONIA MAGNIFICA Morton ex R. R. Smith, sp. nov 128

li^. HELICONIA CURTISPATHA 0. G. Peters 133

15. HELICONIA XANTHOTRICHA R. R. Smith, sp. nov I38

16. HELICONIA PAULII Lane ex R. R. Smith,

sp. nov , IU3

17. HELICONIA LONGA (Griggs) Winkl. var. LONGA I5O

18. HELICONIA COLLINSIANA Griggs var. COLLINSIANA 157

19. HELICONIA CATHETA R. R. Smith, sp. nov. var. CATHETA 168

XI PLATE Page

20. HELICONIA PSITTACORUM L. f 177

21. HELICONIA AURANTIACA Ghiesbr 183

22. HELICONIA LEUCOFLORA R. R. Smith, sp. nov 190

23. HELICONIA HIRSUTA L. f. var. HIRSUTA 197

24. HELICONIA VAGINALIS Benth 20*^

25. HELICONIA TRINIDATIS Lane ex R. R. Smith. sp. nov... 21^

26. HELICONIA SWARTZIANA Roem. & Schult 220

27. HELICONIA LIBRATA Griggs 229

28. HELICONIA lANKESTERI Standi 236

29. HELICONIA MOLINAEANA R. R. Smith, sp. nov 2*^1

30. HELICONIA ADFLEXA (Griggs) Standi 246

31. HELICONIA METALLICA PI. et Lind. ex Hook 251

32. HELICONIA OSAENSIS Cuf 257

33. HELICONIA LINNAEANA Lane ex R. R. Smith, sp. nov «. 262

, 34. HELICONIA SCHIEDEANA Kl. var. SCHIEDEANA '^t. 267

'35. HELICONIA VERACRUZII R. R. Smith, sp. nov. . . , .T. '275

36. HELICONIA lATISPATHA Benth 281

37. HELICONIA TORTUOSA Griggs 293

38. HELICONIA IRRASA Lane ex R. R. Smith, sp. nov 300

Hybrids

39. HELICONIA X DRESSLERIANA R. R. Smith, hybr. nov 308

40. HELICONIA X GILLYANA R. R. Smith, hybr. nov. 310

xli PLATE Page

^H. HELICONIA X MOOREANA R. R. Smith, hybr. nov 312

^2. HELICONIA X PROCERA R. R. Smith, hybr. nov. 3I5

U3. HELICONIA X WOODSONIANA R. R. Smith. hybr. nov 317

HELICONIA X ZACAPAEA R. R. Smith, hybr. nov 319

I

xlll INTRODUCTION

Hellconla Is a tropical American genus consisting of

approximately 100 species, and Is In the order Scltamlneae

(Zinglberales ) . This group of monocots has long attracted

the attention of botanists primarily because of Its unusual

and strikingly handsome Inflorescences. The are

often cultivated In greenhouses and tropical gardens throughout the world.

The genus Hellconla Is easily recognized, but the

species within the genus need a careful study which is

long overdue. Identifications by published keys are

difficult and often erroneous. The present work treats

species of Central America, , and the West Indies,

the region collectively known as Middle America. It

entails a revision of slightly less than half of the number

of species in the genus. About ten taxa have distributions extending from Central America to .

A few species of Hellconla occur in eastern Malaysia

(van Steenis, 195^). The study of these South Pacific species is not within the scope of this work.

Material has been studied from the following herbaria whose listing is alphabetical by abbreviation, with corre- sponding names indicated in Index Herbnrlorum (1959).

1 2

DUKE Duke University

P Chicago Natural History Museum

FIAS Agricultural Experiment Station University of Florida

FSU Florida State University

GH Gray Herbarium, Harvard University

MICH University of Michigan

MO Missouri Botanical Garden

NY New York Botanical Garden

UC University of , Berkeley

US National Museum, Smithsonian Institute

WIS University of Wisconsin

Photographs of pertinent type specimens have been sent from:

B Berlin: Botanlsches Museum, Germany r K Kew: Royal Botanical Gardens, Great Britain

M Munich: Botanlsche Staatssammlung, Germany

P Paris: Museum National d'Hlstolre Maturelle, Laboratoire de Phanerogamle, France

W Wlen: Botanlsches Institut und Botanlscher Garden der Unlversltat, Austria

Representative specimens have been cited at the end of each discussion of species. These include cultivated, as well as native specimens. However, In the m«?pplng of the species, the cultivated specimens were omitted. HISTORY

In 1703 Plumler described three taxa of Hellconia in pre-Llnnaean polynomial form as follows:

Blhal ampllsslmls follls, florum vasculls cocclneis. Blhal ampllsslmls follls, florum vasculls subnlgrls. Blhal ampllsslmls follls, florum vasculls varlegatls.

The nomenclature history of the genus, however, actually begins with a description by Linnaeus in his

"Species Plantarum" (1753)t in which he named the

Nusa Blhal , and assigned three varieties to the species.

These varieties were the same as Pltimlers' taxa, "... varlegatls," "...cocclneis," and "...subnlgrls." This order of names differed from that of Plumler (I703).

Plumler (1755) continued using his pre-Llnnaean polynomials

In the original order.

Linnaeus first used the name Hellconia in the

"Mantissa Plantarum" (I77I). In doing so he created a

new genus to which he transferred his Blhal . This now well-known generic name Hellconia was used by most

19th and 20th century botanists. Hellconia , however, was not the earliest generic name devoted exclusively to these plants, for Miller (175^) had redescrlbed Linnaeus' plant

3 under the generic name Blhai. Hellconla was the more notable name, but Blhal was used by 0. Kuntze (I89I) and

H. F. Griggs (I90if ,1915). Hellconla bec^jne a nomen generlcum conservandum at the International Botanical

Congress of I9O5, Vienna.

The generic name Hellconla Is derived from the word

Helicon, which Is the name of a mountain In central Greece.

In Greek mythology Helicon was the home of Apollo and the

Muses. According to some etymologists (Ghlesbreght , 1862),

It Is probable that Linnaeus desired to relate two taxa

with the names Musa and Hellconla , since the two show strong resemblances in their habit.

As new species were published, botanists discovered that some species were more closely related than others.

0. Kxmtze (I89I) suggested a subgenerlc term, Taenlostrobus, for species with overlapping bracts. The most complete work of the 19th century on the genus Hellconla was done by J. G. Baker (I893) in which he wrote a synopsis of the genera and species of Museae. In this work he presented the subgenus Platychlamys for the deep, boat-shaped branch

-bracted species and the subgenus Stenochlamys for the shallow, boat-shaped branch-bract ed species. He did not utilize Kuntze 's suggestion of a subgenerlc category.

One of the foremost early 20th century workers on the genus Hellconla was R. F. Griggs (1903,1904,1915), 5

His study of the genus was not monographlcal, but he did

suggest Various subgenerlc categories, some of which are used In this taxonomlc revision. He also. In his study

of the genus, named several new species.

Restricted treatments have been made on Hellconla

In various regional floras. Those which are of concern

Include Petersen (I89O), Standley (I928.I937) and Woodson

and Schery (19'+5)» These authors also have described new

species, and have contributed much to a better understanding

of the genus.

No monographic treatment has yet been completed for

the entire genus Hellconla . One recent worker, I. Lane,

began a taxonomlc study of the genus about 1955 • For

reasons \inknown to this author, his study was abandoned,

and the work that had been done remains unpublished.

The most recent work on the genus Is an unpublished

ecological study of the genus Hellconla of Costa Rica by

Sheffy (1965).

I ECONOMIC VALUE

The genus Hellconla has long enjoyed widespread culti- vation, because of Its very tropical nature and showy

Inflorescence. Early botanical explorers of the tropics

returned to Europe with several species of Hellconla to be grown In greenhouses. Many of these plants were described as new species, and were among the more popular

plants of European horticulturists.

Since the habit of Hellconla Is similar to Musa , the

leaves of the former have been used In much the same fashion as those of the latter. The natives of tropical America

have several uses of these plants. Some of these uses

Include wrapping various Items In leaves for storage or

cooking. Leaves are also used as building materials,

providing new roofs and siding for homes. At times the natives apparently utilized the rhizome as a source of

food. The Inflorescences are often used In floral arrange- ments and as votive offerings.

Verbal reports have been made that certain species

of Hellconla are a weed problem In plantations.

Attempts to control the weedy condition would be a serious problem since the two genera are closely related.

6 The upright Hellconlas collect water in their branch

-bracts. Consequently, they will harbor mosquito larvae and other minute animals, k microbiological study of the life in these bracts would be an interesting contribution, analogous to one made on the tropical bromellads (Laessle,

1961). SURVEY OP THE ORDER SCITAMINEAE

The genus Hellconla Is In the natural order of mono-

cotyledons, Scltamlneae (Hutchinson's Zlnglberales ) . As

Hutchinson (1959) points out, it is "a very advanced group

representing the climax of one line of developnent of the

division in which the calyx and corolla have remained in

separate whorls; often regarded as prototypes of orchids,

but here considered to be a parallel group, with a similar

reduction to 1 stamen, a very different stamen from that

of the orchids. In addition the leaf-sheaths of orchids are never ligulate." The general characters of this order are as follows:

"Herbs with rhizomes; leaf-sheath usually open and

with a ligule: calyx and corolla in separate whorls;

stamens 6-5, or often only 1, the remainder staminodes

and petaloid; ovary inferior; seeds with endosperm, often arillate." k brief systematic survey with a key to the families is

provided to show the family relationships within the order and indicates why the present tendency is to place the

Senus Heliconia into its own monogeneric family, ...... ^ -., . , , ^ 9

Hellconlaceae. The key In Fig. 1 Is a modification taken froD Hutchinson (1959).

The following descriptions of the order and families will facilitate an understanding of their relationships.

Order Scitamineae

Usually large herbaceous plants with well-developed rhi- zomes; large glabrous leaves with leaf-sheaths, often open and with a ligule, blades with simple pinnate venation; inflorescence usually showy; flowers unisexual or perfect, zygomorphic or asymmetrical; calyx and corolla in separate whorls; androecium derived from two whorls; stamens free numbering five or six, often one is fertile, others being represented by petaloid staminodes or absent; ovary infe- rior, three carpels, generally trilocular with one to several ovules per locule; style simple, stigma terminal; fruit a berry or capsule; seeds usually with endosperm and often arillate; embryo straight or curved. 10

« 00 0) 4) «> CD m o 0) 0) 0) u o cs 0] 4) OS a> « 0) N •H OS U OS o P «) a> 4) o o OJ W

1 1 vi 1 1 w 1 1 •d 1 1 PI ^ c 1 tt) o . M 1 1 CC o 1 CO «H g 1 1 0) O CO o +> 4J 1 rH 1 O (I) rH 05 1 M 7i (0 N 0) M o a to 1 CO 0) •H ^1 CO o 1 x: .. to

II 1 ce ;3 O -P 4-> CO CO to 1 " V) xi 03 tt) . > 3 - 0) w 1 o Q) -P rH rH rH 1 -a: •p Q) X o -H •H 0) CO w 1 >» ^< P< c O >i O ^4 1 »- -• «^ 0 :3 Pi Q> O O rH o o M j:; »-( M 1 ^ 0) CC CO CO a 1 o CO Ei CC -p j:: 0} a> tt) crj to 1 o rH a ^ to to to 1 a o aT o C 0) C 0) . Ct< iH ho •P ho u CO >» -H -d •H -d 1 Cd rH t: C to to ;Q o o rH •H -H CO tt) Ctf >i C >> c 1 U O ES rH rH o H C to p< H O >H rH tt) tt) trt rH o crt rH CO -H 0) a. tt) 4J 0) rH ci tt) rH CO CO CC OS OS o CC -H a > p» 0) rH C3 a rH CO C 4-> o a-d CO a P -P P -P G •H 4-> tt) 0) 0) O 0) c tt) o > 0) o > tt) O •H 0) to ce P •H ^ CO a o »-J CO o a o O Q. O CO ^^ P< -d j > tt) CO CO 03 rH O ct! o -H H rH ^^ H =J ;Q Otf >: > O o o O o o H CO CO rH

C E] >> tt) I H CNJ CO » CO CO CO > U tt) CO 0) tt) j:: 0) 0) O 0) -d »H 0) t> X C tt) X cC O CC O 4J 2c -p H .c p «) rH rH tt) iH 0) c CO -P t-l V-i V< a C f^ tt) CO CO hO U rH • 3 » CO o »n Pt tH 0) O cC -d H CO to rH CO o I »H :i c c CVJ 0) 0) o tt) ^ £ o 9 P rH o •P C I 4J -P to oJ cvj to to 11

Family Musaceae

Large erect perennial herbaceous plants with stem formed by

Imbricated bases of petioles; leaves spirally arranged, large, with thick mld-rlb; Inflorescence generally compound, not usually terminal, subtended by large bracts, Flovrers usually unisexual, male flowers within upper bracts, female flowers within lower bracts. Calyx elongated, at first tubular, then splitting on one side: corolla often trun- catci Stamens 6, 5 fertile, a rudimentary sixth stamen.

Ovary trllocular with many anatropous ovules, nectar secreted in septal glands.. Fruit a berry, seeds hard, often arlllate. Embryo straight.

Representative general

Musa ^sete

Family Hellconlaceae

Small to large perennial herbaceous plants; leaves dls- tlchously arranged or seemingly so; with medium to large blades; Inflorescence terminal.: Flowers bisexual, arranged

In a clnclnnus In the axial of the bract or spathe. Calyx trlmerous, corolla trlmerous, perianth segments partially united. Stamens five fertile with sixth Imperfect and petalold^- Ovary Inferior with one basal ovule In each loculus. Fruit fleshy, a schlzocarplc berry, splitting

Into three one-seeded parts; seed not arlllate. t

Monotyplc genus

Hellconia

Family Strelltzlaceae

Herbaceous perennial, or with woody stem as In R. mgdagas-

carlensls and S. nlcolal ; leaves variable in size, dls-

tlchously arranged; Inflorescence terminal ( Phenakospermum )

or not terminal ( and Strelltzla ) » Flov/ers bisexual, arranged In a clnclnnus of axial of a bract or

spathe. Calyx trlmerous, petals trlmerous, sometimes unequal; all perianth parts free. Stamens five or six;

if five, all fertile; if six, the sixth imperfect and

petalold* Ovary inferior with many ovules i Fruit capsular;

seeds arlllate with straight embryo.

Representative genera:

Strelltzla Phenakospermum Ravenala

Family Lowiaceae

Acaulescent ; leaves with petioles extended

from imbricated sheathes; Inflorescence terminal, branched

cyme* Flowers bisexual, very zygomorphic, in bracteate

cymes from base of leaf-sheath; bracts 1-2 flowered.

Calyx trlmerous, united below into slender tube* corolla

trlmerous very unequal, two lateral small, third middle

one forming large colored lip (labellum). All floral parts j

13

united Into slender column above the ovary. Stamens five, sixth opposite the lip lacking. Ovary Inferior with numer- ous ovules. Prult woody capsule, seeds with 3-lobed aril.

Monotyplc genus

Orchldantha

Family Zlnglberaceae

Perennial aromatic herbaceous plant. Stems sometimes short, leafy, bearing only flowers; leaves distichous, with open sheath at base; blade sessile or petloled. Flowers solitary or In distinct Inflorescence; mostly bisexual, symmetric or asymmetric. Perianth 6-merous, 2-serlate, outer calyx-like segments united Into a tube. Inner corolla-like segments more or less united, showy, the posterior segments usually the largest. Stamens 1, 2-locular anther sometimes accom- panied by petalold stamlnodes. Ovary Inferior, 3-'locular, with axlle placentas, or 1-locular with parietal or basal placentas; style terminal, free or more or less enveloped

in a groove of fertile stamen. Ovules mostly numerous.

Fruit fleshy and dehiscent; seeds round or angular, mostly covered with large divided aril; endosperm abundemt.

Representative general Hedychlum Kaempf era purcuma Renealmla Alplnia Zingiber Amomum goscoea j •

Family Costaceae

Perennial herbs, not aromatic j stems with leaves spirally arranged, sheaths closed, occasionally opened only by growth. Flowers solitary or in a distinct Inflorescence accompanying or separate from leaves, mostly bisexual, symmetric or asymmetric. Perianth 6-merous, 2-serlate, outer calyx-like, Inner corolla-like and usually showy, the outer segments united Into tube. Stamens 1, with

2-locular anther, lateral stamlnodes often absent or tooth-like, ovary Inferior, 3 or 1-locular; style terminal, free; Stamen sometimes 2-llpped or dentate. Ovules numer- ous. Fruit fleshy and Indehlscent. Seed round or angular, covered by aril.

Representative genera

Costus Monocostus Dlmerocostus Tapelnochllus Mulfordla

Family Cannaceae >

Tall, leafy, perennial herbaceous plant; leaves large, broad, plnnately nerved, with distinct midrib. Flowers racemose or paniculate, bracteate, zygomorphlc, bisexual, mostly large and brightly colored. Perianth double, outer calyx-like. Inner corolla-like, sepals three. Imbricate, free herbaceous. Petals three, connate at base and adnate to stamlnal column. Stamens petalold, shortly connate at base; three outer sterile, Imbricate, two Inner more or less connate, free; anther solitary, 1-locular, adnate to side of petalold portion. Ovary Inferior, 3-locular, ovules numerous on axile placentas. Fruit a capsule opening by the collapse or warted pericarp; seeds many, rounded with hard endosperm and straight embryo.

Monotyplc genus:

panna

Family Marantaceae

Perennial herbs; leaves In two rows, differentiated into an open sheath, stalk, and blade; stalk often winged, te- rete and pulvlnl form towards apex. Flowers bisexual, asymmetric, in a terminal bracteate spike or panicle, or

the Inflorescence arising from fhizome. Perianth mostly

differentiated into calyx and corolla. Outer segments

free. Inner more or less tubular, often divided into three

unequal parts. Fertile stamen one; anther 1-locular;

staminodes variously petalold. Ovary inferior, 3-1-locular,

sometimes two of the locule infertile; ovule solitary,

erect from base of locule. Fruit fleshy or locullcidal

capsule; seed with abundant endosperm and much incurved

or folded embryo, and often a basal, sometimes lamellate

aril. s

Representative general

Calathea Maranta Thalia

Tomlinson (I96I) produced an outline showing the order of families according to Interpretations of various workers In this field of study. This outline Is modified in TABLE I.

Eight major entities are in this order, and whether each group should receive equivalent status is debatable.

Tomlinson treats these as families. All authors, as seen on the chart, agreed in treating two taxa as families

— Cannaceae and Marantaceae. In the early 1900'

Zlnglberaceae is treated as having two sub- taxa, Costoideae and Zingiberoideae. Nakai (Tomlinson, I96I ) and Tomlinson

(1961) call these families Costaceae and Zlnglberaceae.

The Musaceae is the taxon which has undergone the most change in recent years. It consists of three groups, Musa and Ensete ; Phenakospermum , Ravenala , and Strelitzla ; and

Heliconla, Nakai (Tomlinson, I96I ) and Tomlinson (I96I) treat these respectively as Musaceae, Strelltziaceae and

Heliconlaceae. Hutchinson (1959) places Heliconla in Strelltziaceae.

The morphological characters which Heliconla shares

^i-th Phenakospermum . Ravenala and Strelitzla are the similar inflorescence consisting of clnoinnl In the axils of bracts, and five functional stamens. »

TABLE I

«

SUMMARY OF SUBORDER CATEGORIES IN SCITAMINEAE

Bentham & Engler & Engler Hooker, I883 Prantl, 1st ed. 1900-1912 Family 1889 Das Pflanzen- Scitamlneae Relhei Scitamlneae relcht Scitamlneae

Tribe Museae

Museae Musaceae Musaceae (Petersen, I889) (Schumann, 1959) Tribe (Winkler, 1930) Hellconleae

Zlnglbereae Zlnglberaceae Zlnglberaceae (Schumann, 1959) (Loesener, 1930)

Canneae Cannaeeae Cannaceae

Maranteae Marantaceae Marantaceae )

16

TABLE I (extended)

Engler & Prantl, 2nd ed. 1930 Hutchinson Tomllnson oTiH Naturllchen 1959 1961 Pflanzenfamlllen Zlnglberales Scltamlneae

SuDfamlly Musoldeae Musaceae Musaceae Subfamily Strelltzloldeae Tribe Strelltzleae Strelltzlaceae Strelltzlaceae (Strelltzla. (Strelltzla. Phenako- Phenako- spermum^ spermum, Tribe Ravenala. Ravenala Hellconleae Hellconla) Hellconlaceae Subfamily Lovrloldeae Lowlaceae Lowlaceae

Subfamily Zlnglberoldeae Zlngl^eraceae Zlnglberaceae

Subfamily Tribe Costoldeae Costeae Costaceae

Cannaeeae Cannaceae

Marantaceae Marantaceae Heliconla differs from these above mentioned genera In a number of ways. The most obvious of the differences Is the habit. Heliconla Is never arborescent. Other differ- ences are as follows (Tomllnson, 196l)j

Heliconla has but one ovule In each locule within the

ovary; whereas, Strelltzlaceae has several.

Heliconla develops a schlzocarplc-berry type fruit

with exarlllate seeds; Strelltzlaceae develops a

capsule-type fruit with arlllate seeds.

Heliconla produces a poorly differentiated embryo

at time of seed maturity, as compared to the embryo

In Ravenala and Strelltzla .

Heliconla has an unusual orientation of the flower. This

can be observed In Fig. 2, consisting of floral dlagrsuns

of families In the order Scltamlneae. In Hellconlaceae

the adaxlal sepal Is single and free; whereas. In the

other faunllles a pair of sepals occur adaxlally and the

third sepal Is located abaxlally. This peculiar type of

orientation appears to be associated with its mode of

pollination (Rendle, I93O). Further comparative studies

need to be made on pollination mechanism to clarify this

unique floral morphology. .

Fig. 2. Floral Diagrams of Families In the Order Scltamlneae

A. Musaceae and Lowlaceae; a, stem axis; b, position of rudimentary stamen In Musaceae or position of absent stamen In Lowlaceae; c, sepal; d, petal; e, branch-bract.

B. Strelltzlaceae; a, stamlnode.

C. Hellconlaceae; a, stamlnode.

D. Zlnglberaceae and Costaceae; a, stamlnodal labellum; b, sheathing bract.

E. Cannaceae; a, petalold style; b, stamlnode; c, labellum.

F. Marantaceae; a, fertile stamen; b, stamlnode; c, solitary ovule; d, hooded stsunlnode. 21 Upon reviewing the various works on the order

Scltamlneae, It Is the conclusion of the author that the genus, Hellconla . with Its distinguished floral orientation, fruit-type and exarlllate seed should be placed In Its own family, Hellconlaceae, MORPHOLOGY

Information concerning the morphological aspects of

the genus, Hellconla . comes from the study of living plants, preserved material, and herbarium specimens.

Habit i The habit of Hellconla is often termed as musoid

(bsuiana-like ) or cannoid ( canna-like ) . These terms indi- cate that the plant is an erect herb ranging from about one-half meter to more than 12 meters in height with various sized leaves. When the large leaves are long

-petioled, with oblong to oblong-elliptic leaf-blades, the plant habit is said to be musoid. The cannoid habit is evident when smaller and narrower leaf-blades are more or less sessile to the leaf-sheath making the plant appear slender.

All the species studied have a well-developed rhizome, and are perennial. The modes of propagation are by the formation of and dispersal of seeds, and by the formation of new plants from the rhizome.

The vegetative structures are similar in appearance and growth to that of Musa . Therefore, much of the termi- nology in reference to these structures is that of Slmmonds (I960).

23 Roots t The root system Is clpisslfled as an adventitious

one. The seedling root dies early, and Is immediately

replaced by adventitious roots. From these long adventi-

tious roots many secondary roots develop.

Rhizome ; The rhizome is active in vegetative propagation

There have been no studies on the rhizomal development in

Heliconia . However, it is like that of other monocots,

which is sympodlal. Horizontal growth is slight, and as

soon as the rhizome extends beyond the parent plant it

forms a new aerial stem. Similar to Musa , the new buds

that arise are borne in the middle and upper parts of the

rhizome.

Stem: The stem or trunk is formed from the imbricated

sheaths of the leaves. Thus, it is correctly termed a

pseudostem. It will vary in thickness at the base in

direct proportion with the height of the plant. The two

extremes demonstrating this phenomenon are H. psittacorum

and H. mariae .

Leaves ; The leaves consist of the leaf-blade, petiole

when present, and a leaf-sheath.

The leaves form. from the white merlstematic tissue

of the center of the new pseudostem. The first leaves are termed scale leaves. Subsequent development produces

the typical leaf. In which the sheaths forming the pseudostem are completely enclosed. Later, they develop a free margin by continual growth In the heart of the pseudostem. In section, the sheath Is crescent-shaped tapering at each side to a thin papery margin. The top of the sheath Is contracted Into a petiole. The length of the petiole varies. The leaves of some species are sessile or nearly so, an example being H. vaginalis .

Others have a very long petiole, for example H. latispatha

The blade of the leaf Is composed of two halves, usually equal In appearance, separated by the tapering midrib. The base of the blade Is rounded or acximlnate, the apex Is often acuminate.

The development process of the Musa leaf-blade has been studied by Skutch (Slmmonds, I96O). The Hell con la leaf develops In the same manner. When It emerges from the pseudostem the blade (lamina) appears tightly rolled, the right half rolled upon Itself and the left half rolled upon the right half and midrib. It starts unrolling at the top, and progresses downward as the leaf base elongate and clears the blade from the pseudostem.

The leaf-blade Is of taxonomlc value 'In several

species. It Is sometimes glaucous as In H. colllnslana ,

maroon with reticulated cross veins In H. reticulata , and waxy as In H. longa . 26

Inflorescence t The Inflorescence Is formed from the growing point of the stem in the heart of the pseudostem, and undergoes much of its development before emergence.

The inflorescence in Heliconia is terminal, and -is the last structure formed from the growing point of the stem.

The inflorescence consists of a peduncle, rachis,

branch-bracts, pedicels, floral-bracts (bracteoles ) , and flowers. The peduncle is the elongated stem-like structure to which the Inflorescence is attached. The rachis is the stem-like structure to which the branch-bracts are attached.

The branch-bracts in Heliconia are very well developed.

Depending on the species, the branch-bracts may be succu- lent or membraneous, hairy or glabrous, deep or shallow boat-shaped, and spiralled or distichous. Their width and length vary as does their coloration. Often the lowest bract is sterile, possessing no flowers. If it is sterile

it usually extends into a leaf-blade. All other branch

-bracts contain the floral-bracts and flowers which are arranged in a cincinnus on an extremely short, stout,

secondary rachis or rachilla.

The width of the branch-bract is the' greatest distance

across the top. This is usually one or two centimeters

out from the rachis, and is variable between species.

The side-width of the branch-bract is a term used for

the pressed specimens. It consists of a measurement of .

I 27

the greatest distance from the margin to the mld-polnt of the base. Likewise, this characteristic Is readily dis- tinguishable between species.

Many of the Inflorescences are pendent. This Is one of the characteristics which Is used In grouping the spe- cies In their sections and series.

The floral-bract, found at the base of the pedicel.

Is variable within the genus. It Is either deciduous or persistent, It is usually quite thin, membraneous and of a pale color. In species which have a hairy branch-bract, the floral-bract Is also very pubescent, especially along the midrib on the outer surface.

Flower : The flowers In the genus Hellconla are morpholog- ically consistent. They vary In length, size and color, but often they are not readily distinguishable.

Two of the three lanceolate sepals making up the calyx are partially connate. These are called the abaxlal sepals

(see Fig. 2, 6). The adaxlal sepal Is free. This sepal Is also called the unpaired sepal. In few species the unpaired sepal curls adaxlally at anthesls. An example of this

phenomenon Is found In H. latlspatha and H . blhal

The three lanceolate petals are adnate except for the pair of adaxlal ones. These have a free margin. The free margins of these adaxlal petals are opposite the single adaxlal sepal. The length of the flower varies from about 2.5 cm. to 6.5 era. The calyx and corolla are of the came length.

If the species Is of a pubescent nature, the calyx Is also usually pubescent. The corolla Is glabrous In all species.

Androeclum ; The androeclum consists of five separate functional stamens with a sixth sterile one opposite the unpaired sepal. The sterile stamen Is termed a stamlnode.

The size and length of the short petalold stamlnode appar- ently varies In species. It Is suspected that the staml- nodes would be a good taxonomlc characteristic, but they could not readily be studied in herbarium specimens. The f\inctlonal stamens consist of the typical thin, long filaments with the elongated terminal anthers, which make up the upper one-fourth of the entire structure. The length of the stamens is variable among species, but they consis- tently are the same length or extend slightly beyond the perianth.

Gynoeclum ; The gynoeclum consists of 3 carpels. The terminal capitate or slightly lobed stigma Is attached to a style, which Is straight or slightly geniculate at the apex. In turn the style Is attached to a three-loculed.

Inferior ovary. There Is only one anatropous ovule In each locule. The length of the style Is equal to the length of the functional stamens. Fruit t The fruit Is a one- to three-seeded fleshy, sub- globose structure. It has a diameter of about 1.5 cm. or slightly more when fleshy, and 0.7 to I.5 cm. when dried. The coloration of the fruit varies. It begins a green or yellow In the Immature fruit, and changes to a dark blue at maturity. The fruit Is called a berry by

Lane (1955 )t but previous workers erroneously labelled It a schlzocarp or capsule (Rendle, 1930). A. more correct term would be schlzocarplc berry, since It Is fleshy, and yet It does split at maturity.

Seeds t The seeds are of various sizes and shapes In the different species. They are usually of a three-sided, somewhat almond-shaped form. Two sides are flattened, and the third Is rounded. The size ranges from about

0,k - 0.9 cm, diameter and 0.? - 1.0 cm. In length. They all possess a stony seed coat, which Is usually roughened.

They do not have an aril.

Embryo { The embryo embedded In the endosperm, Is straight and oriented In the longitudinal axis of the seed (Lane,

1955). POLLINATION

Hellconla is considered to be a " flower."

There are verbal reports of visiting various species of Hellconla . R. L. Dressier, herbarium sheet

#1^189 (US), noted that he saw the hummingbird, Fhaethomls

superclliosus , visiting a population of Hellconla located

In a limestone area, near Laguna Ocotal Grande, State of

Chiapas, Mexico. Sturgls (1928) found that several species of hummingbirds visit patches of wild plantain, a common name for Hellconla .

Closely related to the phenomenum of pollination Is the quantity of nectar. Hellconla Is In the group of calyclferous (those with sepals and petals).

Like the genus Musa, It secretes large quantities of nectar within the tubular flowers (Perclval, I965). The humming- bird, utilizing the nectar as one of Its staple foods, comes In contact with the pollen while sipping.

The Hellconla Inflorescence satisfies the requirements of belonging to the group of hummingbird flowers. It

secretes large quantities of nectar j the peduncle, rachis, and pedicel are sturdy, and the tissues of the flowers and branch-bracts are moderately thick and coriaceous. The

30 31

ovaries are protected by being Inferior, and located within the brnnch-bracts supplied with additional protection by the floral-bracts.

Undoubtedly, several species of hummingbirds visit

Hellconla . There may also be a number of other factors

Involved In pollination. Pollination studies would be most enlightening In relation to speclatlon. Further

Investigations and observations definitely need to be made for a better understanding of the genus. CYTOLOGICAL ASPECTS OF HELICONIA

Although this author has not performed cytologlcal investigations to date, a few studies have been made on

the genus Heliconia *

Chromosome counts for Helicon ia listed in Darlington and Wylie (1955) are seen in TABLE II.

The Chromosome Number Reports in Taxon, presented by

Love (1967), has the following chromosome count for

Heliconia t

Heliconia caribaea n=ll India: West Bengal Mitra & Datta Indian Botanic Gardens Slbper, Datta 9^81, CBLH

A. K. Chakravorti (I96O) mgde an interesting study on the cytogenetics of Heliconia . He worked with certain species cultivated in India, attempting to find out whether fragmentation of chromosomes played a role in the evolution of the species.

32 ntS • U M O o O •H >H X) OS • • o a > •H M H N < n iH H o o c • ?f 0) m Q w M O CO CO

NO ro NO ON -:} CJN CjN ON J- ON CJN CJN CjN On r-i o • m • CJN CQ 05 M «« ^3 ,o w oa -p an an u to to o CO to S a OS 05 > 05 a a 03 « 0) 0) •p 0) P 05 p 9) p o u w to to U to to CS 0) 0) (U 0) 0) 0) C 05 0) 0) CC bO A 0) 0) 0) E-t -IS o o > 6 > 6 > > O O w CJ o oCO o o CVJ CC CM CM CM (M CM CM CM CM B 00

NO

05

05 •H

•P 0) to to OS I -p CO U o o 05 0) t-i o to 0) Qi o 0) to p 05 •H to p 05 o 05 >• (0 p U 0) 0) 05 (0 ^ m u to CO JO 5 p. Chakravortl listed his species ast

Species Chromosome number

1. Hellconla metalllca 22

2. Hellconla sp, I 22

3» Hellconla pslttacorum Zh

' 4. Hellconla blhal Zk

5» Hellconla rubra 2^

6. Hellconla braslllensls Zk

7. Hellconla sp. II ZU

His study concludes that the 22-chromosomal species

Is the more primitive, and that 2^- chromosomal species arose from the 22-chromosomal species by fragmentation of a particular pair of chromosomes at a secondary constriction region.

It Is unfortunate that there was no mention of voucher specimens. Hellconla sp. I may be H. latlspatha according to his brief description of vegetative characteristics.

H. pslttacorum Is the name that has been used for many of the smaller taxa. The name H. rubra Is a source of confusion. .

PHYLOGENY

Phylogenetlc considerations have not been previously

attempted with the genus Hellconla . Attempts of this nature, when considering a genus In part or on a regional basis, are difficult. In this study all of the major morphological features are found In the species of Central

America and the West Indies with the exception of H.

eplscopalls »

The phylogeny of Hellconla Is best measured by the development characters of the Inflorescence. All species

start out erect and somewhat Imbricated. From this stage

there Is great diversity In the formation of the mature

Inflorescence

There are several natural groups within Hellconla .

These groups are listed below, and are followed by an

arrangement of a phylogenetlc tree (Fig. 3)» The letter

used for the listing below Is repeected within the circles

on the phylogenetlc tree.

A. Branch-bracts numerous. Imbricated; Inflorescence

erect or pendulous i H. Imbrlcata, H. reticulata ,

H. eplscopalls (A- ), and H. marlae (A«).

35 36

B. Branch-bracts nearly Imbricated, puberulent; Inflores-

cence erect: H. carlbaea •

C. Branch-bracts somewhat separate, thick; inflorescence

. erect: H. bihai , H. bourgaeana , H. wagneriana

D. Branch-bracts numerous, closely spaced and reflexed;

inflorescence pendulous: H. curtispatha , H. longa ,

H. vellerigera , and H. xanthotricha .

E. Branch-bracts numerous, more or less reflexed and

widely spaced; inflorescence pendulous: H. catheta ,

« H. collinsiana , H. magnif ica, and H. paulii

F. Branch-bracts few, widely spaced; inflorescence

pendulous: H. allenii , H. colgantea , H. marginata ,

and H. nutans ,

G. Branch-bracts several, close together, curved upward;

inflorescence erect: H. adflexa , H. lankesteri , H.

librata , and H. moling eana »

H. Branch-bracts few, moderately to widely spaced;

inflorescence erect; petiole short: H. linnaeana ,

H. metallium , and H. osaensis .

I. Branch-bracts few to several, widely spaced, rachis

twisted; inflorescence erect: H. irrasa , H. latispatha,

H. schiedeana , H. tortuosa , and H. veracruzii .

J. Branch-bracts few; inflorescence erect; leaves petioled:

H« psittacorum* K. Branch-bracts fewi Inflorescence erect; leaf-blade

sessile or subsesslle: H. aurantlaca , H. hlrsuta ,

li' l^ucoflora , H. vaginalis , H. trlnldatls , and

H . swartzlana *

As seen by the phylocenetlc tree In Fig. 3, It Is thought that the Imbricated, somewhat cone-like Inflores- cence Is the most primitive. This Is In accordance with the statement of Bessey (I915), that polymerous flower structure preceeds the ollgomerous ones. It is followed by the assumption that imbricated species gave rise

Independently to the pendulous and the erect species.

The most advanced form would be the erect plant with few narrow branch-bracts in which lie a few flowers.

Along the pendulous branch of the tree, (D), (E), and (F), the number of branch-bracts and flowers per bract becomes substantially less.

The large fleshy branch-bracted species (B) and (C) gave rise to the remainder of the erect species. These erect species vary in number of branch-bracts, as seen in the distichous species, (I). These distichous species gave rise to those with a spiralled rachis, (H), and those with the candelabrum effect, (G). (J) and (K) represent the smaller species with few branch-bracts. Phylogeneti- cally, species of (J) retain the petiole and those of (K), having lost the petiole, possess a sessile leaf-blade. General Phylogenetlc Tree for Species of Hellconla.

Sections Indicated by dashed lines.

I. Section TAENIOSTROBUS

Above the dotted line (A «&; A2) Is Series Taenlostrobus, and below the dotted line (A^ ) Is Series Eplscopales,

II. Section HELICONIA

III. Section PENDULAE

IV. Section STENOCHIAMYS Above the dotted line (J & K) Is Series

- , Stenochlamys, and below the dotted line (G, H, & I) Is Series Dlstantes.

SYSTEMATIC TREATMENT

HELICONIA L., Mant, 2» 14?. 1771.

Typei Helioonla bihai (L.) !• The type apeolmen for thla

species Is In the Llnnaean herbarium. It Is listed by

Savage (19^*5) as follows i 286,1 Helioonla Blhal Surln,

80.

There had been confusion Introduced by Griggs (I903) and others, as to which species Is the type of Helioonla .

As Hodge {19^1) pointed out, there are two entitles growing together In the British West Indies, which Is the location where Linnaeus probably obtained his specimens.

Linnaeus' first description of his plant. In Species

Plantarum 101^3 (1753)t under the name Musa Blhal, Is

"ampllsslmls follls, florum vasculls varlegatls." This

Is followed by the variety "ampllsslmls follls, florum vasculls, cocclnels." It Is now definitely understood that the species with the variegated bracts Is H. blhal

(L.) L. and the species with the solid colored bracts, whether they be the more common red or the less common green or yellow, Is H. carlbaea Lam. Blhai Miller, Gard. Diet. Abr. Ed. f. Vol It BI.

175^ • Types No type designated. The description

of the first species Is ''Blhal ampllsslmls follls,

floruffl vasoulls cooclnels." Both this description

and the name Blhal are pre-Llnnean, and originated

with Plumler (1703)* The plant represented by

this polynomial Is presently known as H. carlbaea

Lam.

Hellconlopsls Mlq. PI. Ind. . 3t 590. l«5b.

Type I No type designated. Hellconlopsls was

based on a plant from Ambolna, Indonesia* which

from the description may be H. blhal (L.) L.

Blhala 0. Ktze., Rev. Gen. PI. Zt 6tt4, lb91.

Type J B. luteofusca 0. Ktze. This Is a synonym

for H. blhal (L.) L. Griggs (190*^) suggested that

Kuntze added the extra "a" to Blhal In order to

further Latinize the spelling.

Coarse, perennial, herbaceous, erect plants of various heights (1-5 m., sometimes taller);" habit similar to Musa of Carma. Rhlzomatous with adventitious roots. Pseudostem composed of long sheathing bases of leaves. Leaves fre- quently large, distichous or give appearance of being distichous. Inflorescence terminal, erect or pendulous, consisting of few to many, usually brightly colored, cymbl- form branoh-braots, either distichous or spirally arranged. Branch-bracts subtend clusters of flowers, few to many. In close cinclnnus arrangement. Flowers subtended by floral

-bracts ( bract eoles ), usvially the length of the flower or a slightly longer, persistent or deciduous. Flowers perfect. Perianth consists of two whorls; calyx with

connate abaxial sepals and free adaxial sepal t corolla with connate petals except for free margins opposite adaxied sepal; stamens 6,5 fxmctional and one stamlnode which lies opposite adaxial sepal. Anthers linear.

Stigma lobed, clavate or subclavate. Style filiform, straight or geniculate. Ovary inferior; 3-loculed; single ovule erect from base of each locule. Fruit a

1 to 3-seeded berry (also called fleshy schizocarp splitting along 3 sides). Seeds with stony seed coat usually rough and often wrinkled; obtusely trigonous; embryo straight.

In the natural state, these plants are usually found in moist to wet forested areas, moist ravines, or they may form dense thickets along streams and rivers. They have an extensive vegetative growth produced from the activity of the rhl2ome. This results in large populations of a species forming clones, instead of scattered individual plants.

The geographic distribution shows that Heliconia is primarily a Central American and northern South American genus. There are nearly 100 species, 38 species from 43

Central America and approximately 45 species from South

America. A few species remge from Indonesia, New Guinea,

New Caledonia to Samoa, for which Miquel proposed the name

Hellconiopsls t He concluded that they differ considerably from the Western Hemisphere species, but a recent worker

(Lane, 1935) disputes this conclusion.

The subgenerlc categories in the following portion of the systematic treatment consist of sections and series.

As species of Heliconia were studied, they were placed

into four major natural groups. These groups are better treated as sections, and not subgenera, since they have not been proposed as separate genera and thus do not approach this level. Likewise, smaller groiips, series, were recognized within two of the sections.

Type species for the subgenerlc categories, where applicable, were selected by the author.

Key to Sections

A* Branch-bracts forming closely-knit spike; lower fertile

branch-bracts 6-10 cm. long, coriaceous; inflorescence

completely imbricated at anthesls, rachis hidden

I. TAENIOSTRCBUS

A. Branch-bracts distant on rachis or Imbricated only at

base during anthesls; if Imbricated, lower fertile

branch-bracts 12-18 cm. long, thick; rachis usually

exposed, in few species partially hidden. t

B* Branoh-braots large* thlolc and deep boat- shaped}

inflorescence sessile or subsessile, with con-

siderable shrinking upon drying; rachis with

diameter of 1.0 cm. or more II. HELICONIA

B* Branch-bracts variable in size* not large and deep

boat-shaped; inflorescence usually peduncled,

without shrinking upon drying: rachis with

diameter of less than 1 cm.

C. Inflorescence pendulous; rachis usually

pubescent III. FENDUIAE

C* Inflorescence erect, nodding or slightly

drooping; rachis either pubescent or

glabrous IV. STEN0CHIAH7S

I. Section TAENIOSTROBUS 0« Ktze.. Rer. Gen. Fl. Zt 684.

1891.

Subgenus Platyohlamys Baker in part. Ann. Bot. Vol. VII

190. 1893.

Subgenus Taeniostrobus (0. Ktze.) Griggs, Bull. Torr. Bot.

Club 42i 316. 1915.

Stout, herbaceous, perennial plants, 1-12 m. tall; musoid habit. Leaf-blades large, all petioled. Inflores- ence erect or pendulous, somewhat pubescent > rachis straight or nearly so. Branch-bracts IO-7O, red; middle branch-bracts 4-10 cm. long. All branoh-braots closely imbricated. •

*3 I . 1

i

Type species I Hellconia Imbricata

0* Ktze*

This Is a small section consisting of four species.

The Inflorescence Is composed of many branch-bracts, In

which each branch-bract completely overlaps the preceedlng

one. The phenomenon gives the Inflorescence the appearance

of being a single large structure. One species, H. marlae .

Is commonly called the "beefsteak" plant, because the

Inflorescence has the appearance of a hanging piece of

meat*

Key to the Series

A* Branch-bracts persistent, extending out horizontally; Inflorescence composed of distichous branch-bracts....

Imbrlcatae

A« Branch-bracts deciduous, ascending; Inflorescence com-

posed of spiralled branch-bracts, more cone-shaped. ••

Eplscopales

Series Imbrlcatae Griggs

Imbrlcatae Griggs, Bull. Torr. Bot. Club. kZt 316.

1915* (Rank not specified; here Interpreted as

series.)

Inflorescence large, slightly pubescent, erect or

pendulous, 12-60 cm. long. Branch-bracts extending out

horizontally; middle branch-bracts 7.0-10.0 cm. long,

3«0-U.5 8lde-wldthi moderately deep boat-shaped* -

Type species I Hell con la Imbrlcata

0. Ktze.

Three entitles occur In this series, two of which are found In Central America. The third species Is found throughout Central America and the northern portion of

South America*

It Is speculated that this series Is representat Ire of the most primitive type of Inflorescence within the whole genus.

Key to the Species of Series Imbrlcatae

A* Inflorescence pendent; 20-70 branch-bracts per

Inflorescence. • • 1. H. marlae

A* Inflorescence erects 10-20 branch-bracts per

Inflorescence.

B. Pleoit about 1^ m. tall: middle branch-bracts

8-10 cm. long; undersurface of leaves maroon

euid with reticulate veins*. 2. H,. reticulata

B* Plant 3-5 m* tall; middle branch-bracts 6-7 cm*

long; undersurface of leaves green and with

parallel veins, not reticulated

3« H* Imbrlcata 1

1. HELICONIA MARIAE Hooker f.. Jour. Linn. Soc. Bot.

7t 68-69. 1864. Blhala marlae (Hooker f.) 0. Ktze..

Hey. Gen. PI. 2t 66k, I89I. Typet New Granada

(). Bolivar, Betaml on Slnu River* Dr. 1. Antholne

(London, Kew Herb., not seen).

Hellconla elegans Petersen In Mart. PI. Bras. 33

12. I89O. Blhal elegans (Peters.) 0. Ktze.,

Rev. Gen. PI. 2t 684. I89I. Type: Panama,

Paralso, Dec. 1897. Wagner I9OI8 (M).

Hellconla pxmlcea (Griggs) L. B. Smith, Contr. Gray

Herb. 124: 6. 1939. Blhal pxmlcea Griggs, Bull.

Torr. Bot. Club 42 1 321. 1915 • Type: Panama,

between Gorgona and Gatun, alt. 10-50 m., 7 Jan.

1911, H. Plttler 2290 (US).

Large, stout plant, up to 12 m., sometimes more; musold habit. Leaf-blade obiong-elllpt leal, 2-3 n. long,

30-60 cm. wide, acute apex, rounded to obllquely-sub

-cordate base; upper and lower surface green; petiole

prominent, at least 20 cm. long, stout. Inflorescence

pendulous, puberulent-tomentose, I5-6O cm. long (varies

greatly In length), peduncle tomentose, 1 cm. diameter;

rachls puberulent-tomentose, 0.6-0.8 cm., hidden by

imbricated branch-bracts. Bremch-bracts 20-68, deep red*

puberulent, wide-ovate deep boat-shaped; lowest branch

-bracts ovate-acuminate, 12-13 cm. long; lower )

1^8

branch-bracts 6-8 cm. long; middle and upper branch-bracts

'f-5 cm* long. All brem oh- bracts 3-'*^ •5 cm. side-width t all branch-bracts completely overlapping. Ploral-bracts cm. long (slightly shorter than flowers), 1.5-2.0 cm. wide, glabrous. Flowers 10-20 per branch-bract, red to white, exposed portions are red; 'f-5 cm. long; perianth 3-3 cm. long, calyx vlllous-tomentose on outer surface, corolla glabrous; pedicel 1.2-1.8 cm. long glabrous. Fruit blue, subglobose, about 1 cm. diameter. (Plate 1.)

Wet forests, steep wet slopes, edge of bogs, sea level to about 700 m. altitude. British to Panama.

This species also extends to and throughout northern South

America. (Map 1.

There is no taxonomlo confusion with H. marlae. In the past, names which are now synonyms were given to this species for the slightly different color of red or differ- ent length of the Inflorescence. This species was origi- nally named H. marlae by Hooker in honor of the Empress of Russia upon the request of its discoverer. Dr. Antholne.

H. marlae is the only pendent imbricated species.

Therefore, it should not be confused with other species of Hellconla . It is commonly known as the "beef-steak" plant because the closeness of the branch-bracts give the appearance of a "solid red piece of meat hanging from a tree." 49

PLATE 1* HELICONIA MARIAE Hook.

Isotype: Blhal punlcea Griggs, between Gorgona and Gatun. Canal Zone, ? Jan. I9II, h. Pittler 2290 (US).

J t

50

This species with Its Interesting Inflorescence has been used extensively In cultivation.

Representative specimens

BRITISH HONDURAS. District Stann Creek . W. A. Schlpp

8-

(F).

COSTA RICA. Prov. Alajuela . A. Molina R., L. 0. Williams.

W. C. Burger and B. Wallenta 17657. 21 Feb. 1966, edge of bog, wet tropical rain forest near Los Angeles, alt. 100 m.

(F). Prov. Cartago . W. Burger 3875. 1^ Dec. 1966, steep slopes above the Rio Reventeizon and below the Interamerlcan

Institute of Agricultural Sciences near Turrlalba, + 700 m. alt. (F). Prov. Llmon . F. Soils 431, 1936, Toro Amarlllo

(P).

GUATEMALA. Dept. Alta Verapaz . 0. F. Cook and R. F.

Griggs 787. 2 May 1902, road from Sepaculte to Panzos

(US). Dept. Izabal . J. A. Steyermark 39638, 18 April

19'^0, Jungle bordering Quebrada Roseul, lower slopes bordering eastern part of Cerro Santa Cruz, n.e. of San

Felipe, alt. 5O-I5O m. (F).

HONDURAS. Dept. Atlantlda . A. M. Chlckerlng 90, 22 June

- 27 July 1929. near Toloa (MICH). P. C. Standley 53^7^5*

Ik Dec. 1927 - 15 Mar. 1928, wooded swamp, vicinity of

Tela, (F, US). T. 0. Yuncker, J. M. Koepper, and K. A.

Wagner 8529. 19 July 1938, on slopes near Puente Alto stop Map 1. Distribution of Heliconia mariae * 52 on the S. P. Co. R. R., e. of Ceiba« alt. 900 ft. (?, GH,

NY).

PANAMA. Prov. Bocas Del Toro . H. von Wedel 2905. 23 Oct.

19'^i* Bastlmentos Island, Tlclnity of Chiriqul Lagoon (MO).

Canal Zone . W. R. Maxon 678^. 6 June 1923. old site of

Gorgona (US). C. H. Ostenfeld 35» 21 Jan. 1922, In forest,

Gatun (US). C. V. Piper 5^37* 23 Feb. 1923. Empire to

Mandlnga (US). C. E. Smith, Jr., H. N. Smith and T. Arrauz

3292, 26 Dec. 1957. off road K-15 near Hulle on the hills

aroxrnd Gatun Lake; secondary and out-over primary forest

(P). P. C. Standley 26OOO, 3 Dec. 1923 t Cerro Gordo, near

Culebra (US).

2. HELICONIA RETICULATA (Griggs) Winkler In Engl,

and Prantl. Nat. Pflanzenfam. 15ai 536. 1930. Blhal

reticulata Griggs, Bull. Torr. Bot. Club 31 1 kkS, Types

Costa Rica, Buena Vista, road to San Carlos Valley, alt.

600 m., 0. P. Cook and C. B. Doyle 1^9, April I903 (US).

Short plant, approximately 1-1^ m. tall; musold habit.

Leaf-blades elliptical, thin and glabrous, 70-102 cm. long,

30-40 cm. wide, acute apex, cuneate base; upper surface green, lower surface maroon with reddish, parallel veins

connected by cross velnlets giving appearance of reticula- tion; petiole prominent, about 70 cm. long. Inflorescence

erect, 12-15 cm. long, peduncle short to almost lacking. ... . j

puberulent; raohls straight, 0.5 cm* diameter* Branch )

-bracts 10-15, red, smooth, ovate with lowest branch-bracts ! with long acute apex and others with more or less rounded to short acuminate apext deep boat-shaped, branch-bracts pointed upward and Imbricated so that rachls Is hlddeni lowest branch-bract 13-16 cm* long, 3*0 cm. side-width, sterile; middle and upper branch-bracts 7-10 cm. long,

3.0 cm. side-width, fertile. Floral-bracts few, whitish,

4.0 cm. long, 0.5-0.8 cm. wide. Flowers 5-6 per branch

-bract, red and yellow, 3»5'-$»0 cm. long, perianth 2.5 cm. long, glabrous; pedicel 0*9-1 .8 cm* long, glabrous*

Fruit not seen. (Plate 2.)

Deep ravines, moist forest floor, and thickets near rivers. Endemic to Costa Rica. (Map 2.)

The only species with which H. reticulata might be confused Is H. Imbrl cata. Both have erect Inflorescences '4 with red, overlapping branch-bracts. The former Is a much shorter species with a maroon leaf-blade undersurface on which there Is reticulation of the veins. The latter Is a tall species with a green leaf-blade undersurface.

H* reticulata has not been collected very extensively*

It Is a species which, with the red Inflorescence and the unique undersurface of the leaf-blade^ may be of Interest td the horticulturist* 55

PIATE 2. HELICONIA RETICUIATA (Griggs) Wlnkl.

Holotype: H. reticulata (Griggs) Winkl., Buena Vista, road to San Carlos Valley, Costa Rica, April I903, 0. F. Cook and G. N, Collins 149 (US). Map 2» Distribution of Hellconla reticulata.

t

58

Representative specimens

COSTA RICA. ProY. Alajuela * A. Molina R.. L. 0. Williams,

W. C. Burger and B. Wallenta 17^k6, 18 Feb. I966. forest floor along Rio San Rafael, near hot springs. Hacienda La

Marina, ca. 1^ km. n.e* of Villa Quesada, alt. 500 m. (F).

A. Molina R., L. 0. Williams, W. C. Burger and B. Wallenta

17312, 17 Feb. 1966, thickets and wet forests near Rio San

Bafail, 2 km. w. of La Marina, Llanura de San Carlos, alt*

550 m. (F). Prov. Limon . A. Jimenez 2811, 9 Feb. I965, bosque oerca al Rio Toro Amarillo, oerca al puenta de ese mismo riT, unoe 5 km., al SO. de Guapiles, jilt. 300-ifOO a.

(F).

3* HELICONIA IMBRICATA (0. Kuntze) Baker, Ann. Bot.

7t 191. 1893. Bihai imbricata 0. Ktze., Rev. Gen. PI.

2 I 684. I89I. Typex Costa Rica: Port Limon (NY).

Heliconia densa (Griggs) L. B. Smith, Contr. Gray

Herb. 12^*1 6. 1939. Bihai densa Griggs, Bull,

Torr. Bot. Club kZt 321. 1915 • Typet Panama,

Prov. Colon, around Dos Booas, Rio Fato valley,

alt. ifO-80 m. 16 Aug. I9II. H. Pittier s.n. (US).

Stout plant, 2^-5 m. tall; musoid habit. Leaf-blades oblong-ovate, ^^0-130 cm. long, 25-36 cm. wide, acuminate apex, obtuse base; upper and lower surfaces green t petiole prominent, about 75 cm. long* stout smd usually puberulent

-velutlnous. Inflorescence erect, pubescent, 20-30 cm* long; peduncled 1.5 cm. thick; rachls short tomentose

(golden brown), hidden by bracts, 1.3 cm. diameter.

Branch-bracts 10-20, usually 10-lU, red to orange, greenish at base, slightly tomentose, short ovate, subcordate, deep boat-shaped; lowest branch-bract sterile, 11 cm. long, lower fertile bremch-bracts 6-7 cm. long, about 3 "5 cm. side-width; middle said upper branch-bracts nearly same dimensions as lower branch-bracts, 5-6 cm. long, 3*5 cm. side-width; branch-bracts Imbricated so rachls not exposed.

Floral-bracts elllptlc-ovate, k cm. long, 1 cm. wide.

Flowers per branch-bract, green to yellow, 3»5-^«0 cm. long: perianth 2.0-2.5 cm. long, calyx villous, corolla glabrous; pedicel 1.0-1.5 cm. long, glabrous. Fruit blue, subglobose, 0.7-0.8 cm. diameter. (Plate 3*)

In moist forested areas and dry river banks. The known range of this species Is Panama and Costa Rica, but

It may also extend Into South America. (Map 3*)

There Is no taxonomlc confusion with this species.

It Is closely related to the erect H. reticulata and the pendulous H. marlae. The lower surface of the leaves on

H. reticulata are maroon with netted veins. 60

1

PIATE 3. HELICONIA IMBRICAIA (0. KTZE) EAKEH

Holotypej BlhXa densa Griggs, around Dos Bocas. Rio Pato Valley. Prov. Colon. Panama. 16 Aug. I9II. H. Pittier, s. n. (US). t

Representative specimens

COSTA RICA. Prov. Cartago . W. R. Hatch 195. 22 July 1937.

Heventazon valley "Las Animas" (P). Prov. Heredla * P. S.

Rajas ^^31. Aug. 1936, Rio Toro Amarlllo (flnca Vleja) Reg.

Atlant. alt. about 200 m. (MO). Prov. Llmon . C. W. Dodge and V. P. Goerger 9'^18, 27 July 1936 (MO), G. Kuntze 1953.

14 June 187^*, Port Llmon (P, NY).

PANAMA. Prov. Bocaa Del Toro . H. Wedel 400, 9 Aug. I9UO

(MO). Prov. Chlrlqui . P. H. Allen 5021, 1 Aug. 19^7. frequent In forest, Cerro Galera Chorcha, vicinity of

Gualaca, alt. 1000 ft. (US). M. E. Davidson 1177. 18 Aug.

1938, stream banks, Puerto Armuelles, alt. 50 ft. (P).

R. E. Woodson. Jr. and R. W. Shery 862, 28-31 July 19^0, vicinity of Puerto Armuelleo, alt. 0-75 m. (GH, US).

Prov. Darien. J. A. Duke 5299. H Aug. I962, Cerro Plrrei ascent of Cerro Plrre from Rio Pirre, s. of El Real, 100

-600 m. (MO). J. A. Duke 5'^^2. 10-11 Aug. I962, about

10 ml. s. of El Real on Rio Plrre (House no 22) (MO).

Series Eplscopales Griggs

Episcopales Griggs, Bull. Torr. Hot. Club 42i 316.

1915 • (Rank not specif ledi here Interpreted as series.)

Inflorescence erect, up to 10 cm. long. Branch-bracts ascending, deciduous, 6-7 cm. long, ovate, rather shallow boat- shaped. 1

1

I

I Map 3» Distribution of Helloonla Imbrlcata.

i

Type species I Hellconla

epiacopalis Veil.

(Flora Plum. IIIi 22. 1827.)

This Is a monotyplc series In which the entity Is

South American. The species Is approximately two meters tall with an erect Inflorescence on a long, glabrous

peduncle. It Is quite different from the species In « the Series Imbrlcatae. H. eplscopalls possesses deciduous,

OTate, shallow boat-shaped bracts, which are ascending on the Inflorescence.

Since this Is a South American species, H. eplscopalls will not be treated further in this work.

II. Section HBLICONIA

Subgenus Platychlamys Baker In part. Ann. Bot. Vol. VII

190. 1893.

Slender or stout plants, 2-5 m. tall; musold habit.

Leaf-blades usually 1 m. or more In length; petloled.

Inflorescence sessile or subsesslle, occasionally peduncled; glabrous; rachls may be hidden or partially hidden by subcordate branch-bracts. Branch-bracts 7-15, large and thick; middle branch-bracts 8-13 cm. long,

2.5-6 .0 cm. side-width.

Type species t Hellconla

blhai (L.) L. Pour very closely related species occur in this sec-

tion. All have large, erect, and colorful inflorescences,

most of which are sessile. The branch-bracts are thick,

somewhat coriaceous, and fleshy. Upon drying, the branch

-bracts show considerable shrinkage.

Key to the Species of Section HELICONIA

A* Branch-bracts solid color, either deep crimson, yellow

or green, not variegated; subulate tips of branch

-bracts recurved? flowers usually number 10-20 per

branch-bract H. caribaea

A. Branch-bracts consisting of variegated colors, tips

of branch-bracts not recurved; flowers number less than 10 per brsmoh-bract.

B. Branch-bracts very deep boat-shaped, broadly

ovate, tapering to blunt, incurved tip; branch

-bract orange with red splotches, usually with

red margin; perianth 4.5-5.0 cm. long

5» H. bourgaeana

B. Branch-bracts moderately deep boat-shaped, ovate

to acuminate tip; branch-bracts with green or

yellow margin; perianth either 4.8-6.5 cm. long or 3,5-4.5 cm. long.

C. Branch-bracts with green margin; perianth

4.8-6.5 cm. long 6. H. wagneriana

C. Branch-bracts with yellow mairgin; perianth

3*5-4.5 cm. long.... 7. H. bihai 66

k» HELICONU CARIEAEA Lamarck, Encyol. Meth. Bot.

1* t ^26, 1783* Typet Antilles. This type Is based on the solidly red-braoted Tar* B* figured by Plumler*

Not. pi. Am. Gen. 50. 1703* and PI. Am. ed. Burm. k9

Pl« 59. 1755.

Musa blhal Tar. B. L.. Sp. PI. 10^3* Typet based

on seune type as Hellconla carlbaea .

Hellconla blhal L. sensu Jacq., Hort. Schoen. 1. Zk*

1797* Typet based on same type as Hellconla

carlbaea .

Hellconla blhal L. sensu L. C. Rich. In Not. Act. XV.

suppl. 22 pi. 10. I83I. Typet based on same

type as Hellconla carlbaea .

Hellconla conferta 0. G. Petersen, PI. Bras. Tol. 3,

pt. 3t 1^. I89O. Blhal conferta (Peters.) 0.

Ktze., Rot. Gen. PI. 2t 685. I89I. Type: Island

Guadeloupe (Herb. Hus. Munich, tracing of type seen

(P)).

Hellconla blhal L. sensu Griggs, Bull. Torr. Bot. Club

3O1 657. 1903. Blhal blhal (L.) Griggs sensu

Griggs, Bull. Torr. Bot. Club 31 1 445. 190^^.

Typet based on same type as Hellconla carlbaea. 67

Helloonla borlnguena Griggs, Bull. Torr. Bot. Club

30x 657. 1903. Blhal borlnguena (Griggs) Griggs,

Bull. Torr. Bot. Club 31 » ^^^^5. 190»f. Types

Porto Blco, 13 km. m. of Cayey, 24 Jiine 1901«

Underwood and Griggs 363 (US).

large, slender plants m.; musold habit. Leaf

-blades long oblong 1-li m. long, 26-56 cm. wide, short acuminate apex, obtuse-acute baset upper surface green, lower surface either green or glaucous; petiole stout, glabrous, 1 m. or longer. Inflorescence erect, glabrous,

30-4^0 cm. long; sessile or nearly so; rachls often hidden by orerlapplng of base of bremch-bracts. Branch-bracts

6-15, usually more than 10, solid color (not variegated)

deep crimson, yellow or green, glabrous, broadly ovate and fleshy, subcordate around rachls, deep boat-shaped;

lowest branch-bract often sterile and expanded Into leaf

-blade, total length 25-70 cm.; lower fertile branch-bracts

12-18 cm. long with long subulate tips, recurved; 3-4 cm.

side-width; upper branch-bracts 8-10 cm. long, with more

obtuse apices 2.5-3*0 cm. side-width; bases of branch

-bracts often overlap, appearing to be partially Imbricated.

Floral-bracts wide-lanceolate, glabrous 4.5-5.5 cm. long,

O.5-I.5 cm. wide. Flowers 10-20, 4.0-6.5 cm. long; perianth white with greenish apex, 3-5 cm. long, glabrous;

free sepal ca* O.5 cm. wide, straight; pedicel 0.7-1 ,0 om.

1 long, glabrous. Fruit dark; subglobose, 3-sided, about

1 cm. long, 0.8 cm. wide when dried. Seed not seen.

(Plate k,)

In deep ravines, moist forests and slopes of the

Caribbean islands. Cultivated throughout tropics but seemingly naturally endemic to the Greater and Lesser

Antilles. (Map k,)

This is a species with which there has been much taxonomic confusion. This confusion stems from the descriptions of Bihai by Plumier (1703). The first taxon he described was "...vasculis coccineis," followed by

. .vasculis subnigris," and "...rasculis yarlegatis."

Linnaeus (1753) accepted Plumier' s three taxa as varieties, but altered the listing by placing "...vasculis variegatis" first and treating it as the "typical" variant of his Musa

Bihai . This emphasis was not changed in 1771 when he published Heliconia as a distinct genus, based upon his

Musa Bihai . Later workers, desiring the original descrip- tion, went to the earliest publication, that of Plumier

(I703). and gave the name H. bihai to "...vasculis coccineis" and other names to "•• .vasculis variegatis."

Following the rules set down by the International Code of

Botanical Nomenclature (Article 11), Linnaeus' first entity becomes H, bihai . and the earliest name for Plumier' s first entity is H. caribaea . Plumier 's "•• .vasculis subnigrls" appears to be an overly mature specimen of H. bihai. 69

PLATE 4. HELICONIA GARIBAEA Lam.

H. carlbaea Lam,, Sjmdlcate Plantations, Dominica, West Indies, 8 April 19^10, W. H. Hodge 2632 (GH). t

70

As stated previously, H. carlbaea Is closely related to other Heliconla species which possess large fleshy branch-bracts. These are H» blhal, H. bourgaeana said H. wagnerlana * H. carlbaea differs from H. blhal and H. wagnerlana In that the former has solid colored branch

-bracts, and the latter two species are variegated. It

differs from H. bourgaeana In that the tips of the upper

branch-bracts are more acuminate, and the calyces of the

flowers are glabrous. In H. bourgaeana the tips of the

upper branch-bracts are acute to blunt and, the calyces

of flowers are pubescent.

Representative specimens

WEST INDIES. ANTIGUA. H. E. Box 826, 30 May 1937. n.w.

side of Boggy Peak at ca* 900 ft*, mesophytlc woodlands

(PIAS. US).

CUBA. Prov. Orlente . E. L. Ekman 160'<'2, 19 Dec. 1922.

Bio Caoba, In narrow gulch of Rio Caoba below Brune (NY).

Bro. Leon 9985* July 1921, In moist woods, 900 m. alt.

Loma del Gato. Sra. Naestra (GH). J. A. Shafer 8029* 21

Dec. 1910, Arroyo Henequen, foot of El Yunque (NY). C*

Wright 607. I856-7. In Cuba Orlentall (GH).

DOMINICA. W* H* Hodge JlkZ, 10-23 Aug. 1938, rainforest

on the precipitous slopes of Mome Colla Anglais, alt*

610-732 m*. Sylvanla (NY)* W* H. Hodge 263I. 8 April

19'fO, deep rich soil, border of high-stem rainforest. Syndicate Plfiintations, close to n.w. slopes of Nome

Dlablotln, alt. 600 m,, rainfall 500 cm. (GH). W. H. and B. T. Hodge 3'*16. 6-7 May 19^*0, moist forest bordering

Pegona Hlver In vicinity of Deux Branches, Concorde Valley

(GH). W. H. and B. T. Hodge 3658, lO-l^f May 19lf0. moist

forest In ralley of Hsunpstead River, ca. 2 ml. from mouth, alt. ca. 100 m.. La Chandlere (GH). R. L. Wilbur. B. L.

Dunn, H. A. Hespenhelde and D. R. Wiseman 7801, 22 July

196^. Lower montane rainforest on the lower slopes of Trala

Plton along the road to Rosalie about 2.5 ml. from Pont

Casse at elev. of ca. I6OO ft. (DUKE). R. L. Wilbur, E.

L. Dunn, H. A. Hespenhelde, D. R. Wiseman 83^7, 6 Aug.

1964, trail east from Fresh Water Lake (L'etang) towards

Rosalie on the slopes of Mome Macaque (Mlcotrln), alt. ca. 2100 ft. (DUKE).

DOMINICAN REPUBLIC. Prov. La Vega . H. A. Allard 17773.

1^ Dec. 19'^7t In deep ravine, vicinity of Pledra Blanca, alt. 200-500 m. (US). Prov. Lamana . E. L. Ekman 152^5,

7 June 1930. Peninsula de Samana. El Valle (US). Prov.

Santiago . J. Jimenez 370^^, Sept. I558, Santiago, Mr.

Arlza-Julla«s Garden (US).

GUADELOUPE. P. Duss 3838, Sept. 1895, Bams-Lannes,

Houelmont (NY, US).

HAITI. Dept. Nord or L'artlbonlte . G. V. Nash 674. 23 Aug. 1903, on wooded bank, Plalsanoe to Marmalade, about ft. 2500 (NY). Dept unknown . E. L* Ekman H4391, 25 June 1925* Mome Cheseman, oa. ^00 m. (US)* G. V* Nash and N*

Taylor 1139* 22,23 July 1905, In deep gully, alt. 1800 ft,.

Camp No. 2, Mt. Haleuvre (NY)«

JAMAICA. Portland Parrlsh * R* A* Howard, G. B* Proctor, and W. T. Steam 14805. 2k Jan. 1956* The John Crow Mts.,

1.5-2*5 ml* s.w. of Ecclesdown. Alt. 1500-2500 ft. (GH).

A. Skutch s.n., 12 July 1926, Abbey Green, Blue Mts. (P).

West, Arnold and Laessle 721, 18 July 1952, roadside, 12 ml. e. and mi, s. of Port Antonio, Ecclesdown Road

(PIAS, GH). T. G. Yuncker 18837, 4 June 1958. in deep

forest on Joe Hill above Moore Town, John Crow Mts., alt*

1000-2000 ft. (MICH). St. Thomas Parrlsh . W. R. Maxon

9097, 6-12 June 1926, deep ravine in mountain forest above

House Hill, St. Thomas, alt. 500-700 m. (US). E. West and

L. Arnold I70, I5 June 1951* low montane forest between

Blue Mts. and John Crow Mts. (FIAS). T. G. Yuncker 17528,

21 Nov. 1957. on moist slope, ravine along Sxilfur River, above Fountain Inn. alt. 5OO-75O ft. (P, MICH).

MONTSERRAT. J. A. Shafer 570, 16 Peb. I907, Woodlands

Mountain, 1800-2000 ft. (NY).

PUERTO RICO. Dept. Htunacad . H. L. Blomqulst 13154, 19

April 1942, below Big Tree Trail, off road to El Yunque

Mt. (DUKE). N. L. Britton and J. A. Shafer 1684, 2-6 Mar. 1913* hillside forest, Rio Arribe, Pajardo and vicinity

(NY. US). Mr. and Mrs. A. A. Heller 1045. 12 April I899,

in damp ravine. La Perla Estate Luqulllo Mts. (NY). ' 73

p. H. Sargent 682, 1 June 1935* moist wooded ravine,

Barranqultas (US). J. A. Shafer 3352, 21 July 191^*. deep shade In ravine, summit of Loma La Mlna to Rio

Blanco, alt. 9^*5-360 m. (US). W. R. Stlmson 3518, 18

July 1966, El Yxmque National Forest, In rainforest beside

stream above cement bridge beyond second rain shelter from lower exit of Victory Trail (DUKE). Dept. San Juan . P.

Slntennls 1131, 24 Mar. I885, Prope Bayamon In sylvls montanlx (GH). P. Slntenls I588, II 1885, Sierra de

Gugulllo In sylvls In reglone media montes Jlmenes (P).

ST. VINCENT. H. A. Howard 11162, 1-7 April 1950, River

Valley, Silver Spoon district, above Three Rivers (GH).

H. A. Howard 11164, 1-7 April 1950, River Valley, Silver

Spoon district, above Three Rivers (GH). C. V. Morton

5292, 23 April 1947, mountains above Chateaubelalr River,

alt. 400-750 m. (US). J. A. Shafer 570, I5 Peb. I907.

Woodlands mountain 1800-2000 ft. (NY).

TORTOIA. W. C. Plshlook 467, 1919. Sage Mountain (NY, US).

5. HELICONIA BOURGAEANA 0. G. Petersen, PI. Eras.

Ill, pt. 3i 3« 1890. Blhal bourgaeana (Peters.) 0.

Ktze., Rev. Gen. PI. 2i 685. 1891. Type: Habitat in

Mexico ad Tuspango prope Cordoba (State Veracruz), Jun.

flort Bourgeau n. 2502 (not seen). ;

Map 4* Distribution of Helloonla carlbaea* 75 76

Helloonla champnelana Griggs, Bull. Torr. Bot. Club

3O1 657. 1903. Blhal champnelana (Griggs) Griggs,

Bull. Torr. Bot. Club 31 1 kl^5, 190k. Typei

Guatemala. Prov. Alta Verapaz, 8 April 1902, 0. P.

Cook and R. P. Griggs, 528 (US).

Hellconla barqueta Loes. In Verh. Bot. Ver. Branden-

burg 5I1 18. 1909. Blhal barqueta (Loes.)

Griggs, Bull. Torr. Bot. Club k2t 324, I9I5.

Typei Mlxlco, State Veracruz, Jalapa, Coatepec,

Ik Dec. 1902. E. Seler 36O9 (B).

Stout plant about 2-5 m. tall; musold habit. Leaf

-blades long ovate-elllptlcal, 3A-2 m. long, IO-5O cm. wide, cuspidate-acuminate apex, obtuse-acute base; upper surface green, lower surface pale-green, occasionally pubescent along midrib; petiole long and moderately stout, sometimes hirsute. Inflorescence erect, glabrous, about

45 cm. long; sessile or nearly so; rachls red, moderately to slightly flexuose, 1.0-1. 5 cm. diameter. Branch-bracts

5-11, usually bright orange with large red spots or bright red, yellowing along base and deep red along margin toward apex, glabrous, wide-ovate, subcordate around rachls; lowest branch-bract sometimes sterile, usually extending

Into leaf-blade, total length 3O-9O cm.; lower fertile branch-bracts with acuminate apices, gradating to obtuse apices toward top of infloresoenoe, all rery deep 77

boat-shaped; lower branch-bracts 13-16 cm. long, 3.5-6.0

CD. slde-widthj middle an& upper branch-bracts 8-13 cm. long. 3«0-5.0 cm. side-width, decreasing in size toward apex; lower branch-bracts close together, but not imbri- cated, intemode 2.0-3.0 cm. long, rachis often hidden.

Floral-bracts lemceolate, cm. long, 0.5 cm. wide, glabrous. Flowers 8-10, green on outside, whitish inside, glabrous, 6. 0-7. 5 cm. long; perianth if. 5-5.0 cm. long, calyx villose along margin and at apex, pedicel 0.7-1.0 cm. long, glabrous. Fruit blue, subglobose, 1 cm. diameter dried; not exerted from branch-bracts. Seed 1.0-1.2 cm. long, 0.6 cm. wide, 2 sides flattened, third side rounded; seed coat sclerified and rough. (Plate 5.)

Along edges of wet lowland forests, low moist places near rivers and swampy areas. Range extends from Mexico to Honduras. (Map 5.)

Taionomically, this species has been confused in a similar manner as H. bihai . In several floristic studies, it has been placed with or has been called H. bihai .

However, many past workers (Petersen, 1890; Griggs. 1903;

Loesener, 1909) have recognized this taxon as a species in its own right.

H. bourgaeana differs considerably from H. bihai in having deeper boat-shaped bracts and longer, pubescent perianths. It also has a different geographical distribu- tion. The only other deep, boat-shaped species with which 78

m

JL

*

JL »

N « MCW VOKtl aoTAHiCAL OA! «

*^

f: PLATE 5. HELICONIA BOURGAEANA 0. G. Peters.

H. bourgaeana 0. G. Peters.. Stann Creek Valley, British Honduras. 1? May 1932. W. A. Schlpp 979 (NY). i

H. bourgaeana might be confused Is H. wagnerlana * In the

former, the flowers are never exerted, they are hidden within the branch-bract . The latter has flowers that are exerted from variegated, green-margined branch-bracts.

Representative specimens

BRITISH HONDURAS. Dlst. Stann Creek . P. H. Gentle 2117,

9 Aug. 1937. Stann Creek Hallway, 15 ml., base of hill

(MICH, NY). W. A. Schlpp 979. 17 May 1932. 22 mile Stann

Creek valley, 250 ft. alt. (P, GH, MICH, MO. NY). Dlst. unknown . H. P. Smart s.n. 1929. Silk Grass Reserve (NY. US).

GUATEMALA. Dept. Alta Verapaz. 0. P. Cook and R. P.

Griggs 528, 8 April 1902, near the Plnca Sepaculte (US).

J. A. Steyermark t^k^kl, 28 Peb. 19^2, lowland forest In valley, "pantano. " 2i miles w. of Cubllqultz. alt. 250-300 m. (P). H. von Tuerokhelm 83I6, M. April 1902. Cubllqultz, alt. 35 m. (US). H. von Tuerckhelm II 586, Peb. I913,

Cubllqultz (US). Dept. Huehuetenango . J. A. Steyermark

^9351. 23 Jiily 19^2. wooded slopes bordering Rio Lacandon. between Ixcan and Rio Ixcan, Sierra de los Cuchumatanes. alt. 150-200 m. (P). Dept. Izabal . P. C. Standley 73001, May wet forest, 3 1939. Escoba, across the bay (w. ) from

Puerto Barrios, near sea level (P). J. A. Steyermark

38702, 4 April 19^0, between Virginia and Lago Izabal,

Montana del Mlco, alt. 5O-IOO m. (P). J. A. Steyermark

39751. 21 April 19'fO. swampy Jungle In valley of tributary 80

of Rio San Francisco del Mar, 2 ml. n.e* of Hopl, 12 ml* e. of Entre Rlos, alt. 10-20 m. (P). Dept. Pet en . J. A*

Stejrermark 45391, 25 Mar. 19*^2, forest between Plnca

Yelpemech along Rio San Diego and San Diego on Rio Cancuen, alt. 50-150 m. (P, MICH, MO, NY, US). J. A. Steyermark

45956, 25 April 1942, along Rio Machaqullla. n. of El

Camblo, alt. 75-100 m. (MICH).

HONDURAS. Dept. Atlantlda . P. C. Standley 52721, 6 Deo.

1927 - 20 Mar. 1928, Lancetllla Valley near Tela, alt.

20-600 m. (P. US). P. Wilson 331, k Peb. I903, forest

near Elcotts' Plantation, Pt, Sierra (NY). T. G. Yuncker

4927 a and b. 5 Aug. 1934, In low place near the river, vicinity of Lancetllla (MICH, MO). T. G. Yuncker, J.

Koepper and K. A. Wagner 835 0, 10 July 1938. near bank

of Salado river, vicinity of village Salado, vicinity of

La Ceiba (P, GH, MICH, MO, NY, US). Dept. Prancisco

Morozan . A. Molina H. 7 Mar. I962, El Zamorano, alt.

800 m. (US).

MEXICO. State . E. Matuda 3683, 10*.12 July 1939,

in forest, Revlna, Palenque (P, GH). K. Roe, E. Roe. 1379. 23 Aug. 1965, cut-over tropical evergreen forest. Interior lowlands, km. 33 (by road) s. of Surest e on road to Mai

Pasa, near border, alt. ca. 35O m. (WIS), State

Oaxaca. H. E. Moore, Jr. 8110, 10 Oct. 1959, Ravine 35 km. e. of Mathlas Romero. 4 km. w. of Palomares on trans

-Isthmus highway (GH). B. P. Heko 4093. Mar. I919. i

I

Map 5. Distribution of Hellconla bourgaeana .

)

83

Teplnapa, alt. 250 m. (US). State Vera C?ruz . E. W. Nelson

136, 22 Feb. 189'^, near Motzorongo (US). C. A. Purpus

15760, April, May 1931. Zacuapan (MICH). C. A. Purpus

I58II, Aug. 1931* along streams, Zacuapan (MICH).

6. HELICONU WAGNERIANA 0. G. Petersen In Mart. Fl.

Bras. 3 pt. 3» 12. I89O. Blhal wagneriana (Peters.) 0.

Ktze., ReT. Gen. PI. 2t 685. I89I. Typet Panama, prope

Haume et Gorgone, m. Jan. flor. H. Wagner (M).

Hellconla elongata Griggs, Bull. Torr. Bot. Club 30»

653. 1903. Blhal elongata (Griggs) Griggs, Bull,

Torr. Bot. Club 31t kk5. 190**. Type: Guatemala,

Alta Verapaz, 2 May 1902, 0. F. Cook and R. P.

Griggs 790 (US).

Hellconla aurea G. Rodriguez, Bol. Soc. Venez. Clenc.

Nat. 15 » 120. 195^* Type I Aragua, G. Rodriguez

230, Dlclembre de 1953* (Herb. Nac. de Venez., not seen.

Stout plant Z-k m. tall; musold habit. Leaf-blade oblong, about 60 cm. long, 36 cm. wide, short actimlnate apex, long cuneate to rounded base, upper and lower sur- faces greenj long petloled. Inflorescence erect, glabrous,

30-40 cm. long; short pedxinole or sessile; rachls stiff and nearly straight, 0.8 cm. diameter. Branch-bracts 7-14, bright red shading to yellow with green border, glabrous. ek j

all ovate with acuminate aploes; lower branch-bracts long

acuminate, upper branch-bracts shorter acuminate; all deep

boat-shaped; lowest branch-bracts often extends Into leaf

-blade with blade up to k^,^ cm. long, 10 cm. wide. Second

lowest branch-bract 16-18 cm. long: middle and upper branch

-bracts 10-l^f cm. long; all branch-bracts 3.0-4.0 cm. side

-width; Intemode between branch-bracts 1.3-3.0 cm.

PI oral-bracts few, triangular ovate, if-5 cm. long, 1-2

cm. wide, glabrous. Flowers if-S per branch-bract; green, bright green at exposed tips, 6-8 cm. long, projecting

to and above margin of branch-bract; perianth if. 8-6. 5 cm. long, glabrous; pedicel 1.0 cm. long, glabrous. Prult blue, subglobose, 1.3 cm. long; 1.0 cm. diameter. (Plate 6.)

In moist to wet forests, swamps, and along river margins at altitudes up to 900 m., ranges from Honduras

Into northern South America. (Map 6.)

This species has often been confused with and called

H. blhal, since It Is variegated with deep boat-shaped branch-bracts. Those who have studied the H. blhal com- j plex readily distinguish this species by the red branch

-bracts bordered by yellow gradating to a green margin.

The perianth also differs In length. The other closely related species are H. bourgaeana and H. oarlbaea. 85

1507i

HI ft Hi ll 0» CAVAMA

m.V

BrartM rrt »nit y»;ij» witii frrm Mrrlr . tf r<>«l K U mraM llml* on thr ktlU wewJ Cmm L«k»i " PIATE 6. HELICONIA WAGNERIANA 0. G. Peters, H. wagnerlana 0. G. Peters, hills around Gatun Lake. Canal 26 Dec. 1957. C Earle Smith. Jr. et al. 3293 (F). t

86

RepresentatlTe specimens

BRITISH HONDURAS. Dist. Stann Creek . W. A. Schlpp 8-^H^^.

27 Dept. s.d*. in forest shade on hillside, Fainriew, alt.

300 ft. (P).

COSTA RICA. ProY. Alajuela . A. Molina H.. L. 0. Williams,

W. C. Burger and B. Wallenta 17619. 20 Feb. 1966, lowland

rain forest between Los Chiles - Venecia, Llanura de San

Carlos, alt. 200 m. (P). A. Jlminez M. 1767. 20 Feb. 1964,

Bordes del Rio Burio, alt. 200 m. (P). P. C. Standley

40172. 2 April 1924, moist forest, vicinity of Capulin,

on the Rio Grande de Tarooles, alt. 80 m. (US). Prov.

Llmon , P. C. Standley 36723. 6-7 Mar. 1924, wet thicket.

La Colombiana Parm of the United Pruit Co., alt. ca. 70

(US). Prov. Puntarenas . P. H. Allen 5752, 6 Jan. 1951,

Palmar Norte de Osa (GH). W. Burger and G. Matta U. 4654,

26 Jan. I967, at stream edge in deep shade of forest, aboTe

Rio Terraba near the Interamerican Hsy. about 10 km. ENE of Palmar Norte (to San Isidro), alt. + 40 m. (P). Prov.

San Jose . A. Molina R., W. C. Burger, and B. Wallenta

I8254, 2 Mar. I966, thickets along River Pejibaye, edge of Pejibaye Town, tropical rain forest and moist secondary forest. El General Valley, vicinity of San Isidro El General, alt. 340 m. (P), Prov. unknown . P. S. Rojas

200, 23 April 1935. Chitana (Meg Attant). alt, 600 (MO). GUATEMAIA. Dept. Alta Verapaz . P. C. Standley 70686, 5

April 1939* damp thloket, near Fanoajche, alt. ca« 360 h«

(F). P. C. Standley 92011^, 10 April 19^1. along Rio

Polochis, near Pancajche, alt. ca. 900 m. (P). J. A.

Steyermark 45227t 23 Mar. 19'<'2. woods s«e. of Plnca

Yalpemech, near Alta Verapaz - Peten boundary line, alt.

100-150 m. (P). H. Yon Tuerckhelm k052, April 1913. Wald

in Cubllquitz (US). Dept. Izabal . W. A. Kellerman 6I99,

20 Jan. 1905. Puerto Barrios (P). J. D. Smith I83O, M.

April I889. Monte Caohlrulo, alt. 700 ft. (US). P. C.

Standley 23992, 15-31 May 1922, in SKamp. vicinity of

Quiriqua. alt. 75 to 225 m. (GH. NY, US). P. C. Standley

73033. 3 May 1939. dense wet forest, Escoba, across the

bay (west) from Puerto Barrios, near sea level (P). J.

A. Steyermark 38756, k April 19^0, between Virginia and

Lago Izabal, Montana del Mico, alt. 5O-IOO m. (P). Dept.

Peten. J. A. Steyermark '+5956. 24 April 19^*2. along Rio

Machaquila, n. of El Cambio, alt. 75 m. (P. MICH. UC).

HONDURAS. Dept. Atlantida . P. C. Standley 55731. 7 Feb.

1928, wet flat forest. La Fragxia, alt. 20 m. (P, US).

P. C. Standley 767O, 22-27 April 19k7, dense wet forest, icinlty of San Alejo, at base of hills s. of San Alejo near Rio San AleJo, I5O-27O m. (P). Dept. Morazan . A.

Molina H. 102^f9. 7 Marzo I962, cultivada en mi Jardin de la SAP., El Zamorano, alt. 800 m. (F). . Dept. Comarca de El Cabo . A. Molina R. I5077,

22 Aug. 1965, Hatorrales del bosque lluvloso de San Mateo,

16 mlllas al sur de tronquera cerca de Rio Wawa, sabanas 7

plnares (Plnus Carlbaea) de la Costa Atlantlca, alt. 35 m. (P).

PANAMA. Canal Zone . K. E. Blum and E. Tyson 1979, 22 Dec.

1965. vicinity w. end of Gatun Lake dam (PSU). J. E.

Eblnger 184. 25 June i960, along edge of Trail 300 m.-600

m., Barro Colorado Island, Snyder Mollno Trail (MO). A. A.

Hunter and P. H. Allen 455, 9 Feb. I935t In Government

forest along Las Cruces Trail, 75 m. (P). H. Plttler 2291,

7 Jan. 1911. everywhere In clearing, railroad relocation

between Borgona and Gatun, alt. 10 to 50 m. (NY, US).

C. E. Smith, Jr.. H. M. Smith and T. Arrauz 3293, 26 Dec. 1957. off road K-I5 near Hulle on the hills around Gatim

Lake; secondary and cut-over primary forest (F). P. C.

Standley 28832. 28 Dec. 1923, moist thicket. Mount Hope

Cemetery (US). E. Tyson 3504. 22 Peb. I966, 6 ml. n.

Gamboa (PSU). Prov. Chlrlqul . R. E. Woodson Jr. and B.

W. Shery 892, 28 July - 1 Aug. 1940. vicinity of San

Bart dome, Peninsula de Burlca. alt. O-50 m. (MO, US).

Prov. Code. A. A. Hunter and P. H. Allen 455a, 17 Peb.

1935. mountains beyond La Plntada, 400-600 m. (MO, US).

Prov. Colon. P. H. Allen 4199. 3 Peb. 1947. Juan Mlna - Hlo Chagres - 25 m. (P). P. c. Standley 30347, 9 Jan.

1924, brushy slope, between Prance Field, and Catlval, Map 6. Distribution of Heliconia wagneriana . 90 91

Prov. of Colon (GH, US). Prov. Darlen . P. H. Allen ^^313. k Mar. 19^7, vicinity Plnogana, 20 m. (MO). A. Duke 5010,

^ July 1962, km. 16 de Yavlza along Q. Unital off Rio

Chucunaque (MO).

WEST INDIES. TRINIDAD. N. L. Brltton 2153, 6 Mar. 1921, forest border Tal Pane (MY).

7. HELICONIA BIHAI (Linnaeus) Linnaeus., Mant. 2:

211. 1771. Musa blhal L., Sp PI. 11 10^3. I753. Typei

" Hellconla blhal Surln. 80." (London, Linn. Soc. 286.1; microfiche seen.)

Hellconla luteofusa Jacq., Hort. Schoen. p. 25. 1797.

Blhal luteofusa (Jacq.) 0. Ktze., Rev. Gen. PI. 2: 684. I89I. Type: based on Swarz . obs. page 96

/ (not seen) . .

Hellconla nlgrescens Jacq., Hort. Schoen. p. 25.

1797. Typei based on Plum. Gen. page 50.

Hellconla varlegata Jacq., Hort. Schoen. p. 25. 1797.

Type: no type Indicated. Taken from Plumler Gen. page 51. 1703.

Hellconla rut 11a (Griggs, Bull. Torr^ Bot. Club 3O:

657. 1903. Blhal rut 11a (Griggs) Griggs, Bull.

Torr. Bot. Club 3I: ^^55. 190^1. Type: Figure

in Edwards' Bot. Reg. pl.37if. Slender plant, 2-'f m. tall, sometimes slightly taller; somewhat musoid. Leaf-blades oblong, 35*-100 cm. long some- times longer, 10-20 cm. wide, acute-acuminate apex, obtuse

-acuminate base; upper and lower surfaces green and glabrous; long, thick petiole. Inflorescence erect, occasionally bent to nodding position, glabrous, 25-19 em. long; peduncle short or absent, inflorescence extending from sheath; rachis straight to partially fleiuose, 0.9-1.3 cm. diameter; branch-bracts 3-7, rarely more, variegated, bright red with yellow margin, inner surface bright yellow, glabrous, long ovate, thick or succulent, moderately deep boat-shaped, subcordate (somewhat clasping rachis), gently curved out and up; lowest branch-bract usually sterile extending into leaf-blade, total length 27-90 cm.; lowest fertile branch-bracts 15-21, cm. long, 2. 4-3. if cm. side

-width; upper branch-bracts 8 cm. long and 1.5 cm. side

-width; intemode between lower branch-bracts 1.0-1.5 cm.; branch-bracts not overlapping. Ploral-bracts lanceolate, membraneous, 4.0-4.5 cm. long, about 0.5-1.0 cm. wide.

Flowers 4-6 per branch-bract, completely glabrous, greenish

-white to pure white with greenish tips, 5.5-6.0 cm. long; perianth 3 •5-4 .5 cm. long, free sepal curved back; pedicel

0.5-0.8 cm. long. Pruit dark, 3-sided to globose, rough;

1 cm. diameter, 1 cm. long, dried. (Plate 7.)

Common on moist slopes, wet ravines, and wet mountain- sides. 93

PIATE 7. HELICONIA BIHAI (L. ) L.

H. Mhal (L.) L., Silver Spoon Dlst,, St. Vincent. West Indies, 1-7 April 1950. R. A. Howard III63 (GH). I The scope of this study is restricted to the .

However, H. blhal Is much more widespread. The "Flora of

Peru" lists this species, and It Is found throughout

northern South America. (Map 7.) It was one of the first

Hellconlas to be brought Into cultivation during the latter part of the 18th century.

There has been an unusual amount of taxonomlc confu-

sion with this species. This was pointed out earlier

under the discussion of the type specimen of the genus,

since H. blhal Is the type. There have been many horti-

cultural names with little. If any, description given to

this species. These names will be Ignored In this study, since they are of no taxonomlc value.

H. blhal Is closely related In habit to the other species In the Section Hellconla . Aside from having a

different geographical distribution, H. blhal does not

have the extremely deep boat-shaped bracts that are found In other species of this Section, and the variegated

pattern of the branch-bracts Is unique.

Representative specimens:

WEST INDIES. DOMINICA. W. H. Hodge 3^3. U Aug. I938, stream-cut, wooded gorge lying between the estate house and orange plantations, alt. 488 m. Sylvanla Estate (NY). W. H. Hodge 1086, 12 Feb. I9lf0, same location as previous specimen (NY). W. H. and B. T. Hodge 1359, 9 Feb. 19^0, 95

wet wooded ravine at laase of twin waterfalls of Massacre

River between Sylvania and Mt, Jay, alt, 500 m. (GH).

W. H. Hodge 2244. 21 Mar. 1940. upper rainforest-clad

slopes, alt 763 to 1142 m. (GH). W. H. and B. T. Hodge

I63O, 8-10 April 1940. deep rich soil, border of high-stem

rainforests. Syndicate Plantations, close to northwest

slopes of Mome Diablotin, alt. 600 m.. rainfall 5OO cm.

(GH). P. E. Uoyd 123t 1903. Laudat (NY). A. C. Smith

10273, 26 Mar. 1956, vicinity of Fresh Water Lake, near

Laudat, alt. 45O-6OO m.. edge of low forest (US). R. L.

Wilbur, E. L. Dunn, H. A. Hespenheide, D. R. Wiseman 7403,

14 July 1964, forested slopes of Micotrin along trail from

a point about ^ mi. beyond Fresh Water Lake (L»etang),

alt. 2300-2700 ft. (DUKE). R. L. Wilbur, E. L. Dunn, H.

A. Hespenheide, D. R. Wiseman 8211, 3 Aug. 1964, vicinity of Boeri Lake on the northern slopes of Mome Micotrin (DUKE).

DOMINICAN REPUBLIC. Prov. Barahona . P. Fuertes II3, July

1910 (NY, US). Prov. Seibo . N. Taylor 306. 25-26 Nov.

1909 • river bank (F, NY).

GRENADA. J. S. Beard 593. 29 Oct. 1945 (GH). R. A. Howard

10637. 24 Feb. - 6 Mar. 1950, plant often reclaiming hill-

side after landslides, vicinity of the Grand Etang (GH).

St. Andrew Farrish . G. H. Proctor I7O85, 30 Oct. - 11

Dec. 1957. common giant herb in moist gullies. (GH). Distribution of Hellconla Mhal.

1

GU/IDELOUPE. P. Duss 3839. Sept. 1895. Bolsdes (NY). A.

Quested 20^9* 10 Mar. 1938. alt. 600 m. (US).

HAITI. E. L. Ekman H106, 10 June 1917. Dept. of Sud . alt. 800 m. (NY). L. R. Holdrldge 2171. 5-6 Aug. 19^5, RiYlere Glace (MICH).

MARTINIQUE. P. Duss 2119. I896 (NY. US). C. Klmber M, June 1962, roadside verge, dark soil, well-developed

profile. Route de la Trace, Commune de Balata-Tourtet

(WIS).

MONTSERHAT. J. A. Shafer 302. 26 Jan. I907, Chanera Moun-

tain, above 2000 ft. (P, NY, US).

ST. VINCENT. St. Patrick Parish . G. R. Cooley 81 79, 18

Jan. 1962, wet mountainside woods at Lower Bellewood,

elev. 1100 ft. (GH). Silver Spoon Dist . R. A. Howard

11163. 1-7 April 1950, River Valley, above Three Rivers

(GH). R. A. Howard 1120**, 1-7 April 1950. n. slope of

Somma Range at ^•050 ft. alt. Summit of the Soufriere (GH).

C. V. Morton 5^18, 16-25 April 1947. along Chateaubelar

River, alt. 300-400 m. (US). H. H. and G. W. Smith 1699a. March I89O (NY).

III. Section PENDULAE (Griggs) R. R. Smith, sect. nov.

Subgenus Platychlamys Baker in part. Ann. Bot. Vol. VII 190. 1893.

Pendulae Griggs., Bull. Torr. Bot. Club jf2i 316. I915. 99

I (Rank not specified; here Interpreted as series.)

Plants 2-6 m. tall; all with musold habit. Leaf

-blades 50-100 cm. long, sometimes longer; all leaves petloled. Inflorescence pendulous; pubescence varies

from puberulent to woolly; rachls flezuosed to nearly

straight. Branch-bracts from 6 to 30, ovate to ovate

-lanceolate; reflexed to extending out horizontally from rachls. Branch-bracts mostly distichous, few spiralled due to twisting of rachls.

Type species t Hellconla

pendulae Wawra

(Oesterr. Bot. Zeltschr. 13i 8. 1863.)

All the species In this section possess a pendulous

Inflorescence In which the branch-bracts are separated on the rachls. The Inflorescence varies greatly In size, shape and color. There are twelve Central American species

In this section. The type species Is Brazlll8ui.

Key to the Species of Section PENDUIAE

A. Inflorescence with 6-9 widely spaced branch-bracts;

diameter of rachls 0.2-0.4 cm.

B. Branch-bracts red with yellow margin; apices of

branch-bracts blunt 8. H. marglnata

B. Branch-bracts orange to deep red, margin ssune

color as rest of branch-bract; apices of branch -bracts lanceolate to acuminate* 100

C* Rachls puberulent with short white hairs;

perianth k,^ om. long 9» H» colgantea

C* Rachls hirsute with dense brown hairs (Term-

glnous-tomentose ) : perianth either 4.0 cm. or

less or 5*0 cm* or more.

D« Rachls moderately flextiosed, branch

-bracts orange to deep red; perianth

3-^ cm. long 10. H. nuteuis

D» Rachls nearly straight; branch-bracts

deep red; perianth 5 cm. long •

11. H. all en 11

A. Inflorescence with 10 or more closely or widely spaced

branch-bracts; diameter of rachls 0,k cA, or more.

E. Inflorescence completely woolly with golden red

-brown pubescence; branch-bracts strongly or

moderately reflexed.

P. Branch-bracts 20-26, strongly reflexed; lower

breoich-bracts 21-25 cm. long....

12. H. Yellerlgera

P. Branch-bracts l^f, moderately reflexed: lower

branch-bracts 16 om. long. •

13. H. aagniflea

E. Inflorescence villous to tomentose, never woolly;

branch-bracts strongly reflexed to extending out horizontally* 101

G» Branch-lsraots not more than 6*5 cm.

long; all branch-bracts strongly

reflexed l**-. H, curtlspatha

G* Brem oh- bracts more than 6 .5 cm. long;

strongly to moderately reflexed.

H. Inflorescence vlllous-velutlnous,

middle branch-bracts 8 cm. long;

all brsuioh-braots strongly reflexed

giving appearance of being nearly

Imbricated

15. H. xanthotricha

H. Inflorescence tomentose to nearly

glabrous; middle branch-bracts from

8-20 cm. long; branch-bracts

strongly-reflexed to extending out

horizontally from rachis.

!• Inflorescence distichous with

10-27 branch-bracts, usually

more than twenty; lowest fer-

tile breuich-bracts ovate-acute

to OYate-lanceolate; rachis

reflexed.

I J. Peduncle moderately stout

eund pubescent; middle

branch-bracts orate

-acuminate, 4 times 102

longer than wide...

16. H. Paul 11

J* Pedtinole stout and gla->

brous; middle branch

-bracts ovate with blunt

tips, 3 times longer than

wide 17* H. longa

!• Inflorescence spiralled with

8-20 branch-bracts, usually

less than twenty; lowest fer-

tile branch-bract 16-35 om.

long; branch-bracts OYate

-Isuiceolate; rachls somewhat

reflexed and twisted.

K. Dlstsuice between

lower branch-bracts

3 •0-3 •5 OBi»J branch -bracts red....

18. H. colllnslana

K. Distance between

lower branch-bracts

4.5-6.0 cm.; branch

-bracts red and

yellow or solid red.

•• 19* R* oatheta 103

8. HELICONIA MARGINATA (Griggs) Plttler In Manual de las Plant es Usuales de p. 299* 1926. Blhal marglnata Griggs, Bull. Torr. Bot. Club k2t 323 • 1915.

Type I Panama, Marragantl and vicinity, 3-9 April 1908,

R. S. Williams 696 (NY). Isotype (US).

Moderately stout plant, about 4 m. tall; musold habit.

Leaf- blade oblong, 35-60 cm. long, about I5 cm. wide, short acuminate-acute apex, roiinded to acute base; upper and lower surfaces of leaf-blade green; petiole prominent,

^5-50 cm. long, moderately stout. Inflorescence pendulous, puberulent to tomentose, 20-28 cm. long, long peduncle? rachis strongly flexuose, 0.3-0. if cm. diameter. Branch

-bracts 6-9, scarlet with yellow margin, puberulent, OTate

-lemceolate with obtuse apex, moderately shallow boat

-shaped; all branch-bracts strongly reflexed; lowest branch-bract 10 cm. long, next lowest branch-bract about

8 cm. long; upper branch-bracts 6-7 cm. long; all branch

-bracts 1.5-2.0 cm. side-width; branch-bracts angled in various directions. Floral-bracts triangular-lanceolate,

5.5-6.0 cm. long, 1.2-1. 5 cm. wide, slightly pubescent to tomentose on outer surface. Flowers 4-6 per branch-bract, yellow, 5.5 cm. long; perianth 4.0 cm. long, glabrous, free sepal curled toward axis at anthesis; pedicel 1.2-1.5 cm. long. Fruit dark, subglobose 1.1-1. 5 cm. diameter.

(Plate 8.)

Infrequent in wet forests and swamps. i

*• - >

PIATE 8. HELICONIA MRGINATA (Griggs) Plttler

Holotype: Blhal marglnata Griggs, Marragantl and vicinity, Panama, 3-9 April 1908, R. S. Williams 696 (NY). at...

Map 8. Distribution of Helioonia marginata . 106 t

Indicated by the number of specimens collected, this

Is a rare species in Panama. However, it is reportedly common in South America. The Province of Darlen, Panama, may well be the northernmost extension of the range for

H. marginata . (Map 8.)

H. marginata is a clear-out species, and there is no taxonomic confusion involved. Its apparent closest rela-

tives are H. allenii and H. nutans , both of which have solid red branch-bracts.

Representative specimens

PANAMA. Prov. Darlen . J. A. Duke kQl?, 15 June 1962,

Alconorque sweunp on Rio Tuira ca. 3 ml. n.w. of El Real

(MO).

9* HELICONIA COLGANTEA R. R. Smith, sp. nov.

Typet Costa Rica, Prov. Puntarenas, forest and stream edge above Rio Terraba near Interamerican Hwyi, about 10 km. e.n.e. of Palmar Norte (to San Isidro), + kO m. alt.,

26 Jan. I967, W. Burger and G. Matta U. 4662 (P).

Moderately stout plant, 2-'4^ m. tall, musold habit.

Leaf-blade oblong, about 1 m. long, about 24 cm. wide, short acuminate apex, wide obtuse-acuminate base; upper and lower surfaces green; petiole 1 m. or longer. Inflo- rescence pendent, pubescent, 3O-6O cm. longj ped\mcle not seen; rachls nearly straight, puber\aent with short white hairs, 0. 3-0.4 cm. diameter. Branoii- bracts 6, rose-red. 108

puberulent without emd within, ovate-lanceolate, moderately deep boat-shaped, slightly reflexedj lowest branch-bract

25 cm. long, 2.5 cm. side-width, Intemode between branch

-bracts 3.5-5»0 cm. Ploral-bracts 'f-S per branch-bract, pale cream color, 3*5-'f.O cm. long, 0.7-0.8 cm. wide, white pubescence on outer surface, glabrous Inner surface.

Flowers approximately k per branch-bract, yellow, 5«5 cm. long; perleuith ^.5 cm. long, calyx white pubescence, corolla glabrous; pedicel about 1 cm. velutlnous with whltlsh-yellow hairs; ovary yellow; hairy. Maturing fruits dark, Ylllous-velutlnous, somewhat globose, 0.8-0.9 cm. diameter. (Plate 9*)

In moist forests near rivers, endemic to Costa Blca.

(Map 9.)

In general appeare^ice, this new species seems to be quite close to H. allenll and H. nutans « All three species are pendent with similarly shaped branch-bracts widely spaced on the rachls smd showy pale floral-bracts. H. colgsmtea differs from these two species In that the pubescence on the rachls Is shorter, sparse and white, while that on H. nutans and H. allenll Is thick and golden brown. The pedicels and ovaries of H. colgantea are extremely pubescent, while those of the 2 latter species are nearly glabrous. There are also a few less obvious morphological differences. Geographically, H. nutans and COSTA mCA tn ,

PIATE 9. HELICONIA COLGANTEA R. R, Smith, sp, nov,

Holotypei H, colgantea R, R, Smith, Forest above Rio Terra ba, e.n.e, of Palmar Norte, Prov, Puntarenas, Costa Rica, 2 Jan. 1967, W. Burger and G, Matta a ^662 (P). Hap 9* Distribution of Hell con la. colgantea. 111 " t

H. allenll are Panamanian species, while H. colgantea Is a Costa Rlcan species.

The specific name chosen for this new species Is a

Latinized form of the Spanish word "colgante, " which means "hanging.

Representative specimens

COSTA RICA. Prov. Alajuela . A. Jimenez M. 2323. 10

Setlembre I96'i-, Quebrada Zajas. Plnca Los Ensayos, Buena Vista de San Carlos, alt. 85O m. (P). Prov. Cartage . A.

Tonduz 920, XI 1897. Forets de Tuls (US).

10. HELICONIA NUTANS Woodson. Ann. Mo. Bot. Card.

26: 276. 193^. Typei Panama. Prov. Chlrlqul. vicinity of Caslta Alta, Volcan de Chlrlqul. 28 June - 2 July 1938.

Woodson. Allen, and Selbert 968 (MO). Isotypes (GH. MICH, NY).

Moderately slender plant, 2-3 m. tall; musold habit.

Leaf-blade oblong-elllptlc, 35-60 cm. long, 13-2^^ cm. wide acuminate apex, rounded to wide cordate base; upper and lower surfaces green; petiole 20-30 cm. long extending from sheath with purplish membrane. Inflorescence pendu- lous, pubescent, about I9 cm. long; peduncle curved, usually glabrous; rachls moderately fleiuose, 15-25 cm* long, 0.3-0. if cm. thick, long ferruglnous-tomentose.

Branch-bracts 4-8, deep red to orange red, glabrous. .

113

sometimes with tomentose margins, lanceolate, long acumi- nate, shallow boat-shaped} lower breoioh- bracts ll-l? cm. long; upper branch-bracts 6-10 cm. longj all branch-bracts

1.0-1.7 cm. side-width; Intemode between lower branch

-bracts 2.5-3.8 cm.; upper 1.0-2.3 cm. Floral-bracts prominent, ovate-acumlnate, papery appearance, 2-U cm. long, puberulous-vlllous along midrib on outer surface, scattered elsewhere. Flowers 4-7 per branch-bract, yellowish, 3.5-4.7 cm. long; perianth 3-4 cm. long, calyx puberulous, corolla glabrous; pedicel about 1-2 cm. long, sparsely pilose. Fruit dark violet, subglobose, 1 cm. long, 0.8 cm. wide. Seed with sclerlfled, wrinkled seed- coat; 1 cm. long, 0.6 cm. wide. (Plate 10.)

Apparently restricted to cloud forests and moist forests of higher altitudes, 100-2000 m. In the Providence of Chlrlqul, Panama. (Map 10.)

According to collected specimens and literature, this species Is endemic to Panama, and does not extend Into South America.

H. nutsins may be confused with H. allenll . They are both pendent with showy floral-bracts protruding from red branch-bracts. The 2 species differ In that the branch

-bracts of H. allenll are strongly reflexed and those of

H. nutans are not. Other differences, are the longer perianths In H. allenll and the shorter branoh-bracts in

H. nutans PIATE 10. HELICONIA NUTANS Woodson

Holotype: H. nutans Woodson, vicinity of Caslta Alta, Prov. Chlrlqul, Panama, 28 June - 2 July 1938, Woodson et al, 968 (MO), Map 10. Distribution of Helloonla nutans.

Representative specimens}

PANAMA. ProY. Chlrlqul * M. E. Davidson 668, 18 May 1938,

Boquete Dlst. (P. GH. US). H. Plttler 2936. 3 March 1911, forests around El Boquete, alt. 1000-1300 m. (US). J.

Duke, M. Correa, C. Gallegos, and P. Woods I3678 (k), 2

Aug. 1967. TTC - BMI Cloud Forest Litter Study. Cerro

Horqueta, elev. ca. I5OO m. (DUKE). R. E. Woodson, Jr.,

P. H. Allen, R. J. Selbert 968, 28 June - 2 July 1938,

Iclnlty of Caslta Alta, Voloan de Chlrlqul, ca. 1500-2000 m. (GH, MICH. NY).

11. HELICONU ALLENII Standley and Williams, Celba

3t 189. 1953. Type I Panama, Prov. Chlrlqul. Vicinity of Nueva Suiza, Rio Chlrlqul Vlejo, alt. 5OOO ft., 12 June

1951, P. H. Allen 6255 (Herb. Esc. Agr. Panam. and US).

Slender plant about 2.5 m. tall; musold habit. Leaf

-blade long, broadly elllptlc-ovate, about 32 cm. long,

11-18 cm. wide; upper and lower surfaces green and glabrous; petiole at least 14 cm. long. Inflorescence about 25-30 cm. long, pendulous; peduncle short; rachls nearly straight.

0.4-0.5 cm. thick, hirsute with dense brown hairs. Branch

-bracts 6-7, deep red, puberulent along margin and base, lanceolate-ovate, shallow boat-shaped, moderately to abruptly reflexed, 6-14 cm. long, 1.8 cm. side-width;

Intemode between lower branch-bracts 3.5-4.5 cm. Ploral

-bracts numerous, conspicuous, very pale yellow; ovate with •

abruptly narrow, attenxiate-acumlnate tlpj 4 cm. long, 1.5 cm* wldet thickly pubescent along mldveln on outer surface.

Flowers 4-5 per branch-bract, bright yellow, glabrous to hispid; perianth 5 cm. long; pedicel 1.2 cm. long, slightly to moderately hirsute. Ovary yellowish, glabrous, about

0.5 cm. diameter. No fruit examined. (Plate 11.)

Along streams In wet forested areas of Costa Rica and

Panama. Infrequently collected, though one of the more

strikingly beautiful species In the genus. (Map 11.)

The outstanding features are the showy floral-bracts

and the strongly reflexed, widely spaced branch-bracts

This species shows affinities to H* nutans. As stated

previously, H. nutans has smaller floral-bracts and the

orange branch-bracts extend out more horizontally from

the rachls.

Representative specimens*

PANAMA. Prov. Chlrlqul . P. H. Allen 4656, 23 July 1947.

iolnlty of Boquete, liunber road Into the hills e. of Rio

Caldera, alt. 4500-6500 ft. (GH). 119

PIATE 11. HELICONIA ALLENII Standi, and L. 0, Wms.

Holotype: H. allenll Standi, and L. 0. Wmx., vicinity of Nueva Suiza, Prov. Chiriqui, Panama, 12 June 1951, P. H, Allen 6255 (US), Map 11. Distribution of Helicon la allenli. 121 j

122

12. HELICONU VELLERIGERA Poepplg, Reise Chill II

295* I836. Blhal Yellerlgera (Poepp.) 0. Ktze. Rev. Gen.

PI. 2 1 685. I89I. Types Upper , ProT. of Haynas,

Poepplg, In Baker (1893)» Type not seen.

Stout plant k-6 m. talli musold habit. Leaf-blade oblong, 3A- 1 1/2 m. long, 20-30 cm. wide, nearly cordate at base; upper surface green, lower surface green and some-

times with bronze tinge j petloled. Inflorescence pendulous, long golden red-brown woolly pubescence, 75-120 cm. long; long woolly peduncle, 1.2 cm. diameter; rachls fleiuose, woolly, 1 cm. diameter. Branch-bracts 20-26, red or red

-orange, woolly outside, strongly velutlnous-woolly within lower branch-bracts ovate lanceolate, upper branch-bracts ovate, all somewhat cordate; moderately shallow boat-shaped; most branch-bracts strongly reflezed, giving appearance of being almost Imbricated; lower branch-bracts 15-21 cm. long, 3.5-4.0 cm. side-width; intemode between lower branch-bracts 5-7 cm.; middle branch-bracts 8-10 cm. long,

3*0-3.4 cm. side-width; intemode between middle and upper branch-bracts 1.5-2.0 cm. Floral-bracts wide lanceolate, thick, woolly along outer adaxial margin, velutinous over rest of outer surface, glabrous within; 3.5 cm. long, I.5 cm. wide. Flowers 6-8 per branch-bract, yellow; perianth

4,0-4.3 cm. long, calyx with woolly pubescence on upper

1/2 of outer surface of sepals, corolla glabrous; pedicel

1 cm. long, glabrous or with velutlnous-woolly streak. t

123

Fruit deep blue-purple, subglobose about 1 cm* diameter*

Seeds not seen. (Plate 12*)

In moist forests from sea level to 500 m.

In Central America the distribution range Is limited to area near Gulfo Dulce In the south eastern province of

Costa Rica. (Hap 12.)

The description of H* vellerlgera by Poepplg (I836)

Is Vague, but has been accepted by past workers on the

genus. There are other woolly species of Hellconla , but these are quite distinct, and differ considerably from this species In shape of Inflorescence and flower length*

H. vellerlgera Is the largest and most conspicuous species of the pendulous, woolly group.

Further studies on the South American woolly, pendu- lous species are needed In order to verify the conclusion that the entity found In Central America Is H. vellerlgera and not a new species or a possible subspeclflc taxon*

Representative specimens

COSTA RICA. Prov. Puntarenas . P. H* Allen 5235. 8 April 19^9. Golfo Dulce Area, vicinity Esqulnas Experiment

Station; sea level (US). W. Burger and G. Matta U. k772t and k773, 2? and 28 Jan. I967, In shade of (?) secondary forest, steep forested slopes above Golflto along the trail to the television tower, alt* 100-300 m* (F). A* Molina

H., W. C. Burger, A* Jimenez and B* Wallenta 17980, 27 Feb.

1966, dense forest. Tropical Forest of Hlo Convent©, 36 km*

•* of San Isldro El General, alt. 5OO m* (F). PIATE 12. HELICONIA VELLERIGERA Poepp.

H. vellerlgera Poepp., slopes above Golfito Puntarenas, Costa Rica, 2? and 28 Jan. 196?. W. Burger and G. Matta U. 4772 (F). Map 12. Distribution of Hellconla vellerlgera .

127

13. HELICONU MAGNIFICA Morton ex R. R. Smith, sp. nov. Type Panama. I Prov, Panama. Cerro Campana. alt.

1000 m., P. H. Allen ZkZ5, 21 April I9kl (US). Isotype (MO).

Moderately stout plant, about 3 m. tall; musoid habit.

Leaf-blade oblong to oblong-elliptic, 1-1 I/3 m. long, about 50 cm. wide; short acuminate apex, rounded and unequal base; both leaf surfaces green. Inflorescence pendulous, covered with golden-brown woolly-viHous pubescence, about 75 om. long; peduncled; rachis slightly flexuose to straight. Branch-bracts 14, red, woolly; lower ones ovate- lanceolate graduating to ovate, shallow boat-shaped; lowest 16 cm. long, middle about 10 cm. long, upper branch-bracts about 6 cm. long; all branch-bracts about 2 cm. side-width; intemode between branch-bracts on lower part of rachis 3.0-4.5 cm. Floral-bracts 3.5 cm. long; cm. 1 wide, woolly red-brown pubescence along adaxial side of outer surface. Flowers 6-8 per branch -bract, perianth 3.5 cm. long, calyx with woolly-villous golden pubescence covering nearly all of outer surface except lower of lA sepals. Fruit and seed not seen. (Plate 13.)

In dense wet forests of Panama. (Map I3.) This species is apparently quite rare the only col- lection la that of the type. It is a very distinctive ,

128

PIATE 13. HELICONIA MAGNIPICA Morton ex R. R. Smith, sp. nov

Holotype: H. magnlflca Morton ex R. R. Smith, Cerro Campana, Prov, Panama, Panama, 21 April 19^1, P. H. Allen 2425 (US), Map 13 • Distribution of Hellconla magnlflca *

3' 130 131

species In which the outstanding characteristics are the

golden-brown pubescence which covers the entire Inflorescence.

H. magniflca appears to be closely related to H. longa In Its general habit. Both species are pendent, pubescent and have red branch-bracts. H. longa has larger and deeper branch-bracts which are mostly tomentose and usually strong- ly reflexed. H. magniflca possesses shorter and more narrow branch-bracts which are densely woolly and moderately reflexed.

The name for this species was suggested by Morton on the herbarium sheet of the type specimen. There are no representative specimens.

14. HELICONU CUHTISPATHA Petersen In Mart. PI. Bras. 3. pt. 3i 15. 1890. Blhal curtispatha (Peters.) 0. Ktze,, Rev. Gen. PI. 2i 685. I89I. Type, Panama, prope Gatun. m. Jan. Plor, M. Wagner (M. not seen). A tracing of type deposited at Gray Herbarium was seen. large plant, 2-3 m. tall; musold habit. Leaf-blades oblong-ovate. 1 1/2-1 3A m. long, about 48-50 cm. wide, short acute apex, obtuse base; upper surface green, lower surfaces green or slightly glaucous; petiole prominent. Inflorescence pendent, puberulent. I/2-I I/2 m. long; peduncled; rachls usually moderately flexuose. tomentose. 0.5 cm. diameter. Branch-bracts 12-24. deep red. slightly tomentose. ovate, deep boat-shaped; inner surface of 132

branch-bract tomentose; lowest branch-bracts about 5*0~6*5

cm. long, 5-6 cm. side-width; middle and upper branch-bracts

4. 0-5. 5 cm. long; Intemode between branch-bracts 1.5-2.0

cm.; flexuose rachls gives appearance of being almost

Imbricated, but branch-bracts are spaced separately. Floral

-bracts many, light color; some with velutlnous pubescence

on outer adazlal one-half of surface and others uniformly

pubescent; wide lanceolate, 3 cm. long, 1.5 cm. wide.

Flowers about 10 per brsmch- bract , yellow; perianth 3.5

cm. long, slightly vlllous-puberulent ; pedicel 0.8-0.9 cm.

long, glabrous. Fruit dark blue to purple, subglobose to

3>slded, 0.9 cm. In diameter, 1.5 cm. long. Seed sclerl-

fled, rough and wrinkled, 1.5 cm. long, 0.8 cm. wide.

(Plate 14.)

In wooded swamps and wet forested places. Endemic

to Panama. (Map Ik,)

Taxonomlcally, this species Is close to H. rostrata

of Peru. H. rostrata appears to have an erect Inflores-

cence as seen In the Illustration accompanying the original description. The branch-bracts of H. rostrata are larger, and they are variegated, with red sides and green-yellow margins. There are about 18 flowers per branch-bract In

H. rostrata .

As seen In the key to the Central American Hellconlas . H. curtlspatha possesses a habit similar to H. longa . H. xanthotrlcha and H. vellerlgera . Within this group H. curtlspatha has the shortest branch-bracts. 133

PIATE Ih, HELICONIA CURTISPATHA 0. G. Peters.

H. curtlspatha 0, G, Peters,, vicinity of Fort Sherman, Canal Zone, Panansa, 15 Jan. 1924, P. C. Standley 3111? (US), Map Distribution of H*lioonia curtis£atha«

I

t

Some herbarium specimens studied In respect to this species show affinities to other species by having still shorter branch-bracts. This may be a variation within this species or It may be hybridization. Further study and more specimens are needed to understand the species more completely.

Representative specimens

PANAMA. Canal Zone . I. M. Johnston 1639, 10 Nov. 1955, n. base of Mundl Hills, Catlval along Quelvada Morlto, n.w. part of Canal Zone (area w. of Llmon Bay) (GH). ?•

C. Standley 31117. 15 Jan. 1924, wooded swamp, vicinity of Port Sherman (US). Prov. Colon . P. C. Standley 30^^25,

9 Jan. 1924, brushy slope, between Prance Field and

Catlval (US).

15. HELICONU XANTHOTRICHA R. R. Smith, sp. nov.

Type: Panama, forests along R. Boqueron above Peluca

Hydrographlc Station, about 90 m., A. A. Hunter and P.

H. Allen 659. 22 Feb. 1935 (MO).

Stout plant 3-4 m. tall; musold habit. Leaf-blade oblong, 80 cm. long, 36-38 cm. wide, base rounded; upper and lower surfaces green; petiole at least 40 cm. long,

I. 1 cm. diameter. Inflorescence pendulous, villous

-velutlnous, 38 cm. long; peduncle with long golden-brown hairs rachls I flexuose, villous becoming smooth with age. ) 7

0.4 om. diameter. Branch-bracts 15-18, scarlet, outer surface with scattered hairs, Inner surface of branch-bract thickly pubescent with long golden-brown hairs; branch

-bracts strongly reflexed, shallow boat-shapedj lower branch-bracts ovate-lanceolate with obtuse apices, 10-17 cm. long, 2.5 cm. side-width, middle branch-bracts ovate,

7»5-9«0 cm. long; all branch-bracts about 2.5 cm. side

-width; Intemode between branch-bracts 1.5-2.0 cm. Floral -bracts several; yellow; wide lanceolate; woolly-vlllous along narrow, adazlal, marginal strip on outer surface; rest of floral-bract slightly villous to glabrous; k,3 cm. long, 0.6 cm. wide. Flowers yellow, tips exerted, pubes- cent, ^^.0-^f.5 cm. long; perianth 3.5 cm. long; calyx with yellow villous pubescence along upper 2/3 's of outer sur- face of sepals, corolla glabrous; pedicel 0.8-0. 9 cm. long, smooth. Fruit blue, subglobose, about 1 cm. long, 0.6-0. cm. wide. (Plate I5.)

In moist forests of Panama, endemic to Province of Code. (Map I5.

This species Is related to the others with villous floral-bracts and perianths. It Is most closely related to H. vellerlgera In structure and pubescence. It differs from H. vellerlgera In several waysi

1. Generally fewer and shorter branch-bracts. 2. Shorter pedicel.

3» Shorter and more pubescent perianth. 138

H. xanthotrlcha R. R. Smith, El Valle de Anton, Prov. Code. Panama, 5 Aug. 19^6, P. H, Allen 3629 (GH), Map 15. Distribution of Hellconla xanthotrloha*

t a

4. Entire plant of shorter stature.

5* Inflorescence less woolly.

This species appears to be closely related to H.

mutlslana Cuatr. which Is found In Central Columbia. It

characteristics are similar, except the breuich-bracts are villous on the outer surface of H. mutlslana . whereas on

H. lanthotrloha the branch-bracts are neeurly glabrous.

The specific name, xanthotrlcha, was chosen for the

golden-brown and yellow hairs that are abundant on the Inflorescence.

Representative specimens

PANAMA. Prov. Code . P. H. Allen 3629, 5 Aug. 19k6, El

Valle de Anton - Floor - Vicinity Plnca Tomas Arias, alt.

600 m. (GH). R. E. Woodson and R. W. Schery IO5, 30 June.

19^0, north rim of El Valle de Anton, near Cerro Turega, alt. 650-700 m. (MO).

16. HELICONU PAULII Lane ex R. R. Smith, ep. nor.

Type I Panama, Prov. Veraguas, vicinity of Santa Pe, forested slopes of Cerro Tute, 3OOO ft. alt., Zk March 19^7, P. H. Allen 4333 (MO).

Stout plant, about 3-3i m. tall; musold habit. Whole leaves not seen, leaf-blade over 60 cm. long, about 36 cm. wide, base of leaf-blade rounded; upper and lower surfaces green; petiole prominent, striated. Inflorescence pendulous f

142

about 75 cm. long, older Inflorescence not as pubescent as younger ones; peduncle 0.6-0.7 cm. diameter, woolly

-villous with golden-brown hairs; rachls straight 0.5-0.7 cm. diameter, not as villous as pedtincle. Branch-bract a

21-26, terminal half of bracts dark red, basal half of bract yellow or red, ovate-acuminate with obtuse tip,

puberulent to slightly villous along margins, shallow boat-shaped; older branch-bracts becoming moderately

reflexed; lowest branch-bracts 10-15 co» long; middle and upper branch-bracts 5»5-9»0 cm. long; all branch-bracts,

2.0-2.5 cm. side-width; Intemode between branch-bracts

1.5-4.0 cm. Floral-bracts several, ovate-lanceolate,

3. 0-3. 5 cm. long, 1.2-1.3 cm. wide, short tomentose except

along outer, marginal surface which has strip of golden

-brown villous pubescence along adaxlal side. Flowers

3-5 per branch-bract, yellow, 5 cm. long; perlemth about

3*2 cm. long with golden-brown villous pubescence along

outer surface of calyx, corolla glabrous; pedicel slightly

villous, 0.6-0.8 cm. long. Fruit not seen. (Plate 16.)

Moist forest slopes at altitudes of about 1000 meters.

Endemic to Panama. (Map 16.)

This species Is a clear-cut entity. The closest

related plants to this species are H. longa and H.

magnlflea . All three species have similar pubescence

as found on the floral-bracts. H. longa possesses branch

-bracts which are more reflexed and a rachls more flexuose. PLATE 16. HELICONIA PAULII Lane ex R. R. Smith, sp. nov.

Holotype: H. paulil Lane ex R. R. Smith, slopes of Cerro Tutu, near Sante Fe, Prov, Veraguas, Panama, 2h March 19^^?, P. H. Allen 4333 (MO). .1

Map 16, Distribution of Heliconia paixlii .

t •

U6

H. magnlf lea possesses branch-bracts which are generally . smaller and much more Tlllous pubescent.

H. Paul 11 appears to be more toraentose-woolly when young. Therefore, the description In terms of pubescence must be modified when discussing an older Inflorescence as compared to a young one.

• This new species is named in honor of the eminent botemist and collector of the type specimen, Paul H. Allen.

Representative specimens

PANAMA.. Prov. Code . P. H. Allen 2867t 2 Dec. 19^1, region n. of El Valle de Anton on trail to Las Minas, alt. 1000 m. (US).

17. HELICONIA LONGA (Griggs) Winkler in Engl, and

Prantl. Nat. Pflanzenf . 2 Aufl. 15at 536. 1930. Bihai longa Griggs, Bull. Torr. Bot. Club 31 1 I90i»-.

Typet Costa Rica, Prov. Alajuela, El Pez road from Buena

Vista to San Carlos Valley, Cook and Collins U?. I903

(US).

Hellconia pogonanthus Cuf., Archivio Bot. 9t 191

1930. Typet Costa Rica, in reglone Atlantica: in sllva densa ad rirum prope « La Cast ilia - Los Negritos », 12 km. at ore fluminis Reventazon*

Cufodontis 621 (W). Large stout plemts, 2-6 m* tall; musold habit. Leaf

-blade oblong to oblong-elllptlc, 1.5-2.0 m. long, about

50 cm. wide, acute to acuminate apex, rounded at base, sometimes unequally rounded; upper surface green, lower surface waxy, glaucous or green; petiole prominent, about

1 m. long. Inflorescence pendulous, tomentose to veluti- nous, 1/2 -2m. long; peduncle often 1/2-3A ». long, up to 2.5 m. diameter; rachis moderately to slightly flexuose,

tomentose, 0.6-0.8 cm. diameter. Branch-bracts 10-27t

deep red, often yellow near base; lower branch-bracts

ovate-lanceolate, middle and upper branch-bracts ovate

-acute, moderately deep boat-shaped, red-brown tomentose

pubescence without and within; lowest branch-bract 10-18

cm. long, 3*5**'^*0 cm. side-width; middle branch-bracts

b-11 cm. long, 3.5 cm. side-width; upper branch-bracts

5.7 cm. long, 3«0-3»5 om. side-width; intemode between

branch-bracts 2-6 cm. Floral-bracts few to many, 5.O-5.5

cm. long 1/2 cm. wide, dense, light, velvety, woolly

pubescence without, glabrous within. Flowers about 8-25

or 30 yellowish; perianth 3 cm. long, calyx usually with

light woolly pubescence along midrib, corolla glabrous;

on outer surface, pedicel 1.0-1.5 cm. long, glabrous.

Fruit bluish-purple, 1.0 cm. long, 0.6 cm. wide. Seed

not seen.

Inhabits steep moist slopes of forest, wet woods or

swampy forests of Costa Rica and Panama. 148

Collections of this species have been made at alti- tudes of 500 meters on the Pacific side of Central America*

One herbarium label (K. Lems, 5054 (NY)) stated that the habitat was that of subtropical forest zone 600- 7OO m« elevation. Collections have also been made at 25 m* It is apparent that there is diversity in the habitat of the plant.

Tazonomically. H. longa belongs to the pendent group of plants whose branch-bracts are strongly reflezed. There is no confusion with other species.

It is the opinion of this author that, when Cufodontis

(1930) wrote the description of H. pogonanthus , he was not aware of the description of H. longa by Griggs (1904).

Both are very similar in the arrangement of details dis- cussed, and in the measurements taken of the plemt struc- tures. Cufodontis made no mention of Griggs* publication, although he did make reference to Griggs* (I903) publica- tion.

There seems to be a great deal of variation in the length and size of the inflorescence. However, the branch

-bracts remain the same shape and color. The younger ones are more pubescent. This pubescence is somewhat lost on the older plants. The woolly pubescence of the perianth is consistent, and it is an outstanding characteristic of the species. Key to varieties of Hellconla longa

A. About 8 flowers per branch-bract: middle branch-bracts

about 8 cm. long, 3.0 cm. wide var. longa

A. About 20-30 flowers per branch-bract; middle branch

-bracts about 11 cm. long. 3.5-4.0 cm. wide

ar. multlflora

Hellconla longa (Griggs) Wlnkl. van longa

Inflorescence pendulous, variable In size up to 2 meters; peduncle up to 25 om. long. Branch-bracts 7-16 cm. long and 3.0-3.5 cm. side-width. Flowers about 8 per branch-bract, perianth 3 cm. long, pedicel 1.0-1,5 cm. long. (Plate 17.)

Restricted to wet forested areas and slopes of various altitudes from Nicaragua to Panama. (Map 17.)

Hepresentatlve specimens!

COSTA RICA. Prov. Alajuela . A. Molina R., L. 0. Williams.

W. Burger, and B. Wallenta 17219. 16 Feb. I966. near

Artezalea and Methodist Rural Center about 8 km. n.e. of

Villa Quesada, alt. 55O m. (F). A. Molina R.. L. 0.

Williams, W. Burger and B. Wallenta 17422, 18 Feb. I966, forests along Rio San Rafael, near hot springs. Hacienda la Marina, about 14 km. n.e. of Villa Quesada, alt. 5OO

Prov. Cartap^o . K. Lema 5054, 21 Jan. 1964, 150

• as

PIATE 17. KELICONIA LONGA (Griggs) Winkl. var. LONGA Holotype: Bihai longa Griggs var. longa . El Pez road Buena Vista from to San Carlos Valley. Costa Rica. I903. Cook and Collins (US). subtropical forest zone along road from Turrlalba n. to

Pavones and Slqulrres, 600-700 m. elev. (NY). ProY.

Guana caste . P. L. Stevens 766, 8 July 1923, Santa Cecelia

(US). ProY. Llmon . P. C. Standley and J. Valeric i*898'f,

18-19 Feb. 1926, swcunpy forest, Plnca Monte Crlsto on the

Rio Reventazon below Cairo, alt. ca. 25 m. (US). PrOY.

Puntarenas . R. H. Allen 6289, 3 Aug. 1951. Esqulnas

Forest, alt. 30 m. (GH). A. Brenes 12325a, 12 Aptll 1930,

Golflto de Osa (F). W. Burger and G. Matta U. 'f769. 27

and 28 Jan. 1967* on steep slope In forest, aboYe Golflto

along trail to the television tower, 100-300 m. alt. (P),

W. Burger and G. Matta U. 4771, 27 and 28 Jan. I967. In apparently secondary growth forest In shade, steep forested

slopes above Golflto along the trail to the television

tower, 100-300 m. alt. (F). Prov. unknown . A. Tonduz

9433. 1895. Talamanca (US). A. Tonduz 12923. Dec. I898.

Bords du Rio de Las Gueltas, Tucuerlque (US).

NICARAGUA. Dept. Zelaya . P. C. Standley 19^07, 23 April

- 14 May, 1949. wet mixed forest, vicinity of El Recreo,

on Rio Mlco, alt. 30 m. (F).

PANAMA. Prov. xmknown . C. E. Smith, Jr. and H. M. Smith

3447, 4 Jan. 1958, Santa Rita Hills, out-over secondary

forest (F). Map 17. Distribution of Hellconla longa » 153 15^

Hell con la longa (Griggs) Wlnkl. var. mult 1 flora R. R. Smith var. nov~ '

Type: Panama, Canal Zone, Rio Agua Salud, near Prljoles,

6 Mar. 1923. C. V. Piper 6O36 (US).

Inflorescence pendulous, 2 meters or more, peduncle up to 90 cm. long. Branch-bracts 7-18 cm. long; 3 •5-^*0 cm. side-width (always wider than var. longa). Flowers numerous, up to 20 to 25 per branch, perianth not seen, pedicel 2.0-2.5 om.

Infrequent In moist wet forested areas. Panama.

Type specimen Is the only specimen studied. More collections and work needs to be done on this taxon.

18. HELICONU COLLINSUNA Griggs, Bull. Torr. Bot.

Club 30 I 648. 1903. Blhal colllnslana (Griggs), Bull.

Torr. Bot. Club 3I1 445. 1904. Typei Guatemala, Dept.

Alta Verapaz, near Flnca Sepaculte, 0. P. Cook and R. P.

Griggs 352, 30 March I9O2 (US).

Large, stout plant, 2-5 m. tallt musold habit. Leaf

-blades elllptlc-ovate to nearly oblong, i-l^ m. long,

30-50 cm. wide, acmnlnate to acute apex, obliquely cordate to obtuse base; upper surface green, lower surface usually glaucous, sometimes green, often with fine pubescence along midrib; petiole from about 18 cm. to 1 m. long, stout. Inflorescence pendent, tomentose to velutlnous along peduncle and rachls, 35-70 cm. long; peduncle

Tarlable In size, up to about 30 cm. long, glabrous. 156

H. pendula Is a Brazilian species with dark brown to purple branch-bracts, and. reaches a height of approximately 3 m.

H. collinsiana is a northern Central American species extending from Mexico to Costa Rica.

Key to varieties of Heliconia collinsiana

Peduncle smd rachls glabrous to slightly tomentose; pedicel up to k cm. in length, nearly glabrous

var. collinsiana

Peduncle and rachls with long red-brown hairs; pedicels up to 2.5 cm. in length, tomentose var. velutlna

Heliconia colllnsletna Griggs var. collinsiana

Leaf-blades usually glaucous; pedimcle stout. Inflo- rescence pendent, tomentose along peduncle and rachls; usually '^5-70 cm. long; rachls slightly flexuose, 0.3-0.7 cm. diameter. Branch-bracts bright to deep red; glabrous except along outer tomentose surface near base. Flowers bright yellow. (Plate 18.)

This is the more common of the two varieties, and occupies a wider range. (Map 18.)

This is the variety which is believed to hybridize with H. schiedeana . One Mexican specimen (H. E. Moore,

Jr. 620^+ (GH)) was examined which seems to indicate that hybridization took place. Hybridization with other species is discussed in a later chapter. 157

PIATE 18. HELICONIA. COLLINSIANA Griggs var, COLLINSIAM

H. coll Ins lana Griggs var, collinslana t Masagiia, Depart. Esculntla, Guatemala, M. April I89O, J. D. Smith 2102 (US). j

158

Representatl7e specimens

BRITISH HONDUHAS. Dlst. Stann Creek , P. H. Gentle 2106,

20 July 1937. base of hill. Stann Creek Railway (MICH. NY).

W. A. Schlpp 375. k Sept. 1929. growing In secondary forest,

Middlesex, alt. 200 ft. (P, GH. MICH, MO. NY).

COSTA RICA. Prov. Cartago . W. R. Hatch l6l. 22 July 1937,

along trail. "Las Animas" near Turrlalba (P). Prov.

Heredla . A. Jimenez M. 20*^8. 10 Julio 1964. Catarata El

Angel, Rio La Paz Grande, entre Carlblanoo y Vera Blanca

(P).

EL SALVADOR. Dept. La Llbertad . M. C. Carlson 162, 19

Jan. 19^6. Plnoa Santa Maria, w. of Santa Tecla. on

northern slopes of a mountain, alt. 750 m. (P). Dept. San

Salvador . S. Calderon I85I, Sept. 1923. Collna de Santa

Tecla (US). P. C. Standley 2055I. 2-7 Feb. 1922. thicket

along stream, vicinity of San Salvador (US). Dept. San

Vicente . P. C. Standley 21751. 2-11 March 1922. vicinity

of San Vlncente, along stream, alt. 35O-5OO m. (GH, NY,

US). Dept. Sonsonate . P. C. Standley 19330, 26, 27 Feb.

1921, Rocky stream bank, Plnca Chllata (GH, NY, US).

GUATEMALA. Dept. Alta Verapaz . W. R. Maxon and R. Hay

3I87, 5-6 Jan. I905. vicinity of Secanquln, alt. ca. 55O

m. (US). P. C. Standley 70639. 5 April 1939, damp thicket,

near Pancajche, alt. ca. 36O m. (P). H. von Tuerckhelm

8*^68, m. July I903, Cubllqultz, alt. 350 m. (P, GH, NY.

US). Dept. Chlmaltenango . J, R, Johnston 993. 17 Sept. 159

1937. Varledades (P). J. A. Steyermark 31128. k Nov, I939.

Rio Grsmde (Blo Concepolon)* on Socorro Mountain, above

Flnca San Jose, s.e. of Conoepclon de las Ninas, alt.

1200-1700 m, (P), Dept. Esoulntla * J. D. Smith 2102, m. April I89O, Masagua, alt. kOO ft. (US). P. C. Standley

650lf2, 9 Peb. 1939. wet thicket. La Trinidad, on road

between Esculntla and Plnca Zapote, alt. ca. 780 m. (P).

Dept. Huehuetenango . J. A. Steyermark 493^^2, 23 July 19^2,

thickets near Rio Izcan, between Ixcan and Rio Ixcan, Sier-

ra de los Cuchumatanes , alt. I5O-2OO m. (P). Dept. Izabal .

P. C. Standley 2'*'f80, 15-31 May 1922, In swamp, vicinity

of Qulrlgua, alt. 75-225 »>• (GH, US). Dept. Quezaltenango .

W. A. Kellerman 6073. 17 Jan. 1907. El Palmar, alt. 2300

ft. (MICH. US). W. A. Kellerman 6078, 17 Jan. I907, El

Palmar, alt. 2300 ft. (P). Dept. Retalhuleu . P. C.

Standley 87338. 17 Peb. 19^1, wet thicket. Rio Talculan,

along road 5 km. w. of Retalhuleu. alt. ca. 300 m. (P).

Dept. San Marcos . P. C. Standley 68796, 1^ March 1939.

wet thicket Rio Mopa, below Rodeo, alt. ca. 600 m. (P).

Dept. Santa Rosa . P. C. Standley 60706, 20 Dec. 1938,

wet thicket, valley of Rio de los Esclavos, near El Mollno,

alt. ca. 600 m. (P). P. C. Standley 79130, 3 Dec. 19^*0,

region of Platanares, between Taxlsco and Guazacapan, alt.

ca. 220 m. (P). Dept. Suchltepeque . J. D. Smith 280^^,

m. April 1892, Rio Sis, alt. I300 ft. (US). P. C. Standley

622^^^, 6 Jan. 1939* wet thicket, along Blo Madre Vleja, 160

above Patulul, alt. k^O m. (P). J. A. Steyennark ^^7633,

17 June 19^2, virgin forest near Santiago farm, vicinity

of Tlqulsate, alt. 100 m. (P). J, A. Steyermark 'f6777.

28 May 19'*2, eastern slopes of Volcan Santa Clara, above

Chlcacao, alt. 1,250 m. (P).

HONDURAS. Dept. Atlantlda . P. C. Standley 7715. 22-27

April 19^*7. dense wet mixed forest, vicinity of San Alejo,

150-270 m. at base of hills s. of San Alejo near Rio San

Alejo (P). T. G. Yuncker, J, M. Koepper, and K. A* Wagner

8537. 19 July 1938, In rocky ravine, slopes near Puente

Alto on S. P. Co. R. R., e. of Celba. 800 ft. alt. (P,

GH. MICH, MO, UC. US).

MEXICO. State Chiapas . E. Matuda I6693, 11 July 19i^7,

Esperanza, Esculntla (P). E. Matuda I8OI9, 2 July 19k6,

In wet bush, Esperanza, Esculntla (P). E. Matuda 3823, 16-18 July 1939, In advanced forest, Aguacate, Palenque

(GH, MICH). States Mlchoacan and . E. Langlasse

8lif, 28 Jan. 1899. Sierra Madre, 5-800 m. alt. (GH).

State . G. Martlnez-Calderon 99. July 1940 - Peb.

19^1. Dlst. Tuxtepec, Chlltepec and vicinity, alt, ca.

20 m. (US). H. E. Moore 8111. 10 Oct. I959, Ravines 35 kms. e. of Math las Romero, 4 kms. w. of Palomares on trans -Isthmus highway (GH). State Veracruz . J. V. Santos 2239,

18 July 19'*^3. at edge of second growth forest, east side of

Oampo Experimental de Hule, El Palmar, Zongolloa (MICH). I

161

J. V. Santos 3632, 19-20 Nov. 1944, wooded area on n. side

of airdrome, Campo Experimental de Hule, El Palmar, Zongollca (MICH).

NICARAGUA. Dept. Managua . P. C. Standley 838O, 1? May

1947. In forested quebrada along road between El Crucero

and house of Flnca Santa Julia, 6OO-9OO m. (P),

Hell con la coll Ins iana Griggs Var. velutina Lane ex R. R. ' Smith, var. novT " —\

Type J Guatemala, Dept. Quezaltenango; between Plnca

Plrlneas and Finca Soledad, lower south-facing slopes of Volcan Santa Maria, between Santa Maria de Jesus and Calahuacle, alt. 1300-1400 m., 5 Jan. 1940. J. A. Steyer-

' mark, 33575 (p). .

Hellconla platystachys Baker, Ann. Bot. VII: I94, I893. Type: Guatemala, Dept. Guatemala, Feb.

I89O, J. D. Smith I873 (K).

Leaf- blade usually green on under surface; peduncle moderately stout. Inflorescence pendent, velutinous along peduncle and rachis; usually 3O-6O cm. long; rachls straight to slightly flexuose, 0.4-0.6 cm. diameter. Branch-bracts cardinal red; lower portion of branch-bract tomentose. Flowers pale yellow. These two varieties of H. coll ins iana are very simi- lar. The only outstanding characteristic distinguishing the two Is the pubescence found on the peduncle and the 162

raohls of var. velutlna . This variety appears restricted

to Guatemala and .

Representative specimens!

EL SALVADOR. Dept, Ahuachapan * P. C. Standley 20081,

17-19 Jan. 1922. Sierra de Apaneca, in the region of Pinca

Colima (US). Dept. Santa Ana . J. M. Tucker 1291, 25 April

19^2. dense undergrowth, moist canyon slope w. side Cerro

del Aguila, alt. ca. I65O m. (P, MICH, US).

GUATEMALA. Dept. Escuintla . P. C. Standley 64533, 5 Peb,

1939 1 wet forest, Pinca Monterrey, s. slope of Volcan de

Puego, alt. 1140-1260 m. (P). Dept. Quealtenango . W. A. Kellerman 6073. 19 Jan. I907. Volcano Santa Maria, 4500

ft. (P). w. A. Kellerman 6076,19 Jan. I907, Volcano

Santa Maria. 4500 ft, (US). P. C. Standley 68427. 11

March 1939 1 damp forest, Pinca Pireneos, below Santa Maria de Jesus, alt. I35O-I38O m. (P). Dept. Santa Rosa .

J. D. Smith 4635. m. MaJ. 1893. Cenaguilla, alt. 4000 ft.

(US). Dept. Suchitepeguez . A. P. Skutch 2101, 10 Jan.

1935. in bushy growth at edge of Barranoo. Pinca Moca, alt. 5000 ft. (GH). Map 18. Distribution of Hellconla oolllnslana.

165

19. HELICONIA CATHETA R. R. Smith, sp. nov. Typei Panama, Near Old Port Lorenzo, mouth of Rio Chagrea. 8 March 1923. C. V. Piper 6037 (US).

Moderately stout plant. 3-5 m. tall; muaold habit. Leaf-blade oblong. 3A - 1 3A m. long, I5-36 cm. wide, slightly acuminate-obtuse apex, obtuse-truncate base, upper and lower surfaces of leaves green; petiole 1/2 - 1 m. long, stout. Inflorescence pendent, elongated. 70-100 cm. long; peduncled; raohls moderately fleiuose and spiral,

golden-brown puberulent pubescence. 0.*f-0.5 cm. diameter. Branch- bracts 10 to 20. scarlet at base, yellow-green at apex, crimson patches on sides, or completely red. pubes- cent over outer surface of bract or Just around the base,

ovate-lanceolate, moderately deep boat-shaped; lowest branch-bract about kO cm. long, sterile; lowest fertile branch-bract 26-28 cm. long, middle branch-bracts gradating from 18-20 cm. long, upper branch-bracts 6-10 cm. long, all branch-bracts 2.0-3.5 cm. side-width Intemodes between lower branch-bracts 5-6 cm. Ploral-bracts lanceolate with translucent or thick border, puberulent. k cm. long. 0.8 cm. wide. Flowers 6-8 per branch-bract, yellow, 6.0-7.0 cm. long; perianth 4-5 cm. long, lower 2/3rds of calyx puberulent. upper l/3rd glabrous, ovary drying dark, puberulent or glabrous, pedicel 0.9-1.3 cm. long, strongly puberulent. Prult dark blue, hairy, subglobose, 1.2-1.3 cm. diameter when dried. 166

In openings of moist forests at low altitudes. Re-

stricted geographically to Panama. May possibly extend southward into Columbia, South America.

This Is a new species xmllke any of the other Central American Hellconlas . The closest related pendent Hellconla Is H. colllnslana . which possesses long, narrow branch

-bracts. Its range Is considerably north of this Panama- nian plant.

The specific name of this new species Is taken from

the Greek work "cathet, -o," which means "hanging down."

Key to varieties of Hellconla catheta

Branch-bracts red or scarlet at base, yellow to yellow-green

margins, sides glabrous; pedicel tomentose; ovary wall

pubescent v^j.. catheta

Branch-bract red, sides pilose to slightly villous; pedicel velutlnous-tomentosej ovary wall nearly glabrous

var. pubescens

Hellconla catheta H. R. Smith, sp. nov. var. catheta

Inflorescence tomentose along rachls; 70-90 cm. long; peduncles; rachls slightly flexuose, O.k-0.5 cm. diameter.

Branch-bracts 16-20, scarlet or red at base, crimson patches on sides, yellow to yellow-green along margins, wide lanceolate with obtuse apex, tomentose only at base; lower branch-bracts 4.5-6.0 cm. distance apart on rachls; ) t

flowers 6.0-7.0 cm. long; perianth U-5 cm. long; pedicel

1.5 cm. long, tomentosej fruit dark, moderately pubescence (Plate 19.)

Forest floors in low moist areas. Restricted to i

Panama. (Map I9.

This is the more common of the two varieties.

Representative specimens

PANAMA. Canal Zone. A. M. Chlckering 16, 16 Aug, 1928, Barro Colorado Island (MICH). D. Pairchild 17, Aug. l92^^,

Barro Colorado Island (P). L. A. Kenoyer 232, 12 July

1927, in openings in forest, Barro Colorado Island (US),

S. McDaniel 239^. 25 June I966. vicinity Nuevo Emperador

(PSU). C. V. Piper 5856, 6 Mar. 1923. Rio Agua Salud, near Prijoles (US). W. R. Stimson 5068, 12 June I967, in forest, Albrook Air Force Base Research Forest Site (DUKE). Prov. ChiriQUi . M. E. Davidson II79, 18 Aug. Puerto 1938, Armuelles (F). r. e. Woodson, Jr. and R. W. Shery 89I, July 28 - August 1, I940. vicinity of San

Bartolome. Peninsula de Burica, alt. O-50 m. (MO). Prov. Darien. W. L. Stem, K. L. Chambers, J. D. Dwyer, J. E. Ebinger 1*27. 12 June 1959. vicinity of Paya, Rio Paya (GH. MO. US). W. L. Stem, K. L. Chambers, J. D. Dtryer, J. E. Ebinger 925. 4 July I959, vicinity of Oampamento Bueno Vista. Rio Chucunaque above confluence with Rio Tuquesa (US, GH). Prov. Panama . 26726. 11 Dec 168

'^jiiiijiiii|iiri|iiii|iiii|iiiijiiii|iiiijiiii|iiii lllllllll wmf

fcwa eu l*lla»iil% (pp. tnm Cual l»a PUHT* or fAHAUM

C. T. Plpar.

PIATE 19. HELICONIA CATHETA R. R, Smith, sp. nov. var. CATHETA

Holotype: H. catheta R. R, Smith, sp. nov. var. catheta, near Old Fort Lorenzo, mouth of Rio Chagres, Panama. 8 March 1923. C. V. Piper 6037 (US). 169

1923. near big swamp e. of Rio Tecumen (US). Prov. unknoKn , 0. Kuntze 1919. M-8, IB?^ (NY).

Hellconla catheta R. R. Smith, sp. nov. var. pubescens

Smith, var. nov ^ R. R. .

Type: Panama, Prov. Code, P. H. Allen 36l*^, 5 Aug. 19'*6.

£1 Valle de Anton-Floor-Vicinity Flnca Tomas Arias, alt.

600 ffl. (US).

Inflorescence with short scattered hairs (moderately

villous ) along raohls and on branch-bracts with narrow villous margin bordering branch-bract; 75 cm. long

(specimen seen not fully mature); rachls slightly flezuose,

0.5-0.6 cm. diameter. Branch-bracts at least 10, red, pilose to slightly villous; lower bremch-bracts 3»0-3»5 cm. distance apart on rachls. Flowers ^.3 cm. long; perianth 3.0 cm. long; pedicel 0.9 cm. long, velutinous

-tomentose. Ovary nearly glabrous.

Forest floors in El Valle de Anton.

Only one specimen of this taxon was seen, and it appears to be close to the new H. catheta var. catheta .

After a further study of more specimens, it may be decided that H. catheta var. pubescens should become a species in its own right. More collections are needed. 1 i-5

Hap 19. Distribution of Heliconla catheta .

172

IV. Section STENOCHIAMYS (Baker), R. R. Smith, sect. nov.

Subgenus Stenochlamys Baker., Ann. Bot. Vol. VII: 190.

1893.

Plants 1-3 m. tall, musold or cannoid habits. Leaf

-blades elliptical, oblong to oblong-ovate; petloled or

sessile. Inflorescence erect or nodding; rachls straight

or flexuose, sometimes twisted. Branch-bracts k~22, red

to yellow; extremely variable In length.

Type species i Hellconla cannoldea

L. C. Rich.

(Nova Acta 15, Suppl.: t. 9-10. 1831.)

This Is the largest section in the genus. There are

19 Central American and West Indian species, a few of which

occur in South America. All the species have a rachls with

a diameter of one cm. or less, and have an erect Inflores-

cence. In some species the Inflorescence may be extended

at an angle, but It is not of a pendulous nature. The

type species is a South American plant.

Key to the Series of Section STENOCHIAMYS

A. Plants with slender, cannoid habit; leaf- blades less

than 50 cm. long, usually sessile, sometimes petloled:

Inflorescence less than 15 cm. in length (H. vaginalis

sometimes produces sessile leaf-blades of 60-80 cm. In

length, slender habit retained) Cannoideae A* Plants with musoid habit t leaf-blades more than 50 cm.

long, petloled; Inflorescence more than 15 cm. In

length.. Dlstantes

Series Cannoldeae

Cannoldeae Griggs, Bull. Torr. Bot. Club kZt 316.

1915* (Rank not specified; here Interpreted as

serlea)

Plants 1-5 m* tall; slender cannold habit. Leaf-blades less theui 50 cm. long, except H. vaginalis In which leaf

-blades sometimes reach 80 cm.; usually sessile or nearly so, sometimes with slender petioles. Inflorescence erect,

15 cm. long or less, often glabrous. Branch-bracts 3-6, variable In length, red or orange (yellow In H. hlrsuta ).

Type species t Hellconla

cannoldea L. C. Rich.

Seven Central American species are In this series,

four of which also have a range extending Into South

America. They all possess a narrow, lanceolate type of branch-bract and are of a cannold habit.

Key to the Species of Series Cannoldeae

A. Plants up to 1 m. tall (occasionally slightly taller); all leaves with slender petioles

20. H. pslttacorum •

Plants more than 1 m» tall; leaf-blades sessile or lower stem leaves with slender petioles.

B. Branch-bracts orange; flowers white or cream-white.

C. Perianth 4.5 cm. long; lower stem leaves petloled 21. H. aurantlaca

C. Perianth 5.5-6.0 cm. long; all leaf-blades

sessile 22. H. leucoflora

B. Branch-bracts straw-yellow, red-orange or red;

flowers yellow or red.

D. Branch-bracts straw-yellow or red; perianth yellow and hirsute or red

and nearly glabrous.... 23 • H« hlrsuta

D. Branch-bracts red; perianth yellow and glabrous

E. Lowest branch-bract usually ex- panded Into large leaf-like blade;

floral-bracts 3.0-3.5 cm. long

24. H. vaginalis

E. Lowest branch-bract not expemded

Into leaf-like blade; floral-bracts

either less than 3 cm. long or more

than 4.5 cm. long. .

P. Plants 1.5-2.0 m. tall; floral

-bracts 4.5-5»5 cm. long;

perianth 3»5-5»0 cm. long with

yellow-green tip

25. H. trlnldatls .

p. Plants to k m. tall; floral

-bracts 2.8 cm. long; perianth

2.5-3.4 cm. long with black

tip 26. H. swartzlana

20. HELICONIA PSITTACORUM Linnaeus f.. Suppl. Syst.

Veg. 158. I78I. Bihala pslttacorum (L. f.) 0. Ktze. Rev.

Gen. PI. 2: 684. I89I. Typei "Habitat In Surlnamo. C.

G. Dalberg." (London, Linn. Soc. 286.4; microfiche seen.)

Above type Is a lectotype selected here. A second specimen

In the Llnnaean Herbarium, 286. 5 t also Hellconla psltta-

corum , was available to Linnaeus f., and was annotated by

him as "Hellconla 13 nova."

Hellconla pumlla L. ex Alsen, PI. Surln. p. 6. 1785*

Type: Surinam. 13 . Llnnaean Herbarium 286.4,

not seen. Linnaeus first called this species

Hellconla pumlla on specimen 286.4 In the Llnnaean

Herbarium (Savage, 19^5). He did not publish this

species. This name was also used by Jacob Alsen

In a thesis entitled "Plantae Surlnamenses" written

In 1775* and published In I785. Therefore,

Hellconla pslttacorum , published by Linnaeus f

(1781) becomes the valid name for this entity.

Small plant, ^-1 m. tall; somewhat cannold habit.

Leaf-blade oblong-ovate, 25-36 cm. long, 4-7 cm. long,

acuminate apex, obtuse base; upper and lower surfaces 176

green; petiole variable In length, petiole of lower leaves up to 22 cm. long, petioles of upper leaves about 5 cm. long. Inflorescence erect, glabrous, 7-9 cm. long; peduncle 26-55 cm. long, 0.3-0.^4' cm. diameter; rachls slightly flexuose, 0.1-0.2 cm. diameter. Branch-bracts

3-^1, orsuige, glabrous, lemceolate, shallow boat-shaped, pointed In upward direction; lowest branch-bract 7»0-8.5 cm. long; upper brcmch-bracts 2.5-5 cm. long; all branch

-bracts 0.5-0.6 cm. side-width; Intemode between lower branch-bracts 1.0-1.8 cm. Ploral-bracts deciduous, not seen. Flowers ^-6 per branch-bract, yellow with black tip, 4.0-5,0 cm. long; perianth 3 •0-3. 5 cm. long, glabrous; pedicel about 1 cm. long, glabrous. Fruit dark, subglobose^

0.6 cm. long. (Plate 20.)

Moist woods, and along stream banks.

Since this species Is small. It Is used extensively

In cultivation. Throughout the tropics It Is found In public and private gardens, along roadsides and In parks.

The natural range for this species Is northern South

America to Brazil, extending to the Islands of Guadeloupe and Martinique. (Map 20.)

H. pslttacorxun Is a name that has been used for several species. A few Central American species which have been called H. pslttacorum are H. aurantlaca . H. leucoflora . H. hlrsuta and H. vaginalis . A number of

South American species have been Included In this grouping also. .

i 177

PIATE 20. HELIC0N3A PSITTACORUM L. f

H, pslttacorim L. f., Tobago, West Indies, 19 Iferch 1909, W. E. Broadway 2992 (NY). 178 4

The present Interpretation for H. pslttacorum is that it is a small speoies about a meter tall in which the leaves have distinctive petioles, the leaf-blades are rounded at the base, the branch-bracts are orange, and the perianth Is yellowish with a black apex. The species « Is more closely related to H. cannoidea of South America than the species mentioned in the above paragraph*

Representative specimens}

WEST INDIES. GUADELOUPE. P. Duss 3318. 1893-1895 (NY,

US). H. Stehle 8 Jan. 1957. Laterites - Limite des cultures et de I'anciene for%t dense. La Roche Blanche,

Petit Bourg, alt. ^fOO m. (UC, US).

JAMAICA. Vto. Harris 10923. 1*^ Sept. 1910, cult. Hope

Gardens (NY). R. A. Howard and G. R. Proctor 13518, 20-31

Dec. 1953 t growing along stream, cultivated at Shaw Park

(GH). E. West and L. Arnold 778, 21 July I952, Hope

Botanical Garden (PLAS).

MARTINIQUE. P. Duss if 677. 1903 (NY, US). C. Kimber 1457-B,

30 April 1963. Littoral woodland on metamorphosed limestone and derived soil. The lighthouse area, Pointe de la Gbra- velle. Commune de la Trinite (WIS).

TOBAGO. W. E. Broadway 2992, 19 March I909, Scarborough

(MO, NY). Eggers 5if76, Oct. I889 (US).

TRINIDAD. N. L. Britton, E. G. Britton, and T. E. Hazen

90, 27 Feb. 1920, Piarco Savanna, s. of Arouoa (NY). Map 20, Distribution of Hellconla pslttacormn .

E* A. Friend IO5, k Sept. 1939 1 Royal Bot. Gardens, Port

-of-Spaln (NY). 0. T. Pulfer 2, 2^ Nov. I963. shady road-

side, Maraval Rock Quarry (PSU). E. H. Graham 54?, 18

Aug. 1924, 6 ml. n.e. of Saugre Grande (NY).

21. HELICONU AURANTIACA Ghlesbreght. In Lemalre's

L'lllustr. Hortlo. PI. 332. 1862. Blhal aurantlaca

(Ghlesbr.) Griggs, Bull. Torr. Bot. Club 3I1 445. 1904.

Typet Illustration with original description. Specimen obtained "dan les forets dur Mexlque meridional."

Hell con la brevlspatha Hook. In Curtis* s Bot. Mag. t.

5416. 1864. Typet Illustration with original

description. Specimen under the name Helicon la

aurantlaca sent from Verschaffelt • s garden In Ghent to Hooker.

Hell con la choconlana W. Watson In Proc. Amer. Acad.

23i 284. 1888. Blhal choconlana (S. Wats.)

Griggs. Bull. Torr. Bot. Club 31t 445. 1904.

Typet Guatemala, eastern portions of Vera Paz

and Chlqulmula, I885, S. Watson 193 (US. GH).

Hellconla crassa Griggs. Bull. Torr. Bot. Club 3O1

646. 1903. Blhal crassa (Griggs) Griggs. Bull,

Torr. Bot. Club 31» 445. 1904. Types Guatemala.

Alta Vera Paz. road from Secanquln to Sepaclute,

1 April 1902. 0. P. Cook and R. P. Griggs 376 (US). 162

Slender plant, 1-2 m. tall; cannold habit. Leaf-blades oblong to oblong-elllptlcal, 18-35 cm. long, 5.5-8.0 cm. wide, ouspldate-long acuminate apex, obtuse to subcordate base; upper euid lower surfaces of leaves green, upper leaf-blades usually sessile to clasplngt lower leaf-blades with short petiole. Inflorescence erect to slightly nodding, glabrous, about 4-11 cm. long; peduncle about

1-2 cm. long, 0.4-0.5 om. diameter; rachls straight to slightly flexuose, 0.2-0.3 cm. diameter. Branch-bracts

4-5, oremge or greenish with orange base, glabrous, wide lanceolate, shallow boat-shaped, moderately curved upward; lowest branch-bract usually sterile and often expanded

Into leaf-blade, orange colored portion 5-12 cm. long, and green leaf-blade 4-18 cm. long If present; second lowest branch-bract sometimes sterile, lower fertile branch-bracts 5.5-9.0 cm. long, 0.5-1,5 cm. side-width; upper branch-bracts about 3.5-4.0 cm. long, 1 cm. side

-width; Intemode between lower branch-bracts 1.0 cm.

Flowers 4-7 per branch-bract, cream to yellow, 5.0-6.0 cm. long; perianth 3.5-4.5 cm. long, glabrous; pedicel

0.8-1.5 cm. long, glabrous. Prult dark blue, about 1 cm. diameter when dried, somewhat glabrous and 3-sectloned»

Seed 0.6-0,7 cm. long, 0.6 cm. wide, 2 Inner adjacent sides with flat surface, outer surface round, nearly globose; rough with sclerified coat. (Plate 21.) 183

PIATE 21. HELICONIA AURANTIACA Ghiesbr.

H. aurantiaca Ghiesbr. near Tela, Dept. Atlantida, 6 Dec, 1927 - 20 March 1928, P. C. Standley 56717 OF). t

Common In woods near streams, or at edge of moist to wet forest. Distribution from southern Mexico to central provinces of Costa Rica. (Map 21.)

There Is no taxonomlc confusion with this species.

Its outstanding features are the orsoige bracts and cream to yellow colored flowers. It Is closely related to H.

leucoflora , a new species. It Is readily distinguishable from H. leucoflora by having much shorter flowers, which are more Intensely yellow. The range for each species differs. H. leucoflora Is restricted to the southern part of Costa Hlca and Panama*

Representative specimens

BRITISH HONDURAS. District El Cayo . P. H. Gentle 2393.

23 March 1938. on hillside, Vaca (GH, MICH, MO, NY). C.

L. Lundell 6219 1 June->July 193^, In advanced forest, lime-

stone valley, Valentin (GH, MICH, NY. US). District

Corozal . H. H. Bartlett 13030, 6 May 1931. San Antonio

(MICH). District Stann Creek . P. H. Gentle JkBk, 12 Jan.

19^1. Big Eddy Creek (GH. MICH). District Toledo . A. P.

A. Lamb 57, 19^5. in undergrowth (P).W. A. Schlpp 1112,

18 Feb. 1933. growing In dense forest shade, Rio Grande

river, alt. 250 ft. (P, GH, MICH, MO, NY, UC). N. S.

Stevenson 98, 1 Feb. 1929 (P).

COSTA RICA. Prov. Cartage . A. Jimenez M. 1012, 6 Aug.

1963, Bosque en Plorencla, Turrlalba (P), Prov. Guana caster p. C. Standley and J. Valerlo kk306, 10-31 Jan. 1926. wet forest, vicinity of Tilaran, alt, 5OO-65O m. (US).

GUATEM/ILA. Dept. Alta Verapaz . 0. P. Cook and C. B. Doyle

21, 15 April 1904. Sepacuite (US). 0. P. Cook and R. P.

Griggs 757t 25 April 1902, Chirujija Oxec. near Pinca

Sepacuite (US). W. R. Maxon and R. Hay 3100, 1 Jan. I905, moist forest, trail from Pangos to Sepacuite (US). H.

Pittier 267. 7 May I905, forest along Saklak River, alt.

300 m. below Secanquin (NY). J. A. Steyermark kkJl^S, 28

Peb. 19'*^2, lowland forest in valley, "pantano," 2j mi. w, of Cubilquitz, alt. 250-300 m. (P).- H. von Tuerckheim

8019, M. Mart. I9OI. Cubilquitz (GH, US). C. L. Wilson

276, 4 March 1939, Rio Chiacte, near Pinca Volcan, 1600

ft. (P). Dept. Izabal . H. Johnson II30, 25 Jan. 1921,

Jocolo (P, GH). P. C. Standley 2^879, 2 June 1922, wet thicket, vicinity of Escoba, on bay opposite Puerto Barrio^ alt. 150 m. or less (US). J. A. Steyermark 381^*6, 28 March

19^0, in woods around stream, between Bananera and '*La

Presa" in Montana del Mico, alt. ^•0-300 m. (P). District of Peten . C. L. Lundell 2903* 1*^ April 1933, Santa Teresa.

Subin River (P, GH, MICH, US). J. A. Steyermark ^5377.

25 March 19'<'2, forest between Pinca Yalpemech along Rio

San Diego and San Diego on Rio Cancuen alt. 5O-I5O m.

(P, MO, UC). J. A. Steyermark 46084, 30 April 1942. Cerro

Ceibal, between mouth of Rio Santa Monica and mouth of Rio

San Martin, on left side of Rio Cancuen (w. side of river going downstream), alt. 75-150 m. (MICH). Map 21. Distribution of Hellconla aurantlaca.

HONDURAS. Dept. Atlantlda . P. J. Dyer A29. 8 March I916, beside trail In foothills s. of Celba (US). A. Molina H. lOkSj, 19 Marzo I962, en el bosque lluvloso y denso de

Montana Lancet 111a, Cerca de El Portlllo 3 kms. al sur de Lancetllla, alt. 5OO m. (P). P. C. Standley 56717.

6 Dec. 1927 - 20 March 1928, wet forest, Lancetllla Valley, near Tela, alt. 20-600 m. (P, US). L. 0. Williams and A.

Molina H. 130b6, 26 April l9^7t along stream In forest near

Lancetllla, alt. 200 m. (F). P. Wilson 350. 7 Feb. I903, forest near Bolets Plantation (NY).

MEXICO. State Chiapas . E. Seler 5^97 (^18). 2f March

1911. Im Walde, Rulnas de Palenque (GH). State Tabasco .

E. Matuda 33^5. 7-11 June 1939. In virgin forest S. Isldro.

Balancan (P. GH, MICH, NY).

NICARAGUA. Dept. Comarca de El Cabo . A. Molina R. I5O65.

22 Aug. 1965. Matorrales y bosque secundarlo lluvloso de

San Mateo, 16 mlllos al sur de Tronquera cerca de Hlo Wawa, alt. 35 m. (P). Dept. Zelaya . L. E. Long 54, 3

May 1947. vicinity of El Recreo (P).

22. HELICONU LEUCOPLORA .R. R. Smith, sp. nov.

Typej Panama, Prov. Colon, vicinity of Camp Plna, alt,

25 m., 11 July 1946. P. H. Allen 3590 (US). Isotypes

(P, NY. UC).

Slender plants 2-5 m. tall, cannold habit. Leaf-blade lanceolate-elliptic. 17-29 cm. long. 5-7 cm. wide, long acuminate apex, obtuse base; upper and lower surfaces greeni petiole nearly absent or leaf-blade sessile* Inflorescence erect, 10-15 cm. long, glabrous; peduncle short. If present

0.3 cm. diameter; rachls slightly flexuose, 0.2 cm. diam- eter. Branch-bracts ^^-5* orange, glabrous, lanceolate, shallow boat-shaped; lower branch-bracts 9 cm. long; upper branch- bracts 4.5-7.0 cm. long; all branch-bracts 0.6 cm. side-width; Intemode between branch-bracts about 1 cm*

Floral-bracts few, deciduous, lanceolate, membraneous,

cm. long, about O.5 cm. wide, glabrous. Flowers ^-Q, white to cream, glabrous, 7 cm. long; perianth 5*5-^*0 cm. long; pedicel 1.0-1. 5 cm. long. Fruit blue, subglobose to 3-6lde(X

O.ti-l.O cm. diameter. Seeds not seen. (Plate 22.)

Edge of moist forests and along river margins, low altitudes. Found In Costa Rica, Panama, and Nicaragua.

(Map 22.)

Lane In an unpublished study in 1957 annotated

herbarium sheets H. paulll-allenll , noting that this Is

a new species. This name could readily be confused with

species H. paulll and H. allenll . Since the naune was not

validly published, the present author, is calling the same

entity H. leucoflora , because of the notable length of the white flowers.

This species has long been Included with H. aurantlaca.

since It also possesses the orange bracts. The flowers in

H. aurantlaca are cream to yellow colored. In H. 190

ri.ANIs OK C'^-.TA RICA

' T '-a-t, /V.,:. /

PIATE 22. HELICONIA LEUCOFLORA R. R. Smith

H, leucoflora R. R. Smith, Golfo Dulce and Rio Terraba, Prov. Puntarenas, Costa Rica, Dec. 194?. A. P. Skutch 5397 (US). 19i

leucoflora the flowers are white, and then change to a

cream color when they mature. As was pointed out under

the discussion of H. aurantlaca . the flowers of H. leuco-

flora are nearly 2 cm. longer than those of H. aurantlaca .

Representative specimens J

COSTA RICA. ProY. Alajuela . A. Molina R., L. 0. Williams,

W. Burger. B. Wallenta 1757^. 20 Fe'b. 1966, lowland rain

forest between Los Chiles and Venecia, Llanura de San

Carlos, alt. 100 m. (P). Prov. Puntarenas . A. P. Slcutch 5397t Dec. 19^7, edge of forest between Golfo Dulce and

Rio Terraba, alt. 30 m. (US). A. P. Slcutch 4761, March

l9'+0, in forest, basin of El General, alt. 675-900 m. (GH, NY).

NICARAGUA. Dept. Zelaya . L. E. Long 59, 12 Aug. 19'4'7,

open bush, vicinity of El Recreo (P).

PANAMA. Canal Zone . I. M. Johnston 1625, 8 Nov. 1955,

n.w. part of Canal Zone, area w. of Llmon Bay, Gatun Locks

and Gatun Lake (GH). H. Plttler 42b5, Aug. I9II, forests around Puerto Obaldla, San Bias coast, alt. O-50 m. (GH). Prov. Code. P. H. Allen 2692, 31 Aug. 1941, La Mesa (GH, MO). Prov. Colon. P. H. Allen 3590, 11 July 1946, vicin- ity of Camp Plna, alt. m. 25 (P). Prov. Darlen . J. A. Duke 5209, 18 July 1962. Rio Chlco across from Boca de Tcsca along the top of a ridge (MO). J. A. Duke 5304, 11 Aug. 1962. ascent of Cerro Plrre from Rio Plrro b» of Map 22. Distribution of Hellconla leuooflora. 193 El Real. 600-750 m. (MO). J. A. Duke 5^03. 3 Aug. I962. along Pan Am Highway between Pucro and Rio Punusa (MO).

M. E. Terry and R. A. Terry IkZJ^ 9 March 19'*0, Cana

- Cuasl Trail (Camp 2) Cheplgana district, alt, 2000 ft.

(F). Prov. Panama . K. E. Blum, R. K. Godfrey and E. Tyson

1843, 13 Nov, 1965. 0 7 ml. n. Cerro Azu on road to Cerro

Jefe, El. @ 2600 ft. (FSU). J. H. Klrkbrlde, Jr. and

Sister Hayden 297t 23 Aug. I967. on trail to Cerro

Campana (MO).

23. HELICONIA HIRSUTA Linnaeus f., Suppl, Syst. Veg.

158. 1781. Blhal hlrsuta (L. f . ) 0. Ktze., Rev. Gen. PI.

2 I 68k, I89I. Typex "Habitat In America merldlonall.

Mutls." (not seen).

Hell con la stramlnea (Griggs) Standley, Jour. Wash.

Acad. Scl. 171 162. 1927. Blhal stramlnea

Griggs, Bull, Torr. Bot. Club kZt 327. I915.

Type I Panama, Prov, Panama, near Alhajulla. 11

Jan. 1911, H. Plttler 2328 (US).

Slender plant, 1-3 m. tallj cannold habit. Leaf-blade oblong ovate to ovate-elllptlcal, 15-35 cm. long, 5-10 cm. wide, acuminate to long acuminate apex, rounded base; upper and lower surface green; sessile to subsesslle (O.5-O.6 cm. petiole length). Inflorescence erect, nearly glabrous to pubescent, 6-9 cm. long; peduncle 0.2-0.3 cm. thlokj rachls moderately flexuose, glabrous to hirsute, 0.2-0.3 cm. diameter. Branch-bracts 4-9t dark, brick-red, or straw -yellow with greenish tinge, smooth to puberulent arotmd base, lanceolate, shallow boat-shapedt lowest branch-bract

6-11 cm. long, next to lowest 6.0-7,5 cm. long; uppermost branch-bracts 2.5-if.O cm. long; all branch-bracts 0.5-0.6

cm. side-width; Intemode between lower branch-bracts about

1.0-1.5 cm. Ploral-bracts few, deciduous, about 1.5-1.8 cm. long, lacking on many herbarium specimens. Flowers

^-12 per branch-bract, yellow with green tips or brick-red; flower about 3. 5-1^.5 cm. long; perianth 2.2-3.0 cm. long; pedicel O.if-l.e cm. long, slightly pubescence to hirsute.

Prult dark blue, subglobose, 0.6-0.8 cm. dlambtcr dried.

Seed with sclerlfled, rough, slightly wrinkled seed coat;

0.8 cm. long, 0.6 cm. wide.

In moist thickets and woods, along banks of rivers, and edges of forest swamps.

H. hlrsuta Is more common In South America. It extends up Into Panama with a distinct variety found further north In British Honduras and Nicaragua.

This species Is not easily confused with other species of Hellconla since It Is of smaller stature with a small pubescent Inflorescence. H. hlrsuta var. hlrsuta has a very pubescent perianth, while var. rublflora possesses a slightly to moderately pubescent perianth. The two varieties are extremely similar In habit, and occupy nearly the same habitat. The first H. hlrsuta to be described was red-bracted.

Griggs (I915) reported that Plttler had found a specimen with straw-yellow bracts. To this specimen, he gave a new name B. stramlnea * However, only the color Is differ-

ent from the described H. hlrsuta . The present conclusion

Is that both colored bracts belong under the same variety,

H. hlrsuta var. hlrsuta .

There Is an Indication the H. hlrsuta var. hlrsuta hybridizes with H. vaginalis . This Is discussed In a later chapter.

Key to varieties of Hellconla hlrsuta

Plant 1-3 m. tall; perianth yellow with green tip, hirsute var. hlrsuta

Plant approximately 1 m. tall; perianth brick-red, glabrous or nearly so var. rublflora

Hellconla hlrsuta L. f . var. hlrsuta

Plant 1-3 m. tall. Leaf-blades 19-35 cm. long, 6.0

-9.5 cm. wide. Branch-bracts 5-9, red to yellow; lowest branch-bract 9. 5-1 1.0 cm. long; uppermost branch-bracts

3-** cm. long; all branch-bracts O.5-O.6 cm. side-width.

Flowers 4-10 per branch-bract, hirsute 3. 5-'*. 5 m. long; perianth 2.5-2.8 cm. long, pale yellow with green tips; calyx hirsute, corolla glabrous; pedicel 1.0-1.8 cm. long, hlrsute-tomentose with whitish to yellow-brown hairs. (Plate 23.) 197

PIATE 23. HELICONIA HIRSUTA L. f. var. HIRSUTA

Blhal stramlnea Griggs, Corozal, Canal Zone, Panama, 18 Dec. 1923. P. C. Standley 2731^5 (US). t .

198

Primarily In moist thickets and edges of forests.

In Central America, this variety Is found In Panama*

Its range extends throiighout northern South America*

(Map 23.)

Representative specimens

PANAMA. Canal Zone , P. H. Allen 3633. 8 Aug. 19'*^6. vicin- ity of Madden Dam, alt. 25 m. (F). K. E. Blum 1076, 5 Oct*

1965, vicinity of TTC Albrook Tower (PSU). D. S. Correll

12263. 31 Aug. 19^5. high hills In Jungle forest, about

2 ml. w. of Balboa (UC). C. W. Dodge and P. H. Allen

17343. 9 Jan. 1935. forest along banks of Quebrada La Palma and Canon of R. Chagres, 7O-8O m. (MO, UC, US). E. P*

Kllllp 39953. 22 Nov. 19'^8, Ancon Hill, open summit (US).

H. Plttler 2200, 3 Jan. I9II. between Mlraflores and

Corozal. alt. 20-30 m. (US). P. C. Standley 27133. Nov.

1923 - Jan. 1924, Balboa (US). P. White 240, 25 Aug.

1938, vicinity of Mlraflores Lake (MO). W. R. Stlmson

53^5. 12 Aug. 1967. edge of clearing near road. Navy

Pipeline Reserve Area ca. 3 ml. n.w. of Gamboa . ( DUKE )

E. Tyson 1149. 16 Aug. 19^5. Mlraflores Looks area (PSU).

R. E. Woodson, Jr., P. H. Allen, R. J. Selbert I3I6, 20

July 1938, western slope of Ancon Hill (MO). Prov.

Chlrlqul . P. H. Allen 3668, 29 Aug. 1946, vicinity of

Remedies, alt. O-I5O m. (P). M. E. Davidson 1175. 18 Aug.

1938, Puerto Armuelles, alt. 50 ft. (P). Prov. Darlen. 199

E. Tyson, J. Dwyer, K. Blum, and J. Duke ^+651, 15 July

1966, 1 ml. e. of Santa Fe (FSU). Prov. Panama * P. H.

Allen 2961, 7 Sept. 19*^2, Vacamonte Pt. (MICH, MO, NY,

US). P. H. Allen 36IO, 1 Aug. 19^6, vicinity of Vacamonte

Pt. (F, NY). C. W. Dodge, A. A. Hunter, J. A. Steyermark and P. H. Allen I6654, 29 Nov. 193'+. along road between

Panama and Chepo (MO). W. R. Maxon, A. D. Harvey 67^0,

1-3 June 1923. near Tapla River, Juan Diaz region (US).

P. C. Standley 28O85, 7 Dec. 1923 - 11 Jan. 1924, moist thicket, Rio Tapla (US). P. C. Standley 3059O, 11 Jan.

192'i-, moist woods, Juan Diaz (US). R. E. Woodson, Jr.,

P. H. Allen, R. J. Selbert I355, 21 July 1938, thickets and forests near Arraljan, ca. I5 m. (MO, NY).

Hell con la hlrsuta L. f . var. rubl flora R. R. Smith, var. nov.

Type: British Honduras. Dlst. Stann Creek, swampy places,

Stann Creek, alt. 20 ft., I9 Sept. I967. W. A. Schlpp 357

(MO). Isotypes (F, MICH, NY, UC).

Plant about 1 m. tall. Leaf-blades 15-25 cm. long,

5- 6 cm. wide. Branch-bracts 4-5, red; lowest branch-bracts

6- 11 cm. long, uppermost branch-bracts 2.5-4.0 cm. long; all branch-bracts 0.4-0.7 side-width. Flowers 4-8 per branch-bract, red, 3.5-4.0 cm. long; perianth 2.5 cm. long, puberulent to slightly hirsute with short whitish hairs.

Ecology similar to var. hlrsuta. :5

Map 23 • Distribution of Hellconla hlrauta*

t

202

This Variety Is apparently quite rare, and further collections are necessary to understand It fully. Perhaps

It will merit the species level upon future consideration*

Representative specimens

BRITISH HONDURAS. Dlst. Stann Creek . W. A. Schlpp 835.

6 Dec. 1931* growing In open flats, Stann Creek (F, MICH,

MO. NY. UC).

NICARAGUA. Prov. Comarca de El Cabo (?). P. E. Schramm

40, 1926, Sangsangta, Segovia district (US). F. £• Schramm

57. 1927. Sangsangta district (US). Dept. Zelaya . A*

Molina R. 2127. 10 April 19^9. brenales espesos de la

Esperanza, Rio Grande, Guamll o brenas sabre areas pantanosas a lo largo del Rio Grande, alt. O-I5 m. (F).

2k. HELICONU VAGINALIS Bentham, Bot. Voy. Sulph.

171, 184if. Type: Isle of Gorgona (off the coast of

Colombia) (K).

Slender plant, 1^-31 m. tall, cannold habit. Leaf

-blades elliptical to oblong, 32-55 cm. long, 9-1^* cm, wide, cuspidate-acuminate at tip, obtusely to broadly acute at base; upper surface dark green, lower surface paler green; sessile or nearly so. Inflorescence erect or slightly nodding, glabrous or occasionally slightly pubescent* 12-14 cm. long; peduncle variable In length.

0.4-0.5 cm. diameter; rachls moderately flexuose, 0.3-0,4 oa. diameter. Branch-bracts 2-6, deep red, lanceolate. usually with thin membraneous margin, shallow boat-shaped; lowest bracts nearly horizontal or slightly curved upward,

12-17 cm. long or If expanded Into leaf-blade 17-37 cm. long; upper branch-bracts 5 •5-9.0 cm. long; all branch

-bracts about 1 cm. side-width; Intemode of lower branch

-bracts on rachls 2-3 cm. Floral-bracts lanceolate, glabrous, about 3 cm. long. Flowers per branch-bract, yellow and often greenish at tip, 5-6 cm. long; perianth

4-5 cm. long, glabrous, pedicel 1.0-1,5 cm. long, sometimes slightly hirsute. Mature fruit bluish purple, somewhat globular, 1 cm. diameter when dried. Seed about 0.6 cm. wide, 1 cm. long. (Plate 2^.)

Frequent component of moist to wet forests, euid swampy at altitudes up to about 975 m.

The distribution for H. vaginalis Is from the southern states of Mexico to Panama. Its range also extends Into

Colombia and . (Map 2^*.)

H. vaginalis Is In a group of closely related species for which there have been a number of Interpretations.

The following list Is offered to Illustrate the confusion.

1) Baker (1893) recognized H. vaginalis as a variety of

H. hirsuta. In this grouping also was Included another

species, H. swartzlana .

2) Griggs (1903) called the entity In question H. choconlana . which Is undoubtedly a synonym of H. aurantlaca. PIATE 2k, HELICONIA VAGINALIS Benth.

H. vaginalis Benth., n.w, part of Canal Zone, Panama, 9 Aug. 1955* I. M. Johnston 1579 (GH). j

3) Griggs (1915) treats the species In his key as B.

acuminata and B. hlrsuta cannoldea .

4) Standley (1928). In the "Flora of the Panama Canal Zone," Includes this taxon \mder the name H. acuminata *

5) Standley (1927). In his "Flora of Costa Rica." repeats H. acuminata for this taxon.

6) Woodson (19'*5). In the "Flora of Panama." places this

species under H. pslttacorum .

7) Standley and Steyermark (1952). In the "Flora of Guate-

mala." list this entity as H. subulata.

The confusion of this species concept Is a result of misinterpretations of the taxa. In an attempt to under-

stand these entitles, this author has listed his conclu-

sions resulting from investigation of literature and herbarium study

1) Hellconla vaginalis Is the glabrous or nearly so. red

branch- bracted species with yellow flowers. It Is

cannold In habit ranging In height from 1^ meters to

3^ meters. Its range extends from Mexico to Ecuador, 2) Hellconla cannoldea L. C. Rich.. Nova Acta XV: 24,

tab. IX and X I83I. This Is a smaller species than H. vaginalis , reaching a height of approximately one

meter. Its range extends from Peru to French Guiana and Brazil. Synonyms for H. cannoldea are as follows 1 t

Hellconla rlchardlana Mlq, In Linn. XVIIIi 70. IBkk,

Type J Surinam.

Hellconla blcolor Klotzseh In Linn. XX» 465. 1847.

Type I Guiana.

Hellconla hlrauta L. var. cannoldea (L. C. Rich.)

Baker, Ann. Bot. 7i 197. 1893. not validly pub-

lished, though cited as such by Woodson (1945).

3) Hellconla acuminata L. C. Rich., Nova Acta XVi tab

XI-XII. 1831. This Is a South American species with

greenish branch-bracts and red flowers* Its reuige Is

similar to that of H. cannoldea .

4) Hellconla subulata Ruiz at Pavon., Fl. Peruv. 3i 70,

pi. 303b. 1802. This Is a tall petloled Peruvian

species, more dlstsmtly related to the group.

It Is thought that this species hybridizes with other species of Hellconla . This is discussed In a later chapter.

Representative specimens

BRITISH HONDURAS. Dlst. Stann Creek . P. H. Gentle 2119,

9 Aug. 1937, base of hill, Stann Creek Hallway, I5 miles

(P, GH, MICH). W. A. Schlpp 363. 21 Sept. 1929. Middlesex, alt. 200 ft. (P. GH, MICH, NY. UC).

COSTA RIGA. Prov. Alajuela . A. M. Brenes I265O, 26 June

1930, entre Tllaran y Laguna de Arenal (NY). Prov. Cartagg

P. C. Standley and J. Valerlo 47312, 7-8 Peb. 1926, wet forest, vicinity of Pejlvalle, alt. ca. 900 m. (US). Ad.

Tonduz 11385, XI I897. Forest de Tuls, alt. 65 m. (MO).

ProY. Guana caste . P. C. Standley, and J. Valerlo k^lSk,

18-19 Jan. 1926. wet forest. El Arenal, alt. ^85-600 m.

(US). Prov. Heredia . W. Burger and G. Matta U. ^^163,

5 and 6 Jan. 1967» original wet forest and Cocoa plantation

on the property of Dr. L. Holdridge, ca. 2 km. upstream

on the Rio Puerto Viejo, alt. + 100 m. (P). Prov. Limon «

C. W. Dodge and V. P. Goerger 928?, 25 July 1936, forest

near farmhouse at Finca Castilla, 30 m. (F, GH, MICH).

A. Jimenez 2097, 17 Julio 196^. Una« 2.5 km. al Este de

Guapiles, alt. 250 m. (P). J. D. Smith '+97'*, M. April

I894, Jimenez, Llanos de Santa Clara, Comarca de Llmon,

alt. 650 m. (US). P. C. Standley 37330, 12-13 March, 192'*.

wet forest, vicinity of Guapiles, alt. 3OO-5OO m. (US).

P. C. Stemdley, J. Valerlo ^18980, I8-I9 Feb. 1926, Plnca

Monte Cristo, on the Rio Reventazon below Cairo, alt. 25

(US). Prov. Puntarenas . P. H. Allen 5486, 16 March

1950, locally common in wet forest, vicinity Tinoco

Station: Area between Rio Esquinas and Palmar Sur de Osa, 100 ft. (UC). Prov. San Jose . A. P. Skutch 476I, March

19'*0, in forest, basin of El General, alt. 675-9OO m. (US). GUATEMAIA. Dept. Alta Verapaz . 0. P. Cook and R. P.

Griggs 693, 22 April 1902. near Pinca Sepacuite Chirujinja

Oxec. (US). J, A. Steyermark 'f'+876. 10 March 1942, between

Cubilquitz and Yakapur, alt. 3OO-37O m. (P). J. A. ,

20o

Steyennark k^ZZS, 23 March 19^2, woods s.e. of Flnca

Yalpemech, near Alta Verapaz-Peten boundary line, alt.

100-150 m. (P). H. von Tuerokhelm 8317. M. July I9O2. Cubllqultz. alt. 35O m. (GH, NY, US). Dept. Chlmaltenango .

Mrs. B. B. Lewis 270. 11 Jan. 1936 (P). Dept. Huehuete-

nango. J. A. Steyermark 49237t 23 July 19^*2. In woods

between Ixcan and Rio Ixcan, Sierra de los Cuchumatanes alt. 150-200 m. (P). Dept. Izabal . W. A. Kellerman 6I97.

17 Jan. 1905, Livingston (P). J. A. Steyennark 38I33, 28

March 19^0, between Bananera and "La Presa" In Montana del

Mlco, alt. tfO-300 m. (P). J. A. Steyermark 3870^^, April

19^0, between Virginia and Lago Izabal, Montana del Mlco,

alt. 50-100 m. (P). J, A. Steyermark iH532, 18 Dec. 19/*1,

openings on slope, along Rio Prlo and tributaries, alt. m. 75-150 (P. MO). Dept. unknown . S. Watson I90, I885.

eastern portions of Verapaz and Chlqulmula (GH). HONDURAS. Dept. Atlantlda . P. C. Standley 7922, 22-27 April 19'*7. dense wet forest, vicinity of San Ale Jo, I50

-270 m., at base of hills s. of San Ale Jo near Hlo San Alejo (P). T. G. Yuncker, J. M. Koepper and K. A. Wagner July 8697, 30 1938, along Danto River, lower slopes of Mt.

Cangrejal, vicinity of La Celba (P, GH, NY). ^ICO. State Chiapas . E. Matuda 3639, 6-9 July I939, In advanced forest. Javallnero, Palenque (MICH). State Tabasco. C. L. Gllly. Sr. and E. Hernandez X. 301, 20 Sept. I9kk, Mun. Teapai rain forest on ridge Just n. of Cerro de Azufre (MICH). NICARAGUA. Dept. Matagalpa . L. 0. Williams, A. Molina R,,

T. P. Williams 23823. 14-16 Jan. I963. growing In forest,

moist, dense rain forest oa. 6-10 km. n.e. of Matagalpa,

road to El Tuma, alt. 1000 m. (P). Dept. Zelaya . P. C.

Engleslng 25^, 1928, region of Braggman's Bluff (P, US).

L. E. Long 65, 7 Aug. 19'*'?. Cukea, near sea level, vicin-

ity of El Rlcreo (P). P. J. Shank and A. Molina R. 4728.

22 Nov. 1951. bosque lluvloso y brenoso de Montana Esqul-

pulas, alt. 130 m. (P). Prov. unknown . B. W. Taylor ^S^ff,

31 Dec. 1963, Miguel Begem, Teoslnto savanna, alt, 200 m.

(P).

PANAMA. Prov. Bocas Del Toro . H. von Wedel 600, 6 Sept.

19^0, Water Valley (GH, MO). H. von Wedel 1023, 2 Oct.

19'*0. vicinity of Chlriqul Lagoon (MO). R. E. Woodson,

Jr., P. H. Allen, R. J. Selbert 1931. 12-16 Aug. I938,

Rio Crlcamola, between Pinca St. Louis and Konklntoe, ca.

10-15 m. (MO). Canal Zone . A. M. Chickering 2, 12 Aug.

1928, common on Barro Colorado Island (MICH). J. P. Cowell

3^0, 7 March I905, Matachul to Las Cascadas (NY). C. W.

Dodge, J. A. Steyermark and P. H. Allen 16993. 16-21 Dec.

1934. drowned forest of Quebrada Ancha and Rio Salamonca, m. 70 (MICH. MO, UC, US). J. A. Duke 4260. 6 Oct. I96I, near Coco Solo Weather Station (MO). J. E. Eblnger 219. June 28 I96O, Barro Colorado Island, along shore, e. of dock (MO). A. Pendler 443. Jan. I95O, Chagres, Isthmus of Panama (GH, MO, US). I. M. Johnston I579, 9 Aug. I955, north base of Hindi Hills, catlval along Quelrada Morlto

(GH). L. A. Kenoyer 229. July 1927, Barro Colorado Island

(US). E. P. Klllys 12170, lb Oct. 1922, shore of Gatun

Lake, between Prljoles nnd Monte Llrlo, alt. 30 m. (US).

R. J. Selbert 1519. 9 Aug. 19^+0, Coloni Gatun Lake, Good- year, all weather Estate (MO). P. C. Standley 271b2, 17

Dec. 1923, wet forest, hills w. of canal, near Gatun (US).

P. C. Standley 27^92, I9 Dec. I923. wet forest, hills n. of Prljoles (US). P. C. Standley 2B669, 28 Dec. I923, swampy woods, near Port Randolph (US). P. C. Standley

30I8I, 9 Jan. 1924, wooded swamp, between Prance Pleld,

Canal Zone and Catlval (US). W. R. Stlmson 53^+4, 12 Aug.

1967. In moist forest on ridge. Navy Pipeline Reserve Area ca. 3 ml. n.w. of Gamboa (DUKE). E. L. Tyson and K. Blum

3937. 6 May 1966, 12 ml. s. Colon on Rio Providencla (MO).

R. E. Woodson, Jr. and R. W. Schery 990, 3-20 Aug. 19^10, Barro Colorado Island (MO). Prov. Code . P. H. Allen

3599. 20 July I946, region north of El Valle de Anton, alt. 1000 m. (GH). Prov. Colon . P. H. Allen 3585, 11 July 1946, vicinity of Camp Plna, alt. 25 m. (P, NY. US).

Prov. Darlen. H. Plttler 4404. Aug. I9II, forests around Puerto Obaldla, San Bias coast, alt. O-50 m. (NY, US). Prov. Panama. P. H. Allen 2216. 1 Sept. 1940, Summit of Cerro Campana, 800-1000 m. (US). Prov. unknown . P. H. Elmore L17. 3 April 1939. Caladonla Harbor, Mt. Vernon (P, MICH, UC, US). Map 24, Distribution of Hellconla vaginalis

213

25. HELICONU TRINIDATIS Lane ex R. R, Smith. Typei

WEST INDIES. TRINIDAD. Forests at Cumuto. 26 Nov. 1926,

W. E. Broadway 6421 (P). Isotype (MO).

Slender plant, 1-2 m. tall, cannold habit. Leaf-blade

ovate-lanceolate, cm. long. 7-13 cm. wide, acuminate

apex, rounded base; upper and lower surfaces green; petiole

2.5 cm. long to nearly sessile. Inflorescence erect or

nodding, slightly pubescent, 7-12 cm. long; pedxmcle vari-

able, 23 cm. long to subsessile, short tomentose; rachls flexuose. 0.2 cm. diameter, hirsute-short tomentose.

Eranch-bracts 5-6, red, orange at base and red distally.

slightly pubescent along underside and near base, lanceo-

late, shallow boat-shaped, moderately curved upward; lowest branch-bract 8-11 cm. long; middle and upper branch-bracts

4-7 cm. long; all branch-bracts 0.6-0.8 cm. side-width;

intemode between branch-bracts 1.0-1.4 cm. Floral-bracts

few. lanceolate, 4.5-5.5 cm. long. 0.6-0.7 cm. wide, some-

what slightly pilose. Flowers 5-30 per branch-bract,

yellow-orange, greenish distally, 5.5 cm. long; perianth cm. 3.5-5.0 long, nearly glabrous; pedicel 0.6-1.2 cm. long, hirsute. Fruit dark, subglobose to 3-sided, 0,6 cm. diameter, 0.7 cm. long, glabrous. (Plate 25.) In moist forests of . (Map 25.) This species was first recognized by Lane in his unpublished study. It is in the group of plants which have red branch- bracts, yellow flowers, and possess a 2U

k

PIATE 25, HELICONIA TRINIDATIS Lane. ex R. R. Smtth

Isotypej H. trlnldatls Lane ex R, R. Smith. Cumuto, Trinidad, , West Indies, 26 Nov. 1926, .W. E. Broai^way (MO), .

?15

cannold habit. In Middle America three species are found In this group, H. vaginalis . H. svartzlana . and H. trlnl-

dfttls differs from the other two species In the following wayst

1. It Is generally a shorter plemt,

2. The leaves often possess a short petiole.

3« The flowers have dark green apices.

4. The pedicel Is generally shorter.

5. It produces a smaller seed than that found In

H. vaginalis .

Often the secondary rachls to which the flowers are

attached Is elongated within the branch-bract. This gives

the appearance of pedicels extending upward from about the middle to the axis of the branch- bract

Representative specimens!

WEST INDIES. N. L. Brltton and T. E. Hazen 366. 8-9 March 1920. Mora Forest, e. of Sangre Grande (GH. NY). W. E. Broadway s. n. 5 Sept. I911, Emperor (Morrison) Valley, back of Lookout (GH. NY). W. E. Broadway 698O. 25 May 1928. gully, near 1 3A ml, post. Canra road (P. MO). E. A. Friend 28. 29 Aug. I939. Summit of Mt. Tucutche. alt. ft. 3072 (NY). R. A. Howard 10338. 9-23 Feb. I95O. In rain forest. Arlpo Savannah on Waller Air Force Base (GH). L. Rlly 62. May 26, hills behind Port of Spain (NY). Map 25. Distribution of Heliconia trlnidatls .

t)

p. Ross 1052, 26 Jan. 1958, Mount Harris, alt. 200 ft.

(GH). A. C. Smith IOO62. 10 March I956, Arlma Valley

Santa Isabella trail, near Simla, alt. 250-350 m., forest

(GH. NY, UC). TOEAGO. N. Y. Sandwlth 1732, 11 Oct. 1937,

In deep shade In forest (NY).

26. HELICONIA SWARTZIANA Roemer and Schultes. Syst.

Veg. Vi 591. I8I9. Type J Curtis. In Jamalcae sylvls montosls caedals. The type Is an Illustration In Curtis 's

Bot. Mag. t, 502. 1801.

Hell con la pslttaborum Swartz, In Curt. Bot. Mag. t.

502. 1801. (Non Hellconla pslttacorum L. f . Type: same as above.

Hellconla harrlslana (Griggs) L. B. Smith, Contr.

Gray Herb. 12^4-: 6. 1939. Blhal harrlslana

Griggs, Bull. Torr. Bot. Club 42: 328. I9I5.

Type: West Indies. Jamaica, Lelcesterf leld.

Upper Clarendon, alt. 1800 ft., 28 Feb. I9IO,

W. Harris 10841 (NY). Isotype (F. US).

Slender plant, 3-5^ m; cannold habit. Leaf- blade oblong-ovate to broadly elliptic. 23-30 cm. long, 10-11 cm. vrlde, long acuminate apex, rounded base; leaf-blade sessile to sheath, no petiole. Inflorescence erect, glabrous, 13-17 cm. long; peduncle variable; rachls flexuose. puberulent-tomentose; 3 cm. diameter. Branch-bracts 5-7, orange-red or bright crimson above and

greenish-yellow below, narrowly lanceolate, shallow boat

-shaped, horizontal to moderately curved upward; lowest

branch-bract 10 cm. long; middle and upper branch-bract

5.5-8.0 cm. long; all branch-bracts 0.5-0. 7 cm. side-width.

Floral-bracts few, deciduous, slightly pubescent, 2.8 cm.

long, 0.6 cm. wide. Flowers 5-10 per branch-bract, yellow

-gold, fading to black tip. If. 5 cm. long; perianth 2.5-3.^f

cm. long, lower half of calyx slightly puberulent, corolla

glabrous; pedicel 1.0-1.5 cm. long, tomentose. Fruit dark,

0.5 cm. diameter. Seeds not seen. (Plate 26.)

At edge of rich woods at altitudes of 1600 to 2300

feet. Endemic to Jamaica. (Map 26.)

H. swartzlana Is closely related to H. trlnldatls and H. vaginalis . These three species have slender cannold habits, red branch-bracts and yellow flowers. H. swartzl- ana differs from H. trlnldatls and H. vaginalis by the following characteristics!

1) H. swartzlana becomes much taller than the two

species forementloned, up to 5j m. tall. The 2) perianth of H. swartzlana Is 3 cm. or less.

The perianth of H. trlnldatls and H. vaginalis

Is longer than 3 cm. The 3) leaf-blades of H. swartzlana are more rounded, with a wlde-elllptlcal shape. The leaf-blades of H. vaginalis and H. trlnldatls are narrowly elllptio. PIATE 26, HELICONIA SWARTZIANA Roem.and Schult.

Holotype: glhal harrlslana :;rlggs. Leicesterfleld. . upper Clarendon. Jamaica, 2^5 Feb. I910, W91. Harris 10. 84l(NY). Map 26, Distribution of Hellconla sxartzlana . 222 )

^) The lowest branch-bract of H. swart zlana la fer-

tile and never extends Into a leaf-blade. In H.

vaginalis the lowest branch-bract often extends Into a leaf-blade.

Representative specimens}

JAMAICA.. Clarendon Parish. M. H. Crosby and W. R. Anderson 1232. 19 Aug. I963. Peckham Woods, near Aenon

Town. ca. 2200 ft. (MICH). Wm. Harris IIO38. 21 May

1912, Public Gardens, In partial shade In the outskirts

of Woods, stem up to 16 ft. high, alt. 2200-2500 ft.

(P. NY).

Series Dlstantes Griggs, Bull. Torr. Bot. Club kZt

316. 1915. (Rank not specified; here Interpreted as series.

Moderately stout plants 1-4 m. tall; musold habit. Leaf- blades over 50 cm. long; somewhat stout petiole, over

4 cm. long. Inflorescence over I5 cm. long, erect or nodding; glabrous or with varying degrees of pubescence; rachls flexuose or straight, sometimes twisted. Branch -bracts if-22. variable In size, shape and color.

Type species I

Hellconla latlspatha Benth. This series Is represented by 12 species from Central America. Two. and possibly a third, species are found In South America. Aside from the musold habit, the branch

-bracts of the species In this series are generally more

deeply boat-shaped than those In the Series Stenochlamys.

Key to the Species of Series Dlstantes

A. Branch-bracts nimber 12-22} all branch-bracts close

together and strongly curved upward; rachls stiffly erect*

B. Branch-bract yellow; perianth less than 2 cm.

long 27. H. librata

B. Branch-bracts red to dark yellow; perianth more

than 2 cm. long.

C. Plant height 3 m.; lower branch-bracts 18-21

cm. long, cherry red to dark yellow; perianth

^.5 cm. long 28. H. lankesterl

C. Plant height 1.5-2.5 m.; lower branch-bracts

18-35 cm. long; red; .perianth length less

than k,5 cm.

D. Inflorescence extremely pubescent;

rachls straight, woolly with light

tan hairs; branch-bracts highly

villous on sides... 29. H. mollanaeana

D. Inflorescence slightly pubescent; rachls

slightly flexuose, puberulent to tomen-

tose; branch-bracts slightly pubescent

on sides 30.* h. adflexa Branch-bracts niunber ^f-ll; lower branch-bracts widely spaced, extended out horizontally or reflexed; rachis erect or sometimes nodding.

£• Petioles of leaf-blades l|-7 cm. longi Inflores-

cence length up to 25 cm.

P. Perianth bright red to pink, glabrous; floral

-bracts ^ cm. long; underside of leaf usually

tinted purple 31. H. metalllca

P. Perianth yellow, slightly pubescent j floral

-bracts 3*0-3.5 cm. longj underside of leaf

usually green, sometimes tinted purple......

32. H. osaensls

E. Petioles over 10 cm. long, extended; Inflorescence

over 25 cm. long (occasionally less).

G. Branch-bracts 3-5; red with green

margin; rachls slightly flexuose;

Inflorescence with distichous bre^ich

-bracts 33, h. llnnaeana

G. Branch-bracts ^-12; red to yellow;

rachls slightly to moderately flexuose;

Inflorescence with twisted rachls so branch-bracts spiral.

H. Branch-bracts narrow, linear

-lanceolate to oblong lanceolate; red. 226

I. Lower fertile branch-bracts

7-10 cm, long and 0.6-0.8 cm.

side-width; floral-bracts

ovate and extended out from

branch-bracts; branch-bract usually reflexed

3^. H. schledeana

I. Lower fertile branch-bracts

11-16 cm. long and 0.7-1,0

cm. side-width; floral-bracts

4 cm, long, remaining within

branch-bract; lower brsmch

-bracts extend out horizon-

tally from rachls

35* H. veracruzll

H. Branch-bracts wide, ovate-lanceolate

with subcordate base to subovate

-triangular; red to yellow.

J • Hachls glabrous or nearly

80; perianth 3.0-4.0 cm. long

36. H. latlspatha

J. Rachls tomentose; peri-

anth 'f.0-5.2 cm. long. •

227

K. Branch-bracts scar-

let, glabrous or

tomentose only along

lower surface; side

-width of branch

-bracts 2.4-3.0 cm.;

apices of branch

-bracts extending

straight out

37. H. tortuosa

K. Branch-bracts red

or yellow with red

margin, tomentose;

side-width of branch

• -bracts 2.0-2.2 cm.;

apices of branch

-bracts clrclnate . .

38. H. irrasa

27. HELICONIA LIBRATA Griggs, Bull. Torr. Bot. Club

30t 1903. Blhal IJJbrata (Griggs) Griggs, Bull,

Torr. Bot. Club 31: >'t5. 190^+. Typei Guatemala, Cherujlja, Oxec, 0. F. Cook and R. P. Griggs 696 (US).

I 228

Moderately stout plant, 1. 5-3.0 m. tall? musold habit.

Leaf-blade ovate-oblong to elllptlc-oblong, 64-115 cm.

long, 20-25 cm. wide, slightly acuminate to acute apex,

obtuse (occasionally with unequal sides) base; upper

surface glabrous, lower surface glabrous or glaucous;

petiole long, up to 90 cm. Inflorescence erect, glabrous

except short tomentose on rachls; triangular, 20-22 cm.

long; peduncled; rachls barely flexuose, 0.7-0.8 cm.

diameter. Branch-bracts 9-22, yellow, glabrous, lanceo-

late to ovate, subcordate, shallow boat-shaped, strongly

curved upward; lowest branch-bract often extended Into

leaf-blade, reaching total length of about 41-60 cm.,

sterllet lowest fertile branch-bracts 10-11 cm. long;

middle branch-bracts 4.0-6.5 cm. long, upper branch

-bracts gradating from 2.5-^^.0 cm. long; all branch

-bracts 1.2-1.5 cm. side-width; internode between branch

-bracts about 1 cm. Floral-bracts, pale-yellow, thin

membraneous, deciduous; about 2 cm. long, 0.5-0.8 cm.

wide. Flowers 7-15 per branch-bract, yellowish, 1.8

-2.5 cm. long; perianth I.5-I.6 cm. long; pedicel 1.2

-1.5 cm., extending up and out of bract, glabrous to slightly short tomentose. Fruit blue, 0.8-0. 9 cm. diameter when dried, glabrous. Seeds with sclerified. rough and wrinkled seed coat; 0.6-0.8 cm. long, 0.4-0.5 cm. wide. (Plate 27. ) / PIATE 27. HELICONIA LIBRATA Griggs Holotype: Hellconla llbrata Griggs, Chlrujija Oxec. Dept. Verapaz. Guatemala. 22 April V I 1902. 0. F. Cook and tt. r. Griggs 696 (US), i

steep heavily wooded slopes, rich moist forest.

Range extends from Mexico to Nicaragua. (Map 27.)

H. llbrata Is one of the four species with a "candel-

abrum" Inflorescence distinguished by yellow branch-bracts

and flowers which do not exceed a length of two and one

-half centimeters. The closely related species are H. adflexa, H. mollnaeana and H. lankesterl . all of which

have red or rarely dark yellow branch-bracts and flowers which are more them three centimeters long.

Representative specimens

BRITISH HONDURAS. Dlst. El Cayo . C. L. Lundell 6535,

June - August 1936, Camp 6, In low secondary growth (GH, MICH, NY).

GUATEMALA. Dept. Alta Verapaz . H. von Tuerckhelm 83I8,

M. June 1902. CubUqultz, alt. 350 m. (US). H. von Tuerck- helm II 620, August I907, Cubllqultz In Hochwald, 35O m. (GH, NY). Dept. Huehuetenanppo . J. A. Steyermark 49203, July 22 1942, around Ixcan at "Patcushln," Sierra de los Cuchumatanes, alt. 5OO m. (P). Dept. Pet en . A. Molina R. 15558, 11 Nov. 1965, Bosque tropical humedo de Yaltutu entre Crlsto y Poptum, alt. 433 m. (F). A. Molina R.

15843, 18 Nov. 1965, Matorrales y bosque humedo entre orllla Rio Paslon y Ceibal. alt. 60 m. (P). 231

HONDURAS. Dept. Atlantlda . A. M. Chlckerlng 168, 22 June

- 27 July 1929, In forest, Lancetilla (MICH). P. C.

Standley 52704, 6 Dec. 1927 - 20 March 1928, Lancetilla

Valley, near Tela, alt. 20-600 m. (F, US). T. G. Yxmcker,

J. M. Koepper, K. A. Wagner 85^16, 19 July 1938, on mountain slopes near Puento Alto stop on S. P. Co. R. R., e. of

Celba. alt. 800 ft. (F, GH, MO, NY, US). Dept. Cortes ,

P. H. Allen. M. Trafton, and A. Chable 6569. 25 Sept. 1952,

Lot "B, " Research Camp, Lake Yojoa, alt. 66O m. (GH).

T. G. Yuncker 4870 a and b, 30 July 1934, In rich moist forest near Paral, Lake Yojoa (P, MICH, MO).

MEXICO, State Chiapas . D. E. Breedlove, P. H. Raven

13557, 19 Oct. 1965, steep heavily wooded slope, 32 kilo- meters n. of Ocozocoautla along road to Mai Pasco.

Mxmlclplo of Ocozocoautla de Esplnosa (P, MICH). R. L.

Dressier 1515, 20 July - 20 Aug. 195^, limestone area, near laguna Ocotal Grande, ca. 25-30 km. s.e. of Monte

(Cerro) Llbano (which Is ca. 45 km. e. of Ocoslngo), alt. ca. 950 m. (GH). K. Roe, E. Roe and S. Mori 904, 4-5 Aug.

1965, undisturbed Tropical Evergreen forest. Rolling hills of n.w. end of Valley of Chiapas on road to Mai Paso, 41 km. (by road) n.w. of Ocozocoautla, alt. ca. 350 m. (WIS). t

Map 27 • Distribution of Hellconla llbrata .

234

I

State Tabasco . E. Matudo 3'J-65, 19-25 June 1939, In virgin forest, Retiro, Tenoslque (F, GH, MICH).

NICARAGUA. Prov. Comarca de El Cabo . A. Molina R. IU70,

13 Agosto 1965, Matorrales y bosque humedo de Sllma Sla, sabanas y plnares (Plnus carlbaea ) de la costa Atlantlca, alt. 100 m. (F).

28. HELICONIA lANKESTERI Standi ey. Jour. Wash. Acad.

Scl. I7t 162. 1927. Typei Costa Rica, Prov. Cartago, wet forest La Estrella, 26-27 March I92if, P. C. Standley

39'+9^ (US).

Stout plant, 1.5-3.0 m. tall; musold habit. Leaf

-blade oblong-elllptlc, 65-75 cm. long, 20-25 cm. wide, short acuminate apex, rounded base; upper and lower sur- faces green; long slender petiole sometimes with short tomentose pubescence. Inflorescence erect, glabrous, deltoid- shaped, 30 cm. long; pedundle sometimes short tomentose; rachls stout, often short tomentose, 1 cm. diameter, flexuose. Branch-bracts about 16, cherry-red to dark yellow, glabrous, lanceolate, long attenuate apex.

- . ] 235

curved upward from base, usually fertile, total length

35-50 cm. long; lowest branch-bract often extending Into leaf- blade 5 next lowest branch-bract lb-2l cm. long; middle

branch-bracts about 10-12 cm. long; uppermost 5-6 cm. longt all branch-bracts about 2.5 cm. side-width j intemode

between lower branch-bracts 1-3 cm., gradually becoming

shorter. Floral-bracts wide-lanceolate, 3.5 cm. long,

0.6 cm. wide. Flowers ^-6 per branch-bract, red to dark yellow, 4.5-5.0 cm. long; perianth 4.5 cm. long, glabrous; pedicel about 0. 5-1.0 cm. long. Fruit dark, almost ex-

serted from bracts, subglobose to 3-sided, 1 cm. diameter. (Plate 28.)

In moist rain forests, mainly of high altitudes, 2000

-7000 ft. Restricted to Costa Rica and western Panama. (Map 28.)

As stated in the discussion of H. librata, H. lankes-

^gri is one of four species with a "candelabrum" type of inflorescence. It is easily distinguished by its large size and cherry-red to dark yellow inflorescence. It is the only one of the four with a red or dark yellow inflo- rescence that is found in Costa Rica and Panama.

Hybridization is thought to occur between H. lankes- terl and H. vaginalis . This is discussed in a later chapter. 236

±« Illllllllllllllll

•-2

Illlll

III

mill

III |.

-&f ET 3

Illllr

UNITCO STATCS niATlO'VJiL MumCUW

PLANn or CO«T* BIC*

PIATE 28. HELICONIA lANKESTERI Standi.

Holotype: H. lankesterl Standi,. La Estrella. Prov CaTtfto-n * Costa Rica. 26. 2? March 192lf. p! C. Stanley ' 39^94 (S)!^ Map 28. Distribution of Hellconla lankesterl * 238 i

Representative specimens

COSTA RICA. ProY. Cartago . P. C. Standley 39927. 30 March

192^, moist forest, vicinity of Orosi (US).

PANAMA. Prov. Bocas Del Toro . P. ,H. Allen ^^973. 5-7 Aug.

19^7. Robalo Trail, northern slopes of Cerro Harqueta, alt. 6000-7000 ft. (P, MO). Prov. Chiriqui . P. H. Allen 3520, 2k May 19'*6. vicinity of Cerro Punta, alt. 2000-2500

m. (MO, NY). M. E. Davidson 62, 6 Jan. 1938, rain forest,

Bajo Chorro, Boquete District. El. 6OOO ft. (P, GH).

29. HELICONIA MOLINAEANA R. R. Smith, sp. nov.

Type J Honduras, Dept. Cortes, bosque nebuloso de Cusuco

en Montana Idalfonso al norte de Cofradia, alt. I6OO m.,

17-18 Abrll 1957. A. Molina R. (P).

Moderately stout plant, 1-2 m. tall; musoid habit.

Leaf-blade ovate, if6 cm. long, 2^ cm. wide, short acuminate apex, rounded-short acuminate base; upper surface green,

lower surface glaucous; petiole at least 30 cm. long,

somewhat stout. Inflorescence erect, highly pubescent,

30 cm. long, peduncled; rachls straight, 0.7 cm. diameter, woolly with light tan hairs. Branch-bracts 11, red, highly villous along sides and base becoming less so toward apex, narrowly linear-lanceolate, strongly curved upward, shallow boat-shaped; lower branch-bracts I8-30 cm. long; middle and upper branch-bracts 10-12 cm. long; all branch-bracts 1.0-1,3 cm, side-width, Ploral-bracts few, about k,3 on. long and 1 cm. wide; hirsute outer surface, glabrous Inner surface. Flowers 3-5 per branch-bract, yellow, 5.5 cm* long; perianth with calyx strongly hirsute, k cm. long} pedicel 1.0-1. 5 cm. long, vlllous-tomentosej ovary nearly glabrous. Prult not seen. (Plate 29.)

Local, In cloud forests. Endemic to Honduras. \ (Map 29.)

This species Is closely related to H. adflexa In the appearance of the Inflorescence. They both are of a

"candelabrum" shape. They differ In the following wayst

1) The lower bracts are generally longer In H. adflexa.

2) The leaf-blades are more ovate In H. mollnaeana

and more oblong In H. adflexa .

3) H» mollnaeana Is much more pubescent throughout

than H. adflexa.

Only representative specimen for this new species la the type. ^ttlfn HONDURAS

PLATE 29. HELICONIA MOLINAEANA R. R. Smith

Holotype: H. mollnaeana R. R. Smith, Montana Idalfonso norte de Cofradia, Dept. Cortes, Honduras, 17-18 April 1957. A. Molina R. 8198 (F). Map 29. Distribution of Helicon la mollnaeana.

30. HELICONIA ADFLEXA (Griggs) Standley. Jour. Wash.

Acad. Scl. 17t 162. 192?. Blhal adflexa Griggs. Bull.

Torr. Hot. Club i^2l 325. Pig. 5. 1915. Typei Guatemala,

Prov. Alta Verapaz, Coban, alt. 1,600 m.. In deep forest,

June 1908. Coll. H. von Tuerckhelm. II 2356 (NY). Isotypes (US. GH).

Moderately stout plant, 1.5-2.5 m. tall; musold habit.

Leaf-blades oblong, 60-120 cm. long, 20-55 cm. wide, short

acuminate apex, rounded or slightly acute at base, some-

times unevenly; upper surface green, lower surface slightly

paler green or sometimes waxy; petiole prominent, about ^0

cm. long or more. Inflorescence stiffly erect, pubescent,

18-30 cm. long; peduncled; rachls slightly flexuose,

puberulent to tomentose, lower part of rachls 0.5-1.2 cm.

diameter. Branch-bracts lO-l^f, scarlet, slightly pubescent

on sides, moderately to heavily pubescent at base, narrowly

linear- oblong, shallow boat-shaped, strongly bent upward

near base; lowest sterile branch-bract. If present, extend-

ing Into leaf, 40-75 cm. total length; lowest fertile

branch-bract 20-35 cm. long, middle 10-12 cm. long, upper-

most 6-8 cm. long, all 1.0-1,5 cm. side-width; margins

usually turned outward and down; Intemode of lower branch

-bracts on rachls 2-4 cm. Ploral-bracts persistent, k cm.

long, 1 cm. wide, slightly pubescent within and moderately pubescent without. Flowers 7-25 per branch-bract, yellow, sparsely pilose, 4.5-7.5 cm. long; perianth 3.5-4.5 cm. 245

long, calyx tomentose without, corolla glabrous j pedicel

1.0>3*0 cm. long, slightly to densely tomentose. Fruit dark blue and glabrous at maturity, 0.6-0*7 cm. diameter.

(Plate 30.)

Found In deep forests at altitudes of approximately

1500 m. Apparently restricted to southern Mexico and northern Guatemala. (Map 30.)

As the name Indicates, all the branch-bracts are r strongly curved upward. This gives the appearance of a

candelabrum and relates this species to H. llbrata , H.

lankesterl , and H. mollnaeana . H. adflexa Is most closely related to H. mollnaeana . Their relationship Is discussed under the previous section on H. mollnaeana .

One specimen (Steyermark 3OO38 (F)) was examined which

Indicates that H. adflexa hybridizes with H. colllnslana .

This Is treated In a later chapter.

Representative specimens:

GUATEMAIA. Dept. Alta Verapaz . 0. P. Cook and C. B. Doyle

200, 1904. vicinity of Sepaculte. alt. ca. 1000 m. (US).

A. Molina R. and A. R. Molina l2295t 15 May I963, Matorr- alles y bosque mlxto N. 0. de Tactic, 6 kms. a Ester, alt.

1500 m. (F). P. C. Standley 90^95. 30 March 1941, wet forest near Tactic, above the bridge across Rio Frio, alt. ca. 1400-1500 m. (F). P. C. Standley 71230, 9 April 1939, dense wet forest, moimtalns e. of Tactic, on road to • K\\ lOKIIIKII M

FLORA VON GUATEMAL*. DEPT «IT« V£R«PA;

PIATE 30. HELICONIA ADFLEXA Griggs Holotype: H. adflexa Griggs, Coban. Dept. Alta Verapaz. Guatemala, April I9O8, H. von Tuerckheim II 2356 (US). Map 30, Distribution of Hell con la adflexa. Zk8 249

Tamahu, alt. I5OO-I65O m. (P). H, von Tuerckhelm II 2356,

June I9O8. Coban, I6OO m. In Hochwald (GH). Dept. Baja

Verapaz . H. von Tuerckhelm II I762. Panreal (US). Dept.

Huehuetenango . J, A. Steyermark 48673. 15 July 1942, dense

rich wet woods between Yulhultz and Maxbal, Sierra de los

Cuchumatanes, alt. 1400-1500 m. (P). Dept. Quezaltenango ,

P. C. Standley 68II7, 8 March 1939. damp forest, Chlqul- hulte, alt. ca. 1410 m. (P). J. A. Steyermark 35171, 21

Jan. 1940, moist slopes of ravine, western slopes of Volcan

Zunll, opposite Santa Maria de Jesus, alt. I5OO m. (P).

Dept. San Marcos . J. A. Steyermark 37491, 11 March 1940,

Plnca El Porvenlr along Rio Chapal, south- facing slopes

of Volcan Tajumulco, alt. I3OO-I5OO m. (F).

MEXICO. State Chiapas . K. and E. Roe, S. Mori 1255, 13

Aug. 1965, steep slope along eastern ridge of Chiapas

Highlands, I5 km. n. of Pueblo Nuevo on highway 195, alt. ca. 1500 m. (WIS). State Vera Cruz . Botterl 523, s.e., Orizaba (US, GH).

31. HELICONU METALLICA Planchon and Linden ex

Hooker. Curt. Bot. Mag. t. 5315. 1862. Blhal metalllca

(Hooker) 0. Ktze., Rev. Gen. PI. 2t 685. I89I. Typei

Colombia, Santa Marta, at foot of Sierra Nevada, Linden and M. Schllm. The type Is an illustration accompanying the description. Moderately stout plant, 2-3 m. tall; muPold-cannold

habit. Leaf-blade oblong. 5O-8O cm. long, 8-21 cm. long

aciunlnate apex, obtuse base; upper surface rich bright

green, lower surface tinted with purple, dries a yellowish

-tan color; petiole if-10 cm. long. Inflorescence erect,

triangular shape, 23 cm. long, glabrous; peduncle pilose

-tomentose, 0.^* cm. diameter; rachls slightly flexuose,

0.5 cm. diameter. Branch-bracts 5-7, greenish, often

tinted purple, somewhat pllose-tomentose, lanceolate,

shallow boat-shaped; lower branch-bracts 7-20 cm. long;

upper branch-bracts 4-8 cm. long; all branch-bracts 0.9

-1.3 cm. side-width; intemode of branch-bracts on rachls

1.0-2.0 cm. Floral-bracts deciduous, lanceolate, k cm. long, 0.6-0.8 cm. wide. Flowers 2-4, sometimes up to 25 per branch-bract, bright red to pink, 6 cm. long; perianth nearly 4-5 cm. long; pllose-tomentose along lower part of calyx, corolla glabrous; pedicel 0.6-1.0 cm. long with short scattered hairs. Fruit dark, subglobose, O.9 cm. diameter, O.7 cm. long. Seed gray-brown, sclerlfied and rough; 0.6-0.7 cm. long and 0.6 cm. wide, 2 sides flattlsh. third side rounded. (Plate 3I.)

Along forest trails, 10-600 m. altitude. In Central America, H. metalllca is found only in Panama, which is evidently the northernmost extension of the range for this Colombian species. (Map 31.) 251

PIATE 31. HELICONIA METALLICA PI. et Llnd. ex Hook.

H. metalllca PI. et Lind. ex Hook., region n. of El Valle de Anton, Prov. Code, Panama, 13 Jan. 19^1, P. H. Allen 2909 (MO). t

H. metalllca was Introduced into cultivation about

I856. It Is an Interesting species distinguished by the

greenish bracts, red-pink flowers, and leaves In which the lower surfaces are usually tinted purple.

Another name which has been used for this species Is

H, ma rant 1 folia Shaw. The description for this name is

extremely vague, and the location for Its type Is St.

Vincent Island, where H. metalllca Is xmknown. This name has been discarded In reference to H. metalllca .

Representative specimens

PAJIAMA. Canal Zone . H. Pittler 2685, 8 Feb. I9II, around

Frljoles, alt. 10 to 30 ml. (US). E. Tyson, J. Dwyer, K.

Blum and J. Duke ^^838, 18 July I966, woods, 2 ml. e. Sante

Pe (PSU). Canal Zone (?) . R, S. Williams IO3O, 3-9 April I9O8. Marragantl and vicinity (NY, US). Prov. Cocle »

P. H. Allen I669, 12 Feb. 1939. north rim of El Valle de

Anton, alt. 600-1000 m. (MO). P. H. Allen 2909, I3 Jan.

19'*1, north rim of El Valle de Anton (GH, MO, US). J. E.

Eblnger 952, 21 Aug. I96O. El Valle. road behind Pension, secondary growth forest (MO). Prov. Darlen . P. H. Allen

259. 17 March 1937. trail between Plnogana and Yavlsa, ca. 15 m. (F). P. .H. Allen 4282. k March 19^7. vicinity

Plnogana. 20 m. (F). R. A. Terry and M. E. Terry 11*31.

9 March 19^0. Cana-Cuasl Trail. Camp 2 (P, MO). E. Tyson. J. Dwyer. K. Blum and J. Duke 4710. 16 July I966. 3 mi. e. Map 31, Distribution of Helloonia metalllca.

255

Santa Pe (PSU), Prov, Panama . K. E. Blian and K. Miller

2295 t 9 May I966. Cerro Campana, El. ca. 2300 ft. (PSU).

32. HELICONIA OSAENSIS Cufodontls, Archlvlo Bot. 9»

I89. 1933» Type: Cpsta Rica, In region of Pacific,

Peninsula Osa ad Golfo Duloe, In sllva prlnaeva ad Hlo

Nuevo circa Puerto Jimenez* alt. 30 m.. G. Cufodontls

158 (W).

Moderately slender plant 1. 5-3*0 m. tall; somewhat cannoid habit. Leaf-blade narrowly oblong, lower leaf

-blades 80-90 cm. long, upper leaf-blades '4-0-75 cm. long, all leaf-blades 10-12 cm. wide; short acuminate apex, acuminate base; upper and lower surfaces green, sometimes lower surface tinted purple; petiole short, k-7 cm. long or absent. Inflorescence erect, 10-25 cm. long, somewhat pubescent; peduncle 18-25 ciii* long, pubescence peeling off; rachis pubescent, slightly flexuose, 0.3 cm. diameter.

Branch-bracts 4-9, dark red to orange- scarlet, narrowly lanceolate, long acuminate, shallow boat-shaped, moderately curved upward; matted pubescence falling off giving appear- ance of roughness; lower branch-bracts 5*0-13.5 cm. long; upper branch-bracts 4.0-6.5 cm. long; all branch-bracts

0.8-1.0 cm. side-width; intemode between lower branch

-bracts 2-3 cm. Ploral-bracts deciduous, 3. 0-3. 5 cm. long, 1 cm. wide. Plowers 5-10 per branch-bract, yellow. '

5»0-5.5 cm. long; perianth 4.0-4.5 cm. long slightly plloae-puberulent; pedicel 0,9-1,5 cm, long, slightly

pubescent-tomentose. Fruit metallic blue at maturity,

subglobose, about 0,8 cm, diameter. Seed with sclerlfled,

rough seed coat; 0,6 cm, long, 0,5 cm, wide, (Plate 32.) In moist forests and along river margins. Once

thought to be endemic to Costa Rica, now also known from

Nicaragua and Panama, (Map 32,)

H, osaensls Is closely related to H, aurantlaca . H, hlrsuta . and H. vaginalis . It differs from these species

In that It has a long peduncle, all bracts are fertile, the lowest branch-bract does not have a leaf-blade exten- sion, the leaves are narrower and much longer with acximlnate apex and base. The pedicels are short, and

like H. hlrsuta , they are pubescent.

Representative specimens:

COSTA RICA. Prov. Alajuela , A. M, Brenes 22^f67, 20-28

March 1937, La Costa del Paclflco, Los Laros (P, NY),

P. C. Standley 40149, 2 April 1924, moist forest, vicinity of Capulln, on the Rio Grande de Tarcoles, alt, ca, 80 m.

(US). Prov, Puntarenas , A. Tonduz 9968, March I896, forests of Santo Domingo of Golfo Dulce (US), M. Valeric

433. 25 Feb. 1933. Playa Blanca, Golfo Dulce (P).

NICARAGUA. Prov. Zelaya . P. C. Engleslng I38, 11 Feb.

1928, region of Braggman's bluff, growing under and among second growth In heavy well drained clay soil at San 257

PIATE 32. HELICONIA OSAENSIS Cuf.

H. osaensls Cuf., Bahia Honda, Panama, 28 March 1939 P H. Elmore H29 (P). Map 32. Distribution of Helloonla osaensls. 259 260

Antonio de Susum, up to 600 ft. alt. in n.e. Nicaragua

(P. US). L. E. Long k5, 8 April 19^+7. vicinity of El

Recreo (P). A. Molina R. 2332, 23 April 19^9, Guamil de

segunda close. Area del Ocotal, Rio Grande, alt. 0.15

(P, GH). P. C. Standley 19363, 23 April - 1*^ May 19^*9. moist mixed forest, vicinity of El Recreo, on Rio Mico,

ca. 30 m. (P).

PANAMA. Prov. Chiriqui . P. H. Elmore 29, 28 March 1939.

Bahia Honda (P, MICH, US). R. E. Woodson. R. W. Shery

889, 28 July - 1 Aug. 19*^0, vicinity of San Bartolome,

Peninsula de Burica, alt. 0-50 m. (MO),

33. HELICONIA LINNAEANA Lane ex R. R. Smith, sp. nov.

Type I Trinidad. Emperor (Morrison) Valley, back of Look

-Out, 5 Sept. 1911, W. E. Broadway (NY).

Slender plant, 2-3 m» tall; musoid in habit. Leaf

-blade oblong-elliptic, 6O-7O cm. long, 15-18 cm. wide, acuminate apex, rounded or slightly acute base; upper

surface green, lower surface green or glaucous; petiole prominent. Inflorescence erect, 18 cm. long, glabrous,

long peduncled; rachis slightly flexuose, 0.4 cm. diameter.

Breaich-bracts 5-10 red with green margin, lanceolate-ovate with acuminate apex, moderately shallow boat-shaped; lowest branch-bract extends into leaf-blade, total length about

^0 cm., sterile; lowest fertile branch-bract about 12-18 cm. long; middle and upper branch-bracts gradating from

10 cm. to k cm. in length; all branch-bracts 2.0-2.5 cm. i

261

slde-Nldth; internode between lower branch-bracts 2 cm. long. Floral-bracts several, lanceolate, thin, membraneouat

4 cm. long, about 0.8 cm. wide, glabrous. Flowers 5-7 per

branch-bract , glabrous, J,0 cm. long: perianth 2.5-4«5 co* long, glabrous, pedicel 0.5 cm. long, glabrous. Fruit bluish, subglobose to 3-slded, 1.0 cm. long, 0.8 cm. wide.

Seed not seen. (Plate 33»)

Infrequent In moist shaded areas and along stream banks. This species Is found on Trinidad and Tobago.

(Map 33.)

This Is a species which has apparently been long overlooked. It has larger bracts than H. trlnldatls and

smaller bracts than H. blhal , the only other two species which grow within Its range.

H. llnnaeana may be an extension of a South American species. Search through the literature suggests that It

Is a valid species In Its own right. However, the more closely related South American species, such as H. humllls

Jacq., should be thoroughly studied, and more specimens of

H. llnnaeana need be examined.

Representative specimens

WEST INDIES. TRINIDAD. N. L. Brltton, E. G. Brltton and

T. E. Hazen 1951. 18 April 1920, moist shade. Cocoa

Plantation, base of Mount Tamana (NY). W. E. Broadway

69^*9 . 11 May 1928, Orppuche local road, via Valencia, near 262

TRINIDAD

PIATE 33. HELICONIA LINNAEANA Lane ex R. R. Smith

Holotypes H. llnnaeana Lane ex R. R, Smith, Emperor (Morrison) Valley, Trinidad, 5 Sept. 1911, W. E. Broadway s. n. (NY). Map 33* Distribution of Heliconia llrmaeana.

i. /. -:t.

264 265

the stream (MO). R. A. Howard 10339. 9-23 Feb. 1950. edge of thickets In Arlpo Savannah on Waller Air Force Base

(GH).

TOBAGO. Eggers 5768. Nov. 1889 (NY).

3^. HELICONIA SCHIEDEANA Klotzsch. Llnnaea ZOt kSj,

18k7» Blhal schledeana (Kl.) 0. Ktze., Rev. Gen. PI. 2t

685. I89I. Typet Mexico, Bararla de Tloselo, Oxtep.,

Sohlede I03I (B).

Hellconla hlrsuta Cham, and Schlect. In Llnnaea 6t

57. I83I. non Hellconla hlrsuta L. f. Typei

(Berlin, Bot. Mus., not seen).

Hellconla pochutlensls Conzattl, PI. Taxon Mei. 3»

129 • 19'*7» Typei Colectada In Pochutla, Oas.

y ahora cultlvada en el Jardlnj not seen.

Moderately stout plant, 1.5-3«0 m, tall, musold habit.

Leaf-blade oblong-ovate, 5O-8O cm. long, 15-20 cm. wide; upper surface green, lower surface paler green; petiole prominent, 27 cm. long. Inflorescence erect, pubescent.

25-30 cm. long, sometimes longer; peduncled; rachls slightly flexuose to straight, densely tomentose pubescent,

0. if- 0.5 cm. diameter; rachls spiralled so branch-bracts extend out In all directions. Branch-bracts 6-12. dark red. red to green, or bright yellow to orange, oblong

-lanceolate with acute or obtuse apices, shallow 266

boat-shaped, may become strongly reflexed while maturing! lowest branch-bract usually sterile, often with leaf-blade extension, 17-22 cm. long; lowest fertile branch-bract

7-15 cm. long; middle branch-bracts 5-10 cm. long; upper branch-bracts 3-7 cm. long; all branch-bracts 0.6-0.8 cm. side-width; Intemode between branch-bracts 1.0-3.0 cm.

Floral-bracts long ovate to wide lanceolate, J-^ cm. long, 0.5-1*0 cm. wide; often pubescent on outer surface.

Flowers 6-21 per branch-bract, yellow to pale dull yellow, densely villous, 1.5-6. 5 cm. long; perianth 2.5-5*0 cm. long, villous; pedicel 1.0-1.5 cm. long vlllous-pllose.

Fruit dark, subglobose to 3-slded, villous, about 1 cm. diameter. Seed 0.8-0.9 cm. long. (Plate 3^.)

Steep slopes of mountains and ravines, moist forested regions, and banks of streams and rivers. Range for H. schledeana extends over central and southern Mexico,

Giiatemala, British Honduras, and Honduras. (Map 3^*)

H. schledeana Is a species with two varieties with possible hybridization at the varietal level. It Is apparent that It also hybridizes with other species.

This phenomenon Is discussed In the next chapter.

H. schledeana Is not closely related to other species of Helicon la . The outstanding characteristics of the species are the pubescence of the Inflorescence, and showy floral-bracts extending over the long thin branch-bracts.

The fruits are exposed on long, hairy pedicels, and not hidden within the branch-bracts. PIATE 3^. HELICONIA SCHIEDEANA Kl. var. SCHIEDEANA

H, schledeana Kl, var. schledeana , banks of Creek El lissadoo Vera Cruz, May 1933» C. A. Purpus 12092 (US), 268

Key to varieties of Hellconla schledeana

B. Branch-bracts red or red-greenj perianth 3»5-5»0 cm.

long..... var. schledeana

B. Branch-bracts yellow or yellow-green j perianth 2.5

-3.0 cm. long var. aplssa

Hellconla schledeana Kl. var. schledeana

Leaf-blade oblong-ovate, 50-60 cm. long, 16-20 cm. wide, acuminate apex, obtuse base; petiole long. Inflo- rescence erect, pubescent, 15-'+^5 cm. long; rachls slightly to moderately flezuose. Branch-bracts 6-15, dull red to red-green; middle branch-bracts 6-10 cm. long. Flowers

6-21, pale yellow 3 '0-6.5 cm. long;- perianth 3»5-5«0 cm* long, villous. Fruit dark, subglobose and 3-si

Steep, moist slopes of ravines and mountains.

Endemic to Mexico.

This variety Is much more variable than var. splssa .

There Is a greater difference In length of the perianth, number of branch-bracts, and number of flowers per branch

-bract. The pubescence on the leaf-blade and petiole Is also somewhat variable as Indicated by Loesener (1913)«

He described a new form of H, schledeana , forma glabrl- folla , from a specimen collected In the State of Vera

Cruz, Mexico. i

269

Representative specimens

MEXICO. State Chiapas , R. L. Dressier lUO**, 9 July 195'*» hills above Ocosingo, elev. oa. 900 m., oak forest (GH).

State Guerrero . E. Matuda S - 30, 13 May 1937. Atoyao,

(MO). State Hidalgo . 0. M. Clark 7*106. 3 July 1935. n.

Chaphuacan (NY). H. E. Moore. Jr. 2719. 28 April 19^*7.

steep slopes and ledges near km. 340-3^1 on highway below

Chapulhuacan, Jacala district, alt. 2600 ft. (GH, MICH).

H. E. Moore. Jr. 3^03, 16 July 19^7, slopes by km. 341 on highway n. of Chapulhuacan, district Jacala (MICH). State

Oaxaca . B. P. Reko 4112, 29 March 1919. Choapan to Cou-

caltepec, alt. 800 m. (US). State San Louis Potosi . M*

T. Edwards 527. 12 July 1937. Arroyo de Poistapa,

Tamazunchale (MO). M. T. Edwards 625. 22 July 1937.

Barrio de San Juan, Tamazunchale (F, MO). R. M. King

4425. 1 April 1961, locally abundant, open sun, mountains along the gravel road to Jalpan ca. 6 mi. n.e. of Xilitla

(F, NY. US). State Vera Cruz . I. Kelly 128. 1947, plants

used by the Totonac Indians at Taj in, district of Papantla

(WIS). H. E. Moore, Jr. and Max Cetto 62I8, 6 April 1952, moist shaded, steep, narrow ravine near km. 33I about 1

km. before Portln de las Flores on road from Tehuacan to

Orizaba (GH). H. E. Moore and H. 0' Gorman 63IO, I9 April

1952, slopes of ravine about 3 mi. beyond Papantla on road

to Poza Rica. alt. 240 ml. (GH). C. A. Purpus 8268. July t

1919. Zacuapan (MO. NY, US). C. A. Purpus 12092, April

193^, moist location. Banks of Creek El Ulsadol, Zacuapan (F, US).

Heliconla schledeana Kl. var splssa (Gri ggs) R. R. Smith, var. nov ~, ' —

Heliconla splssa Griggs. Bull, Torr. Bot. Club JOt

652. 1903. Blhal splssa (Griggs) Griggs, Bull. Torr. Bot. Club 31 X ^^5» 190^. Type I Guatemala, Dept. Alta Verapaz, near Flnca Sepaculte. 0. P. Cook and R. R. Griggs 359, 30 March I9O2 (US).

Leaf-blade oblong-ovate, 60-75 cm. long, 20 cm. wide, acute apex, rounded base; petiole long. Inflorescence

erect, pubescent. I5-35 cm. long; rachls red. nearly straight. Branch-bracts 9-12. bright yellow to orange; middle branch-bracts 5-7 cm. long. Flowers IO-I7 per branch-bract, yellow. 4 cm. long; perianth 2.5-3.0 cm. long, villous. Fruit dark yellowish color, subglobose •

to 3-slded. 1 cm. diameter; seed 0.8 cm. long. 0.7 cm. • wide; seedcoat sclerlfled, rough, somewhat wrinkled. Rain forests, swampy areas, thickets near rivers, and damp forested slopes. Distribution Includes Mexico. Guatemala. British Honduras and Honduras.

Representative specimens

BRITISH HONDURAS. Dlst. Belize . C. L. Lundell 6950, Jan. - June 1936. In high forest, churchyard on the Slbun River (MICH, NY). Plat. El Cayo , H. H. Bartlett II5II, I5 Feb.

1931. between El Cayo and Benque Vlejo (MICH).

GUATEFALA. Dept. Alta Verapaz . P. C. Standley 7057I,

5 April I939t thicket along river, Pantin, below Tamahu, alt. ca. 600 m. (F). J. A. Steyermark ^3899, I9 Feb.

19^2, damp forested slopes, along road, between San

Cristobal Verapaz and Chlxoy, alt. I2OO-I3OO m. (P).

Dept. Pet en . H. H. Bartlett I2638, 12-15 April 1931,

Tikal (GH, MICH). C. L. Lundell 289O, I3 April 1933,

Santa Teresa, Subin River (F, GH, MICH). J. A. Steyermark

^16089, 30 April 19^2, Cerro Ceibal, between mouth of Rio

Santa Monica and mouth of Rio San Martin, on left side of

Rio Canchen, alt. 75-150 m. (F).

HONDURAS. Dept. Comayagua . A. Molina R. IO873, 18 Julio

1962, frecuente en el matorral de Quebrada Montanuelas, bosque mixto entre Trincheras y Montanuelas, alt. 1400 m.

(F). J. B. Edwards 598, 5 May 1933. In swampy ground in

Palm Grove, San Louis, alt. 2500 ft. (F, GH). Dept. El

Paraiso . A. Molina R. llkOk, I5 Marzo I963, en el bosque denso y humedo Montana Cifuentes entre El Urraco y Cifuentes, alt. 9OO m. (F).

MEXICO. State Chiapas . R, L. Dressier 158if, 20 July

- 20 Aug. 195^. near Laguna Ocotal Grande, ca. 25-30 km. s.e. of Monte (Cerro) Libano elev. ca. 950 m. (GH). Map 3^4'. Distribution of Hellconla schledeana. 273 35* HEITCONIA VOXir^RUZII R* R. Smith, ..p. nov.

Typf... Ke.^.ACo, State Veracruz, V'-lcan San .^iartln, ios

Tuxtlc-j at .iide of stream on south-facing slope of

7olc.-a.io. Dec. 1959. R. L. Kouck 35 (KICH).

Koderately stout plant, about 2 m. tall. Leaf-blades

cblong, 40-8G cm. long, 11-24 cm. wide, acuminate apex,

rounded to broad-octue base; petloled. Inflorescence

erect, pubescent; 20-30 cm. long, peduncled; rachis nearly-

straight, tomentose (more so when young), lower part of

rachis 0.5 cm. diameter; at maturity rachis spiralled so

that branch-braots extend out horizontally in all direc-

tions. Branch-bracts 7-9, bright red, puberulent to

tomentose on outer surface along bottom and near base,

linear- lanceolate, shallow boat-shaped, lower branch-bracts

extend out horizontally, upper ones curve upward; lowest

branch-bract sterile, extending into leaf-blade, total

length about 40 cm.; lowest fertile branch-oraco II-I6 cm. long, 0.7-1.0 cm. side-width, upper branch-bracts 6 cm. long, 0.7-0.8 cm. side-width | intemode between lower branch-bracts 2,5-3.0 cm. Floral-bracts about 4 cm. long, pubescent on margin and tomentose along midrib on outer surface. Flowers 3-4 per branch-bract, yellow, 4.5-5.5 cm. long; perianth 3 cm. long, calyx glabrous to puberulent. corolla appears glabrous; pedicel variable in length 1.0 -2.5 cm., tomentose. Fruit not seen. (Plate 35.) PIATE 35. HELICONIA VERACRUZII R. R. Smith

Holotype: H. veracruzll R. R. Smith, Los Tuxtlas, Veracruz, Mexico, 26-28 Dec. 1959. R. L. Houck 35 (MICH). Map 35» Distribution of Hellconia veracruzil.

.

Frequent along itream margins and In moist forests.

Endemic to Mexico. {'Aa-p 35.)

T'nls species is close to H. vaginalis in general appearf.nce. Both species have red bracts and yellov:

flowers. Outstanding differences are several, H. vera-

<^ruzii is much more pubescent than H. vaginalis . H. Vera -

cruz ii is petioled; H. vaginalis is sessile or nearly so.

One of the more distinct features of H. veracruzii is t^iat the rachis is usually contorted so that the branch-bracts extend out in all directions. H. vaginalis has distichous branch-bracts

Representative specimens!

MEXICO. State Tamaulipas . P. S. Martin 01?, I5 Feb. 1953,

Gomez Farias region, near Aserradero del Paralso, I7 km. by road n.n.w. of Chamal, ca. 12 km. s.w. of Gomez Farios, elev. .500 m. (MICH).

36. HELICONIA lATISPATHA Bentham, Bot. Voy. Sulph.

170-171. 1844. Blhai latlspatha (Benth.) Griggs, Bull.

Torr. Bot. Club 31j 445. 1904. Typei Salango (off the coast of Ecuador) (K).

Heliconla meridensis Kl. in Llnnaea 20 j 463. 1847, Bihal meridensis (Kl.) 0. Ktze. Rev, Gen, PI, 2i 685. I89I. Type I in sylvls umbrosis, humldls,

Columbiae, Moritz n, I287. Flor. in Decembr.

(Vienna Nat. Hist. Mus., not seen). Moderately slender plant, 1.0-2.5 m. tall; somewhat musoid In hadt. Leaf-blade elliptic to elllptlc-ovate,

30-100 cm. long, 13-30 cm. wide, short acuminate apex, obtuse to truncate base; upper and lower surfaces green, aldrib somewhat pubescent; petiole prominent, 20-50 cm» long, 1 cm. diameter. Inflorescence erect, occasionally nodding, glabrous, 20-35 cm. long; peduncled; 0.7-C,8 cm. diameter; rachis moderately to slightly flexuose and spiral, occasionally pilose, 0.6 cm. diameter. Branch

-bracts 4-11, variable in color, greenish yellow, golden orange, orange with greenish tip, or red and yellow, glabrous, narrowly lanceolate, subcordate base, shallow boat-shaped, extends out from base horizontally, then curves moderately upward; lowest branch-bract often extended into leaf-blade, sometimes sterile, total length

30-75 cm., lowest fertile branch-bracts 9-20 cm. long; middle and upper branch-bracts 5-1^ cm. long, progressively becoming shorter; 1.8-2.5 cm. side-width; Intemode between lower and middle branch-bracts 2.5-3.0 cm.; intemode between upper branch-bracts 1.5-2.5 cm. Floral-bracts translucent yellow, lanceolate-triangular ovate, 3.0-4.5 cm. long, 1-2 cm. wide. Flowers 4-6 per branch-bract, yellow to yellow-green. 3.5-4.5 cm, long; perianth 3-4 cm. long, calyx slightly pubescent along margins, free sepal often curved back at anthesls, corolla glabrous; pedicel 0.5-1.5 cm. long, glabrous. Fruit dark, subglobose to 3-slded, O.7 cm, diameter dried, mature fruits usually remain hlGden In braces. Seed sclerlfled, rough seed coat

0.7 iit.. long, 0.6 cm. wide. (Plate 36.)

Found in creek beds, on moist wooded slopes moist to wet forests, swampy Jungles, and wet thickets. Common plant In all of Central America. (Map 36.)

H. latispatha is quite variable In the color of the branch-bracts. The plants of Honduras and the northern part of Central America are much more yellow, whereas, the plants of Panama appear to have a considerable amount of red to red-orange In the branch-bract extremity with the predominant yellow color nearer the base.

H. latispatha Is a well-known species, although In the past several other species have been called by Its name. Within its geographical limits H. latispatha may be confused with H. tortuosa In habit, but tne latter has a tomentose rachis and deep red bracts. It is thought

that this species hybridizes with H. vaginalis , and H. coll ins iana . This is discussed in a later chapter.

This species is known by the upright, widely-spaced, long and somewhat flat, spiralled, branch-bracted Inflores cence. It is widely distributed from Mexico throughout Central America and into South America, 281

\

PIATE 36. HELICONIA lATISPATHA Benth.

H, latlspatha Benth,, between Prijoles and Monte Lirlo, Canal Zone, Panama, 18 Oct. 1922, E. P. Killip 12121 (US). j

Representative specimens

BRITISH HONDURAS. Dlst. Belize . P. H. Gentle 1322. 20

July 193^. Maskall (F, GH, MICH, NY, US). Dist. Corozal .

P. H. Gentle 520, 1931-32 (GH, MICH). Dlst. El Cay6 . H.

H. Bartlett II509, 15 Feb. 1931. between El Cayo and Benque

Vlejo (F, GH, MICH, MO, NY, US). C. L. Lundell 6655, July

-Aug. 1936. Mountain Pine Ridge, San Agustln, In creek beds (GH, MICH, NY). Dlst. Stann Creek . W. A. Schlpp

329, 1 Sept. 1929, Middlesex, 200 ft. alt. (P, GH. MO, MICH, NY).

COSTA RICA. Prov. Alajuela . R. W. Holm and H. H. litis

697, 1 Aug. 19^9. open, wet banks of Rio Frio, vicinity

of Los Chiles (GH). Ad. Tonduz 66O7, Feb. I892, Bord du

Rio Nacum pres de Buenos Aires, 25O m. (US). A. Jimenez

M. 2321, 10 Setiembre 1964. Quebrada Lajas, Flnca Los

Ensayos, Buena Vista de San Carlos, alt. 85O m. (F). Prov. Cartago . Ad. Tonduz II387, Nov. 1897, Bulssans et broussailles a Tuis, alt. 65O m. (MO, US). Prov. Guana - caste. P. c. Standley and J. Valeric 46357, 29 Jan. 1926, brushy slope, Agullares, alt. ca. 600 m. (US). L. 0.

Williams, A. Molina R. and T. P. Williams 26362, 31 Dec. in 1963, swamp, sparse broad-leaf forest 5 km. s. of Pinas Blancas, alt. 300m. (F). Prov. Heredla . A. Jimenez M.

2646, 10 Julio 1964, Rio La Paz Grande, entre Cariblanco y Vara Blanca. alt. I35O m. (P). Prov. Llmon . C. W. Dodge 283

and V. F. Goeger WS, ?9 July 1936, forest near farmhouse at Flnca Castllla, 30 m. (MO). A. Jimenez M. 2083, 1?

Julio 1964, Unos 2.5 km. al Este de Guaplles, alt. 250 in« (F). Prov. Puntarenas . W. Burger and G. Matta U. ^^765,

27 and 28 Jan. I967, along edge of trail In open sun or

partial shade, steep forested slopes above Golflto along

trail to television tower, alt. 100-300 m. (F). Prov. San

Jose . P. C. Standley 36l97t 2-4 March 1924, wet forest.

La Hondura, alt. I30O-I7OO m. (US).

EL SALVADOR. Dept. Ahuachapan . S. A. Padllla 58I, 1923

(US). P. C. Standley 19996, 9-27 Jan. 1922, vicinity of Ahuachapan, alt. 800-1000 m. (GH, US). Dept. La Llbertad .

N. C. Passett 28273. 11 Oct. 1950, wet swale near Ateos.

31 km. w. of San Salvador (F, GH, WIS). Dept. San Salvador.

S. Calderon 898, July 1922, San Salvador (NY). P. C.

Standley 19442, JO and 31 Dec. 1921, In Cafetal, vicinity of Tonacatepeque (GH, NY). Dept. Sonsonate . S. Calderon

1760, July 1923. Sonsonate (GH). P. C. Standley 22128.

22, 25 March 1922, vicinity of Santa Emilia, alt. I35 m. (US). Dept. unknown . W. W. and H. E. Rowlee 42, 16 May I9I8, Peralta (US).

GUATEMAIA. Prov. Alta Verapaz . 0. F. Cook and R. P.

Griggs 262, 29 March I902, Secanqulm (US). H. Plttler

19^, 2 May I905, vicinity of Secanqulm. alt. 400 m. (US). J. A. Steyermark 44645, 4 March 1942, Cerro de Agua Tortuga (Sahacoc), vicinity of Cubllquitz. alt. 350-450 m. (P). ..•3*7.,

284

H. von Tuerckhelm II 5^1^, Aug. I907, Cubilqultz, alt. 35O

m. (F, GH, MO. NY). H. von Tuerckhelm 8^166, M. July I903, Cubilqultz, alt. 350 m. (US). Dept. Esculntla . P. C.

Standley 63^*51. 24 Jan. I939, wet thicket, between Rio

Jute and Rio Pantaleon, on road between Esculntla and

Santa Lucia Cotz, alt. 5^0-720 m. (F). Dept. Huehuetenangg

J. A. Steyermark 49200, 22 July 1942, around Ixcan at "Pateushln," Sierra de los Cuchumatanes , alt. 5OO m. (P).

Dept. Izabal . W. A. Kellerman 6334, 20 Feb. I907, Los

Amates, alt. 90 m. (US). J. D. Smith 1829, M. March I889,

Rio Dulce, Livingston, alt. 0 m. (US). P. C. Standley

23703, 15-31 May 1922, brushy slope, vicinity of Qulrlqua, alt. 75-225 m. (US). P. C. Standley 72771. 30 April 1939, wet forest, near Entre Rlos, alt. ca. 18 m. (P). j. a.

Steyermark 38524, 1 April 1940, between Mllla 49.5 and

ridge 6 ml. from Izabal, Montana del Mlco, alt. 65-6OO m. (F). J. A. Steyermark 39793, 21 April 1940, swampy

Jungle In valley of tributary of Rio San Francisco del

Mar, 2 ml. n.w. of Hopl, 12 ml. e. of Entre Rlos. alt. 10-20 m. (P). Dept. Pet en . M. Agullar 319, 28 July I934, La Llbertad and vicinity (P, MICH, MO, NY). Dept. Retal - liuleu. w. R. Hatch and C. L. Wilson 382A and 282B, 23 Aug. 1936, common In second growth, Flnca San Jose Nil (P, GH). W. A. Kellerman 6040. 12 Jan. I907, Retalhuleu (P, UC, US). Rojas 443, 20 June 1923. San Plllpe (US). 285

P. C. Standley 87265, 16 Feb. 19^1, mixed forest along

road between Retalhuleu and Nueva Linda, alt. 120-330 m,

{F)» Dept. Santa Rosa . P. C. Standley 6O629, 21 Dec.

1938, damp thicket, near Oratorio, alt. ca. 1200 m. (P).

P. C. Standley 78899. 1 Dec. 19^*0. damp thicket, region

of La Morenlta, n.e. of Chlqulmulllla, alt. ca. kOO m.

(P). P. C. Standley 79006, 2 Dec. 19^0, wet forested

quebrada, Rio de la Cruz, e. of Taxlsco, alt. ca. 225 m.

(P). Dept. Suchltepequez . P. C. Standley 62172, 5 Jan.

1939, wet thicket, near Patulul. alt. 33O-6OO m. (P). J.

A. Steyermark ^^7685, 17 June 19^2, along Rio Madre Vlejo

bordering virgin forest near Santiago farm, vicinity of

Tlqulsate, alt. 100 m. (P). S. S. White 5I8O, 12 Jan.

19^*^, Hacienda Traplche Grande, s. of Cuyotenango, roadside (MICH). Dept. unknown . S. Watson 5OO, I885, eastern por- tions of Verapaz and Chlqulmula (GH). HONDURAS. Dept. Atlantlda . A. M. Chlckerlng ifO. 22 June

- 27 July 1929. Lancetllla Valley (MICH). E. R. Mitchell

81, 18 April 1926, growing In damp soil In undergrowth at sea level, vicinity of Tela (P, GH). P. C. Standley 53687.

6 Dec. 1927 - 20 March I928, Lancetllla Valley, near Tela, alt. 20-600 m. (P. US). T. G. Yuncker ^^767 a and b. 25 July 193^1. margin of swamp, 30 km. w. of Tela (P, MICH, MO, NY). T. G. Yuncker, J. M. Koepper. and K. A. Wagner

83^9, 10 July 1938, near the bank of the Salado River, near village of Salado (P. MICH, US). Dept. Cortes . 286

A. Molina R. 6952, 12 Mayo 1956, Matorralea entre Tulian y

Cortes (F). Dept. Yoro . P. C. Standley 53892, Dec. 1927,

Quebrada Seca, alt. 30 m. (P, US).

MEXICO. State Campeche . F. D. Barlow 16/13, 21 April

1963, OJo de agua. Beach ridge forest, Santa Leonor, 25 km. e. of mouth of Rio San Pedro y San Pablo (WIS). State

Chiapas . G. L. Fisher 35320, ? Aug. 1935. Tapachula, alt.

150 m. (F, NY, US). E. Matuda s. n., 6-9 July 1939.

Javalinero, Palenque (MICH). E. Matuda I669I, 11 July

19'*?, Esperanza, Escuintla (MICH). E. Matuda I6693, H

July 19^7, Esperonza, Escuintla (GH). E. Matuda 16735,

1^ July 19^7, SM Terena, Acapetahua (F, GH). E. Matuda

18018, 2 July 19^8, in wet bush, Esperanza, Escuitla (P).

State Oaxaca . R. M. King 889, 29 July 1958, growing in rich moist loam in partial shade, along Trans-isthmian highway (route I85) I3 km. s. of village of Mat las Romero

(MICH). H. E. Moore, Jr. 8IO9, 10 Oct. 1959. ravine 35 km. e. of Matias Romero, if km. w. of Palomares on trans

-isthmus highway (GH). C. V. Morton and E. Makrinius 2370,

1-15 April 1933. vicinity of Cafetal, Concordia, alt. kOO

-650 m. (P, US). State Tabasco . P. D. Barlow 2'+/13. 7

May 1963, marsh in milpa, Wolter finca "La Luz" near

Comalcalco (GH, MICH, WIS). E. Matuda 3494, 27 June 1939.

Estapilla, Teuosique (F, GH, MICH). J. N. Rovirosa 8O9. 17 May I89O, Sanctac Rosae (NY). State Vera Cruz . R. L.

Dressier and Q. Jones 5I, 11 Aug. 1953. thicket by road. 287

region of San Andres Tuxtla, near Tapalapan, n.w. of

Santiago Tuxtla (GH). J. Vera Santos 2238, 8 July 19^4-3,

vicinity of Rio Tonto, EJldo de Almlllnga, 6 km. w. of

Campo Experimental de Hule El Palmar, Zongollca (MICH).

C. L. Smith lOkZ, March I895, Coatzacoalcos, Isthmus of

Tehauntepec. (P, GH, MO).

NICARAGUA. Dept. Chlnandega . C. P. Baker 2016, I5 Jan.

1903. Chlnandega (GH, MO, UC, US). W. R. Maxon. A. D.

Harvey and A. T. Valentine 7188, 19-21 June 1923, In shade,

Ameya, near sea level (US). Dept. Comarca de El Cabo .

A. Molina R. I5096, 22 Aug. I965, Matorrales del bosque

secundarlo lluvloso de San Mateo, 16 ml. al sur de Tron-

quera cerca de Rio Wawa, alt. 35 m. (P). Dept. Granada .

L. 0. Williams and A. Molina R. 20027, 5 Jan. I967, In

forest on Mombacho Volcano, alt. I300 m. (P). Dept .

Managua . Bro. A. Gamier 862, s. d. vicinity of Managua

(MICH, US). W. R. Maxon, A. D. Harvey, and A. T. Valentine

7500. 28 June 1923. Los Nubes and vicinity s. of Managua,

alt. 8OO-9OO m. (US).

PANAMA. Canal Zone . E. P. Kllllp 12121, 18 Oct. 1922.

between Prljoles and Monte Llrlo, alt. 30 m. (NY, US).

C. V. Piper 56I8, 26 Peb. 1923, vicinity of Gatuncello

(US). H. Plttler '+062, I9II, arovmd Culebra, alt. 5O-I5O

m. (US). R. C. Shannon s. n., Aug. 1923. Port San Lorenzo

(US). P. C. Standley 252^*5. 27 Nov. - 10 Dec. 1923, Sosa Hill. Balboa (US). P. C. Standley 26023, 5 Dec. 1923. 268

wet forest, Cerro Gordo, near Culebra (US). P. C. Standley

28326, 26 Dec. 1923. moist thicket, Gamboa (US). P. C.

Standley 29OO9, 3I Dec. 1923, along old Las Cruces Trail, between Fort Clayton and Corozal (US). P. C. Standley

31736, 19 Jan. 192^4-. moist thicket, Obispo (US). E. Tyson

1062, l'*^ kng, 1965, near Survival School, Curundu (PSU).

Prov. Chlrlqul . M. E. Davidson II78, 18 Aug. 1938, open areas, Puerto Armuelles (F). R. E. Woodson, Jr. and R.

W. Shery 86^, 28-31 July 19^0, vicinity of Puerto Armuelles, alt. 0-75 m. (GH, MO, US). R. E. Woodson, Jr. and R. W.

Shery 89O, 28 July - 1 Aug. 19^0, vicinity of San Bartolom^

Penlnsual de Burlca, alt. O-5O m. (MO). Prov. Colon . H.

Plttler 4207, 16 Aug. 1911t around Dos Bocas, Rio Fato valley, alt. iJ-0-80 m. (US). Prov. Darlen . P. H. Allen

91^*, 5 Oct. 1938, vicinity of Boca de Cupe (GH, MO). J.

A. Duke 5011, k July I962, km. 16 de Yavlza along Q. Uvltal off Rio Chucunaque (MO). Prov. Panama . P. H. Allen 26O3,

5 July 1941, Pedro Gonzales, Perlas Islands, 0-20 m. (MO).

C. 0. Erlanson 597, 10 Aug. 19^5. stream bank along coast n. of East Harbor, San Jose Island, Pearl Archipelago (USX

I. M. Johnston 1101, 9 Jan. 19'*6, Old Mortar Road, east end, San Jose Island, Perlas Archlplelago, Gulf of Panama

(ca. 55 ml. s.s.e. of Balboa) (GH). Bro. Paul 219, Dec,

1932, Sabanas, R. P. n.e. Panama City (US). H. Plttler

6723. 10 Aug. 191if, Agriculture Exp. Station at Matlas

Hernandez (US). P. C. Standley 26665, 11 Deo. 1923, wet I

Map 36. Distribution of Helloonla latispatha . 290 291

forest, near big swamp e, of Rio Tecumen (US). P. C.

Standley 28266. ? Dec. 1923 - 11 Jan. 1924, Rio Tapla (US).

WEST INDIES. CUBA. Prov. Santa Clara . J. G. Jack 8287,

12 Aug. 1931, plants from Harvard Tropical Garden, Soledad,

Clenfuegos (GH, NY).

JAMAICA. Wm. Harris IIOO6. 24 Nov. I911. Castleton. Public Gardens (NY, US).

MARTINIQUE. P. Duss 2122, 1882, Introduced In garden (NY).

37. HELICONIA TORTUOSA Griggs, Bull. Torr. Bot. Club

3O1 650. 1903. Blhal tortuosa (Griggs) Griggs, Bull.

Torr. Bot. Club 3I: 445. 1904. Type: Guatemala, near

Flnca Sepaculte. Alta Verapaz, 17-18 March I9O2. 0. F.

Cook and R. F. Griggs s. n. (US).

Moderately stout plant, 1^-3^ m. tall; musold habit. '

Leaf-blade oblong-ovate, 0. 5-1.0 m. long, I8-3O cm. wide, short acuminate apex, rounded to attenuate-oblong base; upper and lower surfaces green, midrib on lower surface may have coarse matted brown hairs; petiole at least 35 cm. long, often tomentose. Inflorescence erect, slightly pubescent, about 40 cm. long; peduncle long and stiff, short vlllous-tomentose with golden-brown hairs; rachls flexuose O.5-O.6 cm. diameter, with same pubescence as on peduncle. Branch-bracts 5-11, red to brilliant scarlet, villous pubescent, often only on underside, subovate-trl- angular. moderately deep boat-shaped, not tapering; 292

branch-bracts arranged In spiral fashion; lowest branch

-bract, usxially fertile, extended Into leaf-blade often as long as leaf-blade of upper leaves; lowest branch-bract minus leaf-blade extension 12.5-16.0 cm. long; all branch

-bracts 2.0-3,0 cm. side-width; Intemode between lower branch-bracts 2-3 cm. Floral-bracts several, lanceolate,

4.5 cm. long, 0.8 cm. wide, about two-thirds as long as flowers; slightly puberulent on outer surface, especially along midrib. Flowers 5-6 per branch-bract, yellow-green,

5.0-7.0 cm. long; perianth 4.2-5.2 cm. long, projecting above edge of branch-bract, calyx puberulent, corolla glabrous; pedicel about 1 cm. long, glabrous or nearly so. Fruit dark, subglobose, 0.? cm. diameter. (Plate 37.) In bottom of wet ravines, moist to wet forests, and moist limestone thickets; growing at various altitudes from about 100 m. up to about I5OO m. The range extends from British Honduras to Costa Rica. (Map 37.)

There Is no taxonomlc confusion Involved with H. tortuosa . It was formerly Included In H. vlllosa Kl.. a much more villous species from Columbia, with more numerous branch-bracts and sessile flowers,

H. tortuosa Is named from the Latin "tortu" which means winding or twisting. It Is one of the few species in which the red bracts spiral. 293

'•K III Kit »K]1 M

it

UN T-O STATts NAT ON*l HfRBAR'UM

PIATE 37. HELICONIA TORTUOSA Griggs Holotype: H. tortuosa Griggs, near Finca Sepacuite. Alta Coorand p!'" Griggs'l? (Ss)f^^ °- r! »

29*

This species seems to be somewhat related to H. Irrasa.

The latter Is much more pubescent, with yellow to red branch-bracts, and It Is restricted to Panama. Like H.

Irrasa , H. tortuosa Is more pubescent In the early growth stages than it Is after the Inflorescence matures.

Representative specimens

BRITISH HONDURAS. Dlst. unknown . W. A. Schlpp 8-730.

10 June 193^, forest shade. Camp 35 BE - G survey, alt.

2^100 ft. (F).

COSTA RICA. Prov. of Alajuela . A. M. Brenes 13697a, 25

March 1931. La Palma de San Ramon (NY). A. Smith HU68, s. e,, region of Zarcero (P). Prov. Cartage . P. C.

Standley and R. Torres 51711, 6-7 March 1926, moist forest.

El Muneco, on the Rio Navarro, alt. 1400-1500 m. (US).

P. C. Standley 35'+97, Feb. 192^+, wet forest, Cerro de La

Carplntera, alt. I5OO-I85O m. (US). P. C. Standley 399^*5,

30 March 192^^. moist forest, vicinity of Orosl (US). Prov. Heredla . V. I. Sullivan 362, 18 Aug. I965, Puerto Vlejo (FSU). Prov. San Jose . A. P. Skutch 2685, July

1936, In deep shade, vicinity of El General, alt. 975 m.

(MICH, NY, US). P. C. Standley 41756a, 14-26 Dec. I925, moist forest, vicinity of Santa Maria de Dota, alt. I5OO -1800 m. (US).

GUATEMALA. Dept. Alta Verapaz . A. Molina R. and A. R.

Molina I2O87, 11 Mayo I963, en el matarral humedo entre San Pedro Carcha y Sacoyou, alt. I3OO m. (F). A. Molina

R., and A. R. Molina 12273. 1^ Mayo I963, margenes del Rio

Prlo entre Tactic y Santa Cruz, alt. l'4-60 m. (F). J. D.

Smith I828, M. April I889. Pansamala, alt. 38OO ft. (US).

P. C. Standley 70066, 2 April 1939, In damp limestone thickets, Saqulja, ^^3 km* n.e. of Coban, alt. ca. 1200 m.

(F). P. C. Standley 70299. 2 April 1939. damp forest, region of Cocola, n.e. of Carcha, alt. ca. 1200 m. (F).

P. C. Standley 92278, 1** April 19^^-1. wet forested ravine, between Santa Cruz and Tactic, alt. 1375 m. (F). H. von

Tuerckhelm 876I, M. June 190*4-, Coban, alt. 800 m. (US).

Dept. Baja Verapaz . H. von Tuerckhelm II I76O, April

1907. Panreal (US). Dept. Chlqulmula . J. A. Steyermark

31570, 10 Nov. 1939. Cerro Tlxlxl (Tlshlshl), 3-5 ml. n. of Jocotan, alt. 5OO-I5OO m. (F). Dept. Huehuetenango .

J. A. Steyermark U8713, I5-I6 July 19^2, In wet woods about 2 ml. s.e. of Maxbal, ca. I7 ml. n. of Barillas,

Sierra de los Cuchumatanes, alt. I5OO m. (F).

HONDURAS. Dept. Atlantlda . T. G. Yuncker '^7^8 a-b, 2k

July 193'+. In wet forest above Lancetllla, alt. 5OO ft.

(F, MICH, MO). T. G. Yuncker ^+959-3, 7 Aug. 193**. In low places near Tela River, vicinity of Lancetllla, alt. 90 ft. (P, MICH, MO). Dept. Conayagua . A. Molina R. 7O95.

22 Mayo 1956, bosque de plno - llquldambar de Montana La

Choca, Cordillera Comayagua, cerca de Coyocutena, alt.

1200 m. (F). T. G. Yuncker, R. P. Dawson, and H. H. Youse 58^1, 13 July 1936, bottom of wet ravine near El Achote;

hills above the plains of Slquatepeque, alt. I35O m. (P,

GH, MICH, MO). Dept. Cortes . A. Molina R. 11^129, 26

Marzo I963, bosque mlxto y lluvloso Montana San Idalfonso

entre Banaderos y Cusuco, alt. IkOQ m. (P). Dept. El

Paralso . A. Molina R. IklkJ, 12 Jimlo 196^*. Matorrales

cenagosos del bosque mlxto Sierra El Chile entre El Junqulllo El y Robledal, alt. I30O m. (P). Dept. Morazan .

A. Molina R. 15237, 23 Sept. I965, cultlvada en ml Jardln

compus de la EA.P. El Zamorano, alt. 800 m. (P), NICARAGUA. Dept. Jlnotega . P. C. Standley 1072^, 3 July

19^7, region of Las Mercedes, sierra e. of Jlnotega,

chiefly In dense wet mixed virgin forest (cloud forest), alt. ca. 1200-1500 m. (P). Dept. Madrlz . L. 0. Williams

and A. Molina R. 2027^*, 12 Jan. I967, In forest, cut-over

cloud forest area on Volcan Somoto. ca. 10 km. s. of Somoto alt. 1400 m. (P). Dept. Matagalpa . L. 0. Williams. A.

Molina R. and T. P. Williams 24827, Feb. I963, In moist swale In forest, along road to La Pundadora, cloud forest area n. of Sta. Maria de Ostuma, Cordillera Central de

Nicaragua, alt. I3OO-I5OO m. (P). Map 37« Distribution of Hell con la tortuosa.

299

38. HELICONIA IRRASA Lane ex R. R. Smith, sp. nov.

Typet Panama, Prov. Code, El Valle de Anton, North Hills,

June 29, l9'+6, P. H. Allen 35^7 (MO). Isotypes (P, GH, US),

Moderately stout plant, 1.5-2.0 m. tall; musold habit.

Leaf-blade oblong to oblong-ovate, 35-71 cm. long, 7-15 cm.

wide; acuminate apex, obtuse base; upper surface green,

lower surface slightly paler; petiole prominent, about U8

cm. long, woolly around sheath. Inflorescence erect, some-

times nodding, villous, 25 cm, long; peduncle nearly woollyj

rachls slightly flexuose and spiral; vlllous-woolly, O.^l

-0.5 cm. diameter (the latter more common), moderately

vlllous-woolly, ovate-lancGolate with acuminate apex

clrclnately curved, shallow boat-shaped; lowest branch43ract

often extends Into leaf-blade, often sterile, total length

16-60 cm. long; lowest fertile branch-bract 10-1? cm. long; middle branch-bracts 8-12 cm. long; all branch-bracts I.5

-2.2 cm. side-width; lower branch-bracts separated by

Internodes of 2-3 cm. Floral-bracts wide-lanceolate, U cm. long, 0.8 cm. wide, membraneous, puberulent to slightly pubescent. Flowers about 6 per branch-bract, yellow; 5-6 cm. long, perianth if.0-5.2 cm. long, nearly glabrous to slightly tomentose; pedicel 0.5 cm. long, slightly tomen- tose; ovary smooth. Fruit dark, subglobose. O.7 cm. diameter. Seed not seen. (Plate 38.) PLATE 38. HELICONIA IRRASA Lane ex R. R. Smith

Holotype: H. Irrasa Lane ex R. R. Smith, El Valle de Anton, Prov. Code, Panama, 29 Jvme 19^6, P. H. Allen 35^7 (MO). »

Foxmd In woods and forests In Panama. This species

Is endemic to Panama. There are no South American species

of Helicon la that fit this description. (Map 38.)

The species name of H. Irrasa comes from the Latin

"Inrasa" which means unshaved. As the name Implies, this

species Is very vlllous-woolly and easily recognizable.

There are two variations In color; red, and yellow with a red border.

H. Irrasa has also been called H. tortuosa and H.

vlllosa . H. tortuosa Is geographically distributed In northern Central America. The species has red branch

-bracts, which are not vlllous-woolly, but possess

scattered short hairs on the branch-bracts . H. vlllosa

Kl. may be confused with H. Irrasa since both are villous

-woolly. However, H. vlllosa Is of shorter stature, has

more and shorter branch-bracts, sessile flowers, and Is

pendent. H. vlllosa Is a South American species.

Representative specimens

PANAMA. Canal Zone . S. Avlles 38, 1931. Barro Colorado

Island (P). J. A. Duke 4if26, 6 Oct. I96I, Camp Plna,

vicinity of hill C-6, Fort Sherman (MO). J. E. Eblnger

^72, 17 July i960, Fort San Lorenzo (MO). D. Falrchlld

7. Aug, 1924, Barro Colorado Island (F). G. Hager 363,

Nov. I859, In woods near Gatim Sta. P. R. R. (NY). D.

E. Starry 5I, July 1931. Armour Trail, Barro Colorado 302

Island (P). R. E. Woodson, Jr. and R. W. Schery 983, 3-20

Aug. 19'*0, Barro Colorado Island (MO). Prov. Code . P.

H. Allen 216?, 23 June 19^0, hills n. of El Valle, alt.

1000 m. (GH, MO, NY, US). P. H. Allen 2490, 11 Kay 19^1. hills n. of El Valle de Anton, vicinity of La Mesa, alt.

1000 m. (US). P. H. Allen 35^7. 29 June 19'+6, North

Hills, El Valle de Anton (P, GH, MO, US). Prov. Colon .

P. H. Allen 3582, 11 July 19^6, vicinity of Camp Plna, alt. 25 m. (P, GH, US). J. E. Eblnger 10k, 20 Jtine I96O,

Portobello, Las Cruces Trail (MO). R. J. Selbert 1520,

9 Aug. 19^0. Gatun Lake, Goodyear, Allweather Estate (MO).

Prov. Panama . K. E. Blum, S. Olson and R. Rasmussen 2377,

19 June 1966, Cerro Campana, El. ca. 2300 ft. (PSU, MO). Map 38. Distribution of Hell con la Irrasa.

HYBRIDIZATION

During this taxonomlo study no attempts were made to

hybridize Hellconla. Of the flfty-slx clones representing

approximately twenty species maintained under greenhouse

culture as part of the present study, only four flowered.

The factors restricting flowering were not determined, but

It Is hoped that a continuation of this aspect of the study

will permit hybridization attempts to be conducted In the future.

It has long been thought that species of Hellconla do

hybridize. This conclusion has been set forth verbally by

Various collectors of tropical plants; It Is also the con-

clusion of the author. Upon establishing the morphological

entitles through studying and measuring the herbarium

specimens, and comparing the findings with the literature.

It was discovered that there are certain plants which are

Intermediate In many ways. These plants are within the

range of the putative parents, a general requirement of hybridization.

One curious factor Is that several of these hybrids are found near the edge of the distribution of the Involved species. The habitat of the putative parents appears to be restricted, with a small number of plants of both species

305 in a limited area, there Is a greater chance that cross

-pollination may occur.

In hybridization the mode of pollination must be

considered. It Is known, as stated previously, that

Helicon la is pollinated by hummingbirds. How selective

they are as pollinators is yet to be determined.

The following specimens are considered to be hybrids:

1. HELICONIA X DRESSLERIANA R. R. Smith, hybr. nov.

Helicon la latispatha x H. vaginalis

Type: Mexico, State Chiapas, near Laguna Ocotal Grande,

ca. 25-30 km. s.e. of Monte (Cerro) Llbano (which is ca.

45 km. e. of Ocaslngo), elev. ca. 950 m., 20 July - 20

Aug. 1954. R. L. Dressier lif89 (US). Isotype (GH).

Moderately stout plant, 1.0-2.5 m. tall; musold habit.

Leaf-blades oblong, I2-65 cm. long, 6-20 cm. wide, acumi-

nate apex, ro\inded base; upper and lower surfaces green;

petiole prominent, 6-3O cm. long, slender, about O.3 cm.

diameter. Inflorescence erect, slightly tomentose to

glabrous, 10-25 cm. long, pedvmcled; rachls slightly to

moderately flexuose and spiral, O.3 cm. diameter. Branch

-bracts 4-6, red to red-orange, glabrous, lanceolate,

shallow boat-shaped; lowest branch-bract often extending into short leaf-blade, total length I3-I6 cm; middle branch-bracts 8-10 cm. long; upper branch-bracts 4-6 cm. long; all branch-bracts 1.0-1. 5 cm. side-wldth> Intemodes i

307

between branch-bracts 2.0-3.5 cm. Floral-bracts wide

-lanceolate, deciduous, 3«5 cm. long, 1 cm. wide, glabrous.

Flowers 4-5 per branch-bract, yellow, k,0-k,5 cm. long,

glabrous; pedicel 0.5-0. 7 cm. long. Fruit dark, subglobose^

0.6 cm. diameter. Seed not seen. (Plate 39.)

These plants show the obvious characteristics of the

two species Involved. The characteristics of H. latlspatha are the presence of pronounced petioles, the short pedicels and spiral rachls. The curved, upright, red to red-orange

branch-bracts, which upon maturing become more horizontal, are characteristic of H. vaginalis . This author observed

four separate collections of this hybrid; one from Mexico,

one from Costa Rica and two from Panama, and all four showed the above characteristics.

Representative specimens

COSTA RICA. Prov. San Jose . D. E. Stone 21k9, 29 March

1965, road from San Isldro del General to Cartago, ca.

15.1 ml. s. of Villa Mills (DUKE).

PANAMA. Prov. Bocas del Toro . H. von Wedel 2389, k May 19^1, foothills and lowlands. Fish Creek (GH). Prov .

Panama. S. McDanlel 6852, 28 Aug. I965, near creek.

Cerro Campana, alt. 29OO ft. (PSU). 308

PIATE 39. HELICONIA X DRESSLERIAM R. R. Smith, hybr. nov.

Holotypei H. x dresslerlana R. R, Smith, near Laguna Ocotal. State Chiapas, Mexico, 20 July - 20 Auk. R. L. Dressier 1489 (US). * 2. HELICONIA X GILLYANA R. R. Smith, hybr. nov.

Helicon la colllnslana X H. latlspatha

Type: Mexico, State Tabasco, Mun. Macuspanaj road along

Rio Chlnal (Rio Macuspana) between Macuspana and El Carmen,

29 Sept. 19^+^, C. L. GUly. Sr. and E. Hernandez X. 386

(MICH).

Leaf- blade oblong, 55 cm. long, 19 cm. wide, short acuminate apex, rounded base; upper and lower surfaces green; petiole 26 cm. long, moderately stout. Inflores-

cence erect, slightly puberulent to glabrous, 35 cm. long, peduncled; raohls nearly straight, 0.4-0.5 cm. diameter.

Branch-bracts 7, red, glabrous, lanceolate-ovate, subcor- date, shallow boat-shaped, extending out horizontally; lower branch-bracts slightly curved upward, 14-19 cm. long; middle and upper branch-bracts 7-11 cm. long; all branch

-bracts 2.2-1.5 cm. side-width; Intemodes between branch

-bracts from 2-3 cm. In length. Floral-bracts present.

Flowers 6-7 per branch-bract, yellow, 4-5 cm. long. Fruit not seen. (Plate 40.)

This plant possesses an erect Inflorescence and rather short pedicels, characteristics of H. latlspatha . The sub- cordate, red, branch- bracts are characteristics of H. colllnslana .

Only one specimen of this taxon was seen, and It was

In flower. Therefore, other characteristics such as fruit size could not be observed. 310

Holotype: H. X glllyana R, R, Smith, Mun. Mocuspana, State Tabasco, Mexico. 29 Sept. 19kk, C. L. Gilly, Dr. and E. Hernandez X. 386 (MICH), 9

311

3. HELICONIA X MOOREANA R. R. Smith, hybr. nov.

Heliconla coll Ins iana X H. schledeana

Type: Mexico, State Guerrero, steep sides of shady ravine

In pine woods near km. 339 on highway to Acapulco below

Acahulzotha, alt. 9^0 m., 31 March 1952, H. E. Moore, Jr.

6204 (GH).

Stout plant, about 2m. tall; musold habit. Leaf

-blade oblong-ovate, 1-2 m. long, 12-50 cm. wide, acuminate

apex, rounded base; upper surface green, lower surface

somewhat paler; petiole about 90 cm., stout. Inflorescence

pendulous, strongly hirsute with golden brown hairs, about

45 cm. long, peduncled; rachls crimson, moderately flexuose,

hirsute, 0.4 cm. dlamter. Branch-bracts 7t crimson,

slightly hirsute on bottom near base, narrowly linear with

obtuse apices, shallow boat-shaped, moderately reflexed;

lowest branch-bracts 13-16 cm. long; middle and upper

branch-bracts 7-11 cm. long; all branch-bracts 0.7-0.

cm. side-width; Intemode between branch-bracts gradating

from 3-5 cm. Floral-bracts many, lanceolate, 5.0-5. 5 cm.

long, hirsute-tomentose outer surface and glabrous Inner

surface. Flowers 6-7 per branch-bract, orange-yellow,

6 cm. long; perianth 5 cm. long, hirsute-tomentose calyx,

glabrous corolla; pedicel 1.0-1. 5 cm. long, strongly

hirsute. Fruits dark, subglobose to 3-slded, O.9 cm.

long, 1.0 cm. wide. (Plate 41.) PIATE 41. HELICONIA X MOOREANA R. R. Smith, hybr. nov.

Holotype: H. X mooreana R. R, Smith, below Acahuizotla, State Guerrero, Mexico, 31 March 1952, H, Moore, Jr. 6204 (GH). 313

Found In shady moist ravines at altitudes from about

50-950 meters.

The Inflorescences of these plants are pendulous,

with moderately subcordate branch-bracts and moderately

long pedicels, characteristics of H. coll Ins lana . The

Inflorescence possesses a pubescent rachls with narrow,

somewhat elongated branch-bracts, which are typical of

H. schledeana.

Representative specimens:

MEXICO. State Guerrero . E. Palmer 311, Oct. 189^+ - Mar.

I895. Acapulco and vicinity (GH). State Nayarlt . R.

McVaugh 121^3. 22 April 1951 1 about if ml. e. of Jalcocotan,

on road to Teplc (MICH).

4. HELICONU X PROCERA R. R. Smith, hybr. nov.

Hellconla lankesterl X H. vaginalis

Type I Costa Rica, Prov. Alajuela, near La Lagvina, 6 to

8 km. s. of Villa Quesada, alt. 1000 m., I9 Feb. I966,

A. Molina R., L. 0. Williams, W. Burger and B. Wallenta

17^^77 (F).

Somewhat slender plant 3-4 m. tall; cannold habit.

Leaf-blade oblong, 22-40 cm. long, 9-11 cm. wide, acuminate apex, acximlnate or rounded base; leaf-blade sessile. In-

florescence erect, 14-18 cm. long, glabrous to slightly

pubescent, peduncled or not; rachls flexuose, 0.6 cm. i

31^^

diameter. Branch-bracts 6-7, red, glabrous to slightly hlrsute-tomentose, linear- lanceolate, shallow boat-shaped, extending out horizontally or curved upward; lowest branch

-bracts 10-20 cm. long; middle branch-bracts 8-10 cm. long; uppermost branch-bracts 5«0-5»5 cm. long; all branch-bracts about 1.0 cm. side-width. Floral bracts 3.5-4.0 cm. long,

0.5-1,0 cm, wide, glabrous. Flowers 12 to about 25 per branch-bract, yellow, 5 •5-6,0 cm, long, glabrous to nearly so; perianth 4,5 cm. long; pedicel 1-2 cm. long, slightly hirsute to glabrous. Fruit dark, subglobose to 3-slded, about 1 cm. diameter. (Plate 42.)

The habitat of these plants Is moist forest at altitudes of approximately 1000 m.

The leaf-blades of this hybrid are sessile to the sheaths, a characteristic of H. vaginalis . These plants generally are more robust and taller than H, vaginalis .

The branch-bracts are more cymblform, characteristic of

H, lankesterl . The branch-bracts and the rest of the

Inflorescence are Intermediate between the two species.

Representative specimen

COSTA RICA. Prov. Puntarenas . A. Jimenez M. 2456, 22

Oct. 1964, Entre Agua Buena y San Vlto de Jaba, alt.

1200 m. (P). ; 315

rr«ct;rUk vtth '> - 7 tmttte about 10 cite, or lo«|ti cueuUat*, with unj flow«ra in tl^ UL«. r«d| 1««VMI 30-l»0 CM. lour, l«nc#olnte. PIATE k2, HELICONIA X PROCERA R. R. Smith, hybr. nov.

Holotype: H. X procei^ R, R. Smith, near La Laguna, Prov. Alajuela, Costa Rica, 19 Feb. I967, A. Molina R. et al. 17*^77 (P). 316

5. HELICONIA X WOODSONIANA R. R. Smith, hybr. nov.

Hellconla hlrsuta X H. vaginalis

Type I Panama, Prov. Bocas del Toro, Rio Crlcamola, between

Finca St. Louis and Konklntoe, ca. 10-50 m., 12-16 Aug.

1938, R. E. Woodson, Jr., P. H. Allen, R. J. Selbert 1931

(MO). '

Slender plant, i.0-1.5 m. tall; cannold habit. Leaf

-blades elliptic, 30-50 cm. long, 8-12 cm. wide, acuminate apex and base; upper and lower surfaces green; petiole

prominent, 9-1^ cm. on lower leaves, upper leaves nearly

sessile. Inflorescence erect, slightly tomentose to

glabrous, on long slender peduncles, 0.2-0. 3 cm. diameter.

Branch-bracts ^l-, red to red-orange, glabrous, linear-lance-

olate; lowest branch-bract extending Into small leaf-blade,

total length 12-16 cm.; middle to uppermost branch-bract

5-10 cm. In length; all branch-bracts 0.5-1.0 cm. side

-width. Floral-bracts lanceolate, deciduous, about 3 cm.

long. Flowers 2-3 per branch-bract, yellow, 4.5 cm. long;

perianth 3 •6-3 "8 cm. long, glabrous; pedicel about 0.7 cm.

long, somewhat tomentose. Fruit not seen. (Plate 43»)

This hybrid possesses moderately short, extremely

narrow-lanceolate branch- bracts . The branch-bracts are

similar to those of H. hlrsuta with the exception that

they are red or red-orange and glabrous. The lower leaves

also possess a petiole which Is similar to leaves of H. 317

PIATE 1^3. HELICONIA X WOODSIANA. R. R. Smith, hybr. nov.

Holotype: H. X woodsonlana R. R. Smith, Hlo Cricamola, Prov. Bocas del Toro, Panama, 12-16 Aug, 1938, R, E. Woodson et al, 1931 (MO). hlrsuta. The leaf shape and size, branch-bract color, and flower color are more like that of H. vaginalis .

Representative specimen!

PANAMA. Prov. Panama , P. H. Allen 2216, 1 Sept. 19'4-0, summit of Cerro Campana, 800-1000 m. (GH).

6. HELICONIA X ZACAPAEA R. R. Smith, hybr. nov.

Hellconla coll Ins lana X H. adflexa

Type J Guatemala, Dept. Zacapa, pine-covered canyon border- ing Rio Lima, Sierra de las Mlnas, below Flnca Alexandria, alt. 2000 m., Ik Oct. 1939, J. A. Steyermark 3OO38 (P).

Stout plant, 2-3 m. tall; musold habit. Leaf-blades oblong, 100-130 cm. long, 25-30 cm. wide, short acuminate apex, rounded to acuminate base; upper and lower surfaces green; petiole 1 cm. diameter. Inflorescence pendulous, glabrous, 30 cm. long; peduncle 0.1 cm. diameter; rachls thick, glabrous, 0.9 cm. diameter. Branch-bracts I3, rose

-red, glabrous, linear- lanceolate, shallow boat-shaped, curved upward; lowest branch-bract about 16 cm. long; middle and upper branch-bracts 5-10 cm. long; all branch

-bracts about 1 cm. side-width. Floral-bracts deciduous; not seen. Flowers 10-12 per branch-bract; perianth not seen; pedicel 2.0-2.5 cm. long, glabrous. Fruit large, subglobose, I.5 cm. long, 1 cm. wide. (Plate kk,) PIATE 44. HELICONIA X ZACAPAEA R. R. Smith, hybr. nov. ?^<^apaea nfi?*^?®' ^ R. R. Smith. Sierra de las Minas. ^^^^'^^^^ 1939, J. A. Steyermark 30038 (F)!^' 320

This plant was growing along the wet banks of the Rio Lima, Guatemala.

The Inflorescence Is relatively short with branch

-bracts positioned like those of H. adflexa . It Is pendent with extremely long pedicels and large fruits, as found In H. coll Ins lana .

Only a single plant of this hybrid was seen. »

APPENDIX I

New Names and Combinations

Hellconla catheta R. R. Smith

var * catheta

var. pubescens R. R. Smith

Hellconla colgantea R. R. Smith

Hellconla coll Ins lana Griggs

var. velutlna Lane ex R. R. Smith

Hellconla hlrsuta L. f

var. rublflora R, R. Smith

Hellconla Irrasa Lane ex R. R. Smith

Hellconla leucoflora R. R. Smith

Hellconla llnnaeana Lane ex R. R. Smith

Hellconla longa (Griggs) Wlnkl.

var. mult 1 flora R. R. Smith

Hellconla magnlf lea Morton ex R. R. Smith

Hellconla mollnaeana R. R. Smith

Hellconla paulll Lane ex R. R. Smith Hellconla schledeana Klotzsch

var. splssa (Griggs) R. R. Smith

Hellconla trlnldatls Lane ex R. R. Smith

Hellconla Veracruz 11 R. R, Smith

Hellconla xanthotrlcha R. R. Smith

321 . •

APPENDIX II

Doubtful Species and Rejected Names

Helicon la flora Jacq., Hort. Schoen. Ii 25 • 1797

There Is no description nor Illustration for this name

Helicon la marantl folia G. Shaw, Clmel Phys. 7^t 7« 38. 1796*

The description and Illustration are taxonomlcally obscure. The Illustration shows a likeness to Hellconla aurantlaca . The location, St. Vincent Island, Is not correct for this species or any other resembling the accompanying Illustration.

Hellconla rubra Sesse and Moc, PI. Mex. Ed. 2. p. 65-66.

189^*.

Hellconla pslttacorum Sesse and Moc, Fl. Mex. Ed. 2. p.

65-66. 189*^.

Hellconla hlrsuta Sesse and Moc, Fl. Mex. Ed. 2. p. 66.

1894.

There are two specific entitles under the names of

H. rubra and H. pslttacorum , and one under the name H. hlrsuta . The descriptions are not clear, and the herbarlvim specimens. If they exist, would have to be studied to determine the specific synonym.

322 323

The following tentative deductions are made from the

descriptions J

1) Rubra p. 65 " Hellconla vaginalis Benth.

2) Pslttacorum p. 65 = Hellconla aurantlaca Ghelsbr.

3) Pslttacorum p. 66 = ?

^) Hlrsuta p. 66 = Hellconla schledeana Kltz.

5) Rubra p. 66 = Hellconla bourgaeana 0. G. Peters.

Hellconla sanderl Sand. In Gard. Chron, 1. 365. I899.

This Is merely a brief description of vegetative

structures.

Hellconla purpurea Griggs, Bull. Torr. Bot. Club Vol, 30j

656. 1903.

This Is the name that Griggs gives to a H. blhal found

Illustrated In Andrew's Repository, pi. 6kO, 1812. A

similar specimen Is yet to be found In this present study. The bracts are said to be purple with yellow margins. A

study of South American species may be more revealing in

reference to this taxon.

Hellconla dlstans Griggs Bull. Torr. Bot. Club Vol. 3O:

657. 1903.

This species name Is proposed for the Illustration,

H. blhal Sw., (Petersen, I89O). The taxon Involved is

South American. Therefore, this name will merit further

consideration upon the study of South American species of Hellconla. It Is suspected that Hellcon la dlstans Is a synonym for Hellconla latlspatha * APPENDIX III

Lists of Species by Country

British Honduras Costa Rica

H. aurantlaca H. aurantlaca

H. bourgaeana H. colgantea

H. coll Ins lana H. coll Ins lana

H. hlrsuta var. rubl flora .H. Imbrlcata

H. latlspatha H. lankesterl

H. llbrata H. latlspatha

H. marlae H. leucoflora

H. schledeana var. splssa H. longa

H. vaginalis H. marlae

H. osaensls

H» reticulata

H. tortuosa

H. vaginalis

H. vellerlgera

H. wagnerlauia

El Salvador

H. coll Ins lana

H. latlspatha

325 326

Guatemala Honduras

H. adflexa H. aurantiaca

H. aurantlaca H. bourgaeana

H» bourgaeana H. coll ins iana

H. coll Ins lana H. latispatha

H. latlspatha H. libra ta

H. librata , H. mariae

H. marlae H. molinaeana g.. schledeana var. splssa H. schiedeana var. spissa

H. tortuosa H . tortuosa

H. vaginalis H. vaginalis

H. wagneriana H. wagneriana

Mexico

H . adflexa

H_. aurantiaca

H. bourgaeana

JH • coll ins iana

H. latispatha g.. libra ta

S.. schiedeana var. schledeana

H. schiedeana var. spissa

H. vaginalis

H. veracruzii ,

Nlcaragxm Panama

H. aurantlaca H. allenll

H. coll Ins lana H. catheta

H. hlrsuta var. rubl flora H, curtlspatha

H, latispatha H. hlrsuta var. hlrsuta

H. leucoflora H, Imbrlcata

H. librata H. Irrasa

H. longa H • lankesterl

H» osaensls H. latispatha

H. tortuosa H. leucoflora

H. vaginalis H. magnlf lea

H. wagnerlana H. marglnata ,,

H. marlae West Indies H. metalllca

H. blhal H . nutans

H. carlbaea H. osaensls

H. Indlca (cult.) H. Paul 11

.' H. latispatha (cult.) H. vaginalis .

H. llnnaeana H. vellerlgera

H. pslttaconm H. wagnerlana

H. trlnldatls H. xanthotrlcha

H. swart z lana . .

APPENDIX IV

A Tentative List of South American Species That Do Not Occur In Central America

Type

H. af finis Loes. Peru

H. angusta Veil. Brazil

H. angustlfolla Hook, f Brazil

H. aureo-roseo Loesn. Peru

H. braslllensls Hook f Brazil var. concolor Peters. Island of Cayenne, ' French Guiana

H. burchelll Baker Brazil

H. cannoldea L. C. Rich.

H. Cardenas 11 L. B. Smith Bolivia

H. costanensls Arlsteg. Venezuela

H. dasyantha C. Koch et Bouche Bras 11, Guiema var. rosea Loes. in Dials. Ecuador

- H. denslflora Verlot Guiana ,

H. dlelslana Loes. In Dlels Ecuador *

H. eplscopalls Veil. Brazil

H. senlculata (Griggs) L. B. Smith Colombia

H. glauca Polt. ex Verlot British Guiana H. grlggslana L. B. Smith Colombia

H. humllls Jacq. Venezuela and Guiana

328 329

H. Juriiana Loes. Brazil

H. llngulata Ruiz et Pav. Peru

H. mutlslana Cuatr. Colombia

H. nana G. Rodriguez Venezuela

H. pearcel Rusby Bolivia

H. pendula Wawra Brazil

H. penduloldes Loes. ' Peru

H. poepplglana Elchl. Peru

H. prulnosa Loes. Peru

H. pulverulenta Llndl. Brazil? •

H. revoluta (Griggs) Standley Colombia

H. robusta Pax. Bolivia

H. rodrlguensls Arlsteguleta Venezuela

H. rosea- flava Loes. Peru

H. rostrata Ruiz et Pav. Peru

H. schaeferlana G. Rodriguez Venezuela

H. schneana Steyermark Venezuela

H. schomburgklana Kl. Guiana

H. schumannlana Loes. Peru and Brazil var. acreana Loes. Brazil var. baslrubra Loes. Peru

H. spatho-clrclnada Arlsteguleta Venezuela

H. standleyl Macbr. ' Peru

H. steyermarkll Arlsteg. Venezuela

H. stricta Ruber. Peru

H. subulata Ruiz et Pav. Peru

H. sylvestrls (Gleason) L. B. Smith British Guiana 330

H. uleana Loes, Peru

H. varlegata Loes. Peru

H. vlllosa KL. Colombia

H. weberbauerl Loes. Peru APPENDIX V

Presentation of Form for Field Study

The following form Is suggested to be used when one

Is doing a field study of the genus Hellconla. The major- ity of the species are much too large to be studied thoroughly from herbarium sheets. In the process of drying, many of the characteristics are lost. Therefore, good field data are essential In order to understand fully the entitles.

V/hen pressing a specimen of this genus, the following suggestions might be considered:

1) As the Inflorescence Is being pressed, a mature

branch-bract should be split open so that the

Inner surface Is exposed.

2) A few flowers and floral-bracts should be pressed separately.

3) The color should be stated on the label for the

branch-bracts, rachls, floral-bracts, and flowers.

The color of the fruit and leaves should also be stated when pertinent.

^) A second sheet, and possibly a third, should be

utilized for the pressing of a mature leaf. As the leaf Is pressed, the shape of the base and

331 332

the apex should be made readily evident along

with the upper and lower surfaces of the leaf*

If a petiole Is present, a portion of It attached

to the base of the leaf-blade should be pressed.

5) If possible, a few mature flowers should be pre-

served In a liquid preservative, so that the

anthers, and especially the stamlnode, might

be studied at a later date.

I i

Form for field notes when collecting Helicon la

Name of colleotort Exact location t Collection number

Species

Plant height Diameter of pseudostem at base

LEAVES Number of leaves on stem

Length of longest leaf-blade

width of longest leaf-blade

Length of shortest leaf-blade

width of shortest leaf-blade

Shape of leaf-blade apex

Shape of leaf-blade base

Comments on lower surface

Comments on upper surface

Petiole length of lower leaves

Petiole diameter of lower leaves

Petiole length of upper leaves

Petiole diameter of upper leaves

Coloration markings

General remarks on vegatatlve structures i (margins, leaf angle from stem, etc.) ) )

33k

INFLORESCENCE Pendulous or erect

Length (lowest branch-bract to top of Inflorescence)

Length of peduncle

Width of peduncle just below lowest branch-bract

Pubescence of pedvincle (If any

Color or peduncle

Diameter of rachls above lowest branch-bract

Rachls straight or flexuose

Bachls spiralled or not

Rachls color *

Rachls pubescence (If any

A) BRANCH-BRACTS Number of branch-bracts

Arrangement of branch-bracts (distichous, three-ranked, or

other )____

Lowest branch-bracts J Length

width across top Color of outer surface

Color of Inner surface

Shape of branch- bract

Pubescence (If any) of outer surface

Pubescence (If any) of Inner surface

Remarks! (leaf-extension, sterile or fertile, etc.) 335

Lower fertile branch-bracts i Length

width across top

Color of outer surface

Color of Inner surface

Shape of branch-bract

Pubescence (If any) of outer surface

Pubescence (If any) of Inner surface

Middle branch-bracts t Length_

width across top Shape of branch-bract

Upper branch-bracts t Length

width across top

Shape of branch-bract

Distance between lower branch-bracts

Distance between middle branch-bracts

Distance between upper branch-bracts

B) FLORAL-BRACTS Number per branch-bract

Color Shape

Length Width

Pubescence (If any)

Deciduous or persistent t i

336

i

C) FLOWER

Number per branch-bract

Col or

Length of whole flower

Length of perianth

Pubescence of perianth

Sepalsi (free sepal curled at anthesls, etc.)

Color length

Pubescence

Remarks on petals

Pedicel length pedicel color

Pubescence on pedicel (If any)

General remarks on flower structure

D) FRUIT

Color Shape

Size

Pubescence (If any)

Other remarks I ^ ' E) SEED '

Color Shape

Slze__

Other remarks!

ECOLOGY

1) Altitude

2) Area type (disturbed- pasture, forest, other) (primary or secondary)

3) Sunlight (full, degrees of shade)

k) Slope

5) Drainage

6) Surface characteristics (rocky, smooth, etc.)

7) Soil type (sand, silt, clay)

8) Pollination I

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338 j i

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340

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WOODSON, Jr., R. E. and SCHERY, R. W. 19^*5. Flora of \ the^Missourl Botanical Panama. Annals of j Garden, XXXII. part 3» ^8-57. BIOGRAPHICAL SKETCH

Robert R. Smith was born September 11, 193^t at

Stamford, New York. In June, 1952, he was graduated from Stamford Central School. Prom 195**- until 1956 he served in the Medical Corps of the United States Army.

Prior to emd upon completion of his Army service, he attended the College of Wooster, Wooster, Ohio, and in

1958 he transferred to Hartwick College, Oneonta, New

York, where he received the degree of Bachelor of Arts in June, I96O. In I96O he enrolled in the Graduate School of the University of Florida, and received the degree of

Master of Science in June, I962. From I96O to I962 he worked as a graduate assistant in the Department of Botany, and in I963 he served as herbarium assistant. From the academic years of I963 to I966 he held a position of '

Instructor in Biology at Orlando Jionior College, Orlando,

Florida. In June, I966, he returned to the University of

Florida in order to complete the requirements for the • degree of Doctor of Philosophy.

He is a member of Hartwick College Honor Society,

Beta Beta Beta Honorary Fraternity, Phi Sigma, and an associate member of Sigma Xi.

He is married to the former Julia Mae Darling of

Orlando, Florida.

31*4 This dissertation was prepared under the direction of

the chairman of the candidate's supervisory committee and has been approved by all members of that committee. It was submitted to the Dean of the College of Agriculture and to the Graduate Council, and was approved as partial

fulfillment of the requirements for the degree of Doctor

of Philosophy.

March 1968 19. ^ .

Dean, Graduate School

Supervisory Committee:

Chairman