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Botany

O. 18

FLORA COSTARICENSIS

William Burger, Editor

Family #200 Acanthaceae

L. H. Durkee

Family #201 Plantaginaceae

William Burger

December 31. 1986

Publication 1372

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

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FLORA COSTARICENSIS

Family #200 Acanthaceae

Family #201 Plantaginaceae

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FIELDIANA

Botany

NEW SERIES, NO. 18

FLORA COSTARICENSIS

William Burger, Editor

Family #200 Acanthaceae

L. H. Durkee

Professor of Biology

Department of Biology

Grinnell College

Grinnell. Iowa 501 12

Family #201 Plantaginaceae

William Burger

Curator, Vascular Plants

Department of Botany

Field Museum of Natural History

Chicago, Illinois 60605-2496

Accepted for publication March 14, 1985

December 31, 1986

Publication 1372

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

Library of Congress Catalog Card Number: 78-172358

ISSN 00 15-0746

PRINTED IN THE UNITED STATES OF AMERICA

Table of Contents

List of Illustrations

ACKNOWLEDGMENTS vi

ACANTHACEAE by L. H. Durkee 1

Conspectus of the Subfamilies and Genera

in Costa Rica 2

Illustrations of Acanthaceae 3-20

Key to the Genera of Acanthaceae in Cos-

ta Rica 21

Acanthus 23

Aphelandra 23

Asystasia 30

Barleria 31

Blechum 32

Bravaisia 33

Buceragenia 33

Carlowrightia 34

Chaetochlamys 35

Chaetothylax 35

Chamaeranthemum 36

Dicliptera 37

Dyschoriste 40

Elytraria 41

Graptophyllum 41

Habracanthus 42

Hansteinia 43

Henrya 45

Herpetacanthus 46

Hygrophila 47

Hypoestes 47

Justicia 48

Louteridiwn 62

Megaskepasma 62

Mendoncia 63

Nelsonia 65

Odontonema 66

Poikilacanthus 67

Pseuderanthemum 67

Razisea 70

Ruellia 72

Sanchezia 81

Spathacanthus 81

Streblacanthus 82

Teliostachya 83

Tetramerium 83

Thunbergia 84

Trichanthera 86

PLANT AGINACEAE by William Burger .. 87

Plantago 87

Illustration of Plan taginaceae 89

INDEX . 91

1. Acanthus mollis; Aphelandra: A. auran-

tiaca var. aurantiaca; A. aurantiaca var.

stenophylla; A. campanensis; A. scabra\ A.

dolichantha 3

2. Aphelandra: A. golfodulcensis; A. leo-

nardii; A. lingua- bovis; A. seibertii; A. sin-

clairiana; A. storkii 4

3. Aphelandra: A. tonduzii; A. tridentata;

Asystasia gangetica; Barleria micans;

Blechum: B. costaricense; B. brownei ... 5

4. Bravaisia integerrima; Buceragenia glan-

dulosa; Chaetothylax leucanthus; Cha-

maeranthemum durandii; Dicliptera: D.

imbricata; D. iopus 6

5. Dicliptera: D. pallida; D. podocephala; D.

skutchii; D. trifurca; D. unguiculata; Dys-

choriste valeriana; Elytraria imbricata . . 1

6. Graptophyllum pictum; Habr acanthus sil-

vanticus; Hansteinia: H. blepharorhachis;

B. sessilifolia; H. stricta; H. ventricosa . . 8

7. Henrya scorpioides; Herpetacanthus pan-

amensis; Hygrophilia costata; Hypoestes

phyllostachya; Justicia: J. angustibractea-

ta; J. aurea 9

8. Justicia: J. brenesii; J. candelariae; J. car-

thaginensis; J. chamaephyton; J. comata;

J. costaricana 10

9. Justicia: J. crenata; J. brandegeana; J.

macrantha; J. metallica; J. oerstedii; J.

orosiensis 11

10. Justicia: J. parvibracteata; J. pectoralis; J.

pittieri; J. refractifolia; J. sarapiquensis; J.

skutchii 12

1 1 . Justicia: J. spicigera; J. tinctoria; J. ton-

duzii; J. trichotoma; J. urophylla; J. val-

erii 13

12. Louteridium costaricensis; Mendoncia: M.

brenesii; M. costaricana; M. lindavii; M.

retusa; M. tonduzii 14

1 3. Megaskepasma erythrochlamys; Nelsonia

canescens; Odontonema tubiforme; Po-

ikilacanthus macranthus; Pseuderanthe-

mum atropurpureum; Pseuderanthemum

cuspidatum 15

14. Pseuderanthemum: P. pittieri; P. praecox;

P. standleyi; Razisea: R. citrina; R. spi-

cata; R. voilburi 16

15. Ruellia: R. biolleyi; R. cooperi; R. gemi-

niflora; R. golfodulcensis; R. inundata; R.

jussieuoides 17

16. Ruellia: R. metallica; R. nudiflora; R. pa-

lustris; R. paniculata; R. pittieri; R. prae-

clara 18

17. Ruellia: R. stemonacanthoides; R. ton-

duzii; R. tubiflora var. hirsuta; R. tubiflora

var. tetrastichantha\ Spathacanthus hoff-

manni; Streblacanthus monospermus ... 19

18. Teliostachya alopecuroidea; Tetramer-

ium nervosum; Thunbergia alata; Tri-

chanthera gigantea 20

19. Plantago australis 89

Acknowledgments

This is the fifth issue of the Flora Costaricensis.

The first dealt with Piperaceae (Fieldiana, Bot. 35,

1 97 1). The second included families numbered 42

through 53, Chloranthaceae through Urticaceae

(Fieldiana, Bot. 40, 1977). The third issue con-

tained the Gramineae by Richard Pohl (Fieldiana,

Bot., new series, No. 4, 1980). The fourth issue

included families numbered 54 through 70, Podo-

stemaceae through Caryophyllaceae (Fieldiana,

Bot., new series, No. 13, 1983). We are especially

grateful for the financial assistance of the National

Science Foundation, which has aided this program

for many years, both at Field Museum and in Cos-

ta Rican fieldwork. The project at Field Museum

has been supported most recently by NSF grant

DEB-8103184, through the Biological Research

Resources Program. The staff and the facilities of

the Museo Nacional de Costa Rica have been a

central resource and most helpful to our work for

more than two decades. This issue includes the

large and attractive Acanthaceae family. We are

especially fortunate to have the account of this

family prepared by Dr. Durkee, a specialist in the

family who has spent considerable time in Costa

Rica and earlier published an account of the Acan-

thaceae for the Flora of Panama.

Technical assistance and fieldwork for Durkee

have been financed by grants from Grinnell Col-

lege and the Associated Colleges of the Midwest.

Work on the Acanthaceae was supported by loans

from the following herbaria: Museo Nacional de

Costa Rica, Duke University, Field Museum of

Natural History, Missouri Botanical Garden, New

York Botanical Garden, and the Museum of Nat-

ural History, Smithsonian Institution. Robin

Chazdon and Sarah Dewey provided able tech-

nical assistance; Susan Durkee (SD), Laurie Engle,

Kelly Hanick (KH), Lyla Kimura (LK), Kathleen

Neff (KBN), Christopher Rasmussen (CAR), and

Anthony Sheeder (TS) were the illustrators for the

Acanthaceae. All of these individuals were un-

dergraduates at Grinnell College during the time

of their work.

VI

FLORA COSTARICENSIS

Family #200 Acanthaceae

Family #201 Plantaginaceae

ACANTHACEAE

By L. H. Durkee

REFERENCES— C. E. B. Bremekamp, The delim-

itation of the Acanthaceae. Verh. Kon. Ned. Akad.

Wetensch., Afd. Natuurk., Tweede Sect. C. 56:

533-546. 1953. C. E. B. Bremekamp, Delimita-

tion and subdivision of the Acanthaceae. Bull. Bot.

Surv. India 7: 21-30. 1965. A. Cronquist, Acan-

thaceae, pp. 963-966. In An Integrated System of

Classification of Flowering Plants. 1981. L. H.

Durkee, Acanthaceae. In Flora of Panama. Ann.

Missouri Bot. Gard. 65: 156-283. 1978. E. C.

Leonard, Acanthaceae. In P. C. Standley, Flora of

Costa Rica. Publ. Field Mus. Nat. Hist., Bot. Ser.

18: 1188-1263. 1938. E. C. Leonard, The Acan-

thaceae of Colombia. Contr. U.S. Natl. Herb. 31.

1951-1958. G. Lindau, Acanthaceae. In A. Engler

and G. Prantl (eds.), Nat. Pflanzenfam. 4(3b): 274-

354. 1895.

Herbaceous to suffrutescent plants, some twining, less

often shrubs, rarely trees. Leaves simple, opposite (ex-

cept Elytraria of the subfamily Nelsoniodiae), exstipu-

late, often with cystoliths on the upper surfaces of lam-

inae (most evident in dried specimens). Flowers irregular

to nearly regular, perfect, borne in spikes, racemes, pan-

icles, cymes, or singly and often subtended by a con-

spicuous bract with 2 bracteoles; calyx synsepalous at

least basally, persistent, the segments (3, 4) 5(-16); the

corolla gamopetalous, the limb 5-lobed or 2-lipped, rare-

ly 1 -lipped; stamens adnate to the corolla tube, 4 and

didynamous or 2 only, staminodes often present in flow-

ers with 2 stamens; rarely all 5 stamens fully developed

(none in ours); the anther cells 2 or 1, longitudinally

dehiscent, the pollen sacs juxtaposed or sometimes su-

perposed; ovary 2-celled, superior, 2-10 ovules in each

cell, the style filiform, simple, the stigmas 1 or 2. Fruit

a loculicidal capsule (except Mendoncia with its dru-

paceous fruits), often explosively dehiscent; seeds usually

flattened, mostly borne on hook-shaped funicles, some-

times on papilliform funicles, the testa smooth or rough-

ened, often mucilaginous when wetted.

The family Acanthaceae includes about 250

genera and 2,500 species, widespread in both New

and Old World Tropics, with only a few species

occurring in warm temperate regions. The family

is divided into four subfamilies according to Lin-

dau. More recent considerations have raised each

of the subfamilies Mendoncioideae, Nelsonioi-

diae, and Thunbergioideae to family ranks. I am

in agreement with Cronquist who recognizes the

Mendonciaceae, but who feels that the Nelson-

ioideae and Thunbergioideae represent stages along

the way between the Scrophulariaceae and the

Acanthoideae, the fourth subfamily of Lindau.

While they have clear affinity with the Acanthoi-

deae, they are anomalous with either family, but

insufficiently distinct to warrant separate status.

Furthermore, Cronquist does not feel that the con-

necting forms are sufficiently numerous to warrant

uniting the Acanthaceae and the Scrophulariaceae.

The subfamily Mendoncioideae is included in this

treatment as a matter of convenience since it fol-

lows traditional usage.

The Acanthaceae can be recognized by the

prominent cystoliths appearing with magnifica-

tion as small, cylindrical concretions or short, raised

lines on the upper surface of the dried leaves,

younger stems, the branches of the inflorescence,

and the calyx. Genera lacking these are few and

have other exceptional features such as the fruit.

The genus Mendoncia is unique, with a drupa-

ceous fruit, for example. The only genera lacking

BURGER: FLORA COSTARICENSIS

cystoliths and having the more typical capsular

fruit with hooklike funicles are the genera Aphe-

landra and Megaskepasma. Seeds are frequently

ejected forcefully by the hook-shaped funicles when

the mature capsules are moistened by rainfall. In

opened or dehisced capsules, the funicles can be

seen as hook-shaped projections. Many species of

the family also have imbricate bracts subtending

the flowers. These bracts may be green, yellow,

bright red, or orange.

While most genera of the family are restricted

to either New or Old World Tropics, a few are

found in both. These include Justicia and Ruellia,

the largest genera in the family, with about 410

and 250 species, respectively. Typically, most of

these species are found in the tropics and sub-

tropics, with a few extending into temperate areas.

The major economic importance of the family

is horticultural. Cultivation is often restricted to

greenhouses, however, since in spite of attractive

foliage and flowers, they are difficult to grow. Jus-

ticia brandegeana, the shrimp plant, with its showy

spikes of reddish bracts is perhaps the best known

in the United States. Seeds of Thunbergia alata,

the clock vine or black-eyed susan, are frequently

sold in gardening stores in the United States, and

in the tropics several other species are grown for

their attractive foliage and flowers. Many other

genera are cultivated; these include: Acanthus,

Aphelandra, Crossandra, Eranthemum, Fittonia,

Graptophyllum, Hypoestes, Pachystachys, Ruellia,

Sanchezia, and Strobilanthes.

The following summary characterizes each of

the four subfamilies as recognized here and indi-

cates the Costa Rican genera belonging to each.

Conspectus of the Subfamilies and

Genera in Costa Rica

SUBFAMILY I. Ac ANTHOIDEAE— Cylindrical silic-

ified cystoliths commonly present in epidermal

cells of stems and leaves. These are best observed

under magnification of dried specimens. They are

sometimes mistaken for strigose hairs. Among the

Costa Rican genera, Aphelandra and Megaske-

pasma lack cystoliths. Seeds borne on hooklike

funicles in explosively dehiscent capsules. Most of

the genera and species of the family are found in

this subfamily: Acanthus, Aphelandra, Asystasia,

Barleria, Blechum, Bravaisia, Buceragenia, Car-

lowrightia, Chaetochlamys, Chaetothylax, Cha-

maeranthemum, Dicliptera, Dyschoriste, Grapto-

phyllum, Habracanthus, Hansteinia, Henrya,

Herpetacanthus, Hygrophila, Hypoestes, Justicia,

Louteridium, Megaskepasma, Odontonema, Poi-

kilacanthus, Pseuderanthemum, Razisea, Ruellia,

Sanchezia, Spathacanthus, Streblacanthus, Te-

liostachya, Tetramerium, Tricanthera.

SUBFAMILY II. MENDONCIOIDEAE— Mostly high-

climbing vines without cystoliths. Upper surfaces

of laminae sometimes with stellate bases of the

hairs. Flowers enclosed by two fairly conspicuous

bracts that are at first connate; calyx reduced to

an entire or low-lobed annulus. Fruit a drupe:

Mendoncia.

SUBFAMILY III. NELSONIOIDEAE— Herbs without

cystoliths in stems or leaves. Leaves opposite or

alternate. Inflorescences mostly in bracteate spikes.

Flowers with a more or less equally five-parted

calyx. Fruit a capsule with papilliform funicles:

Elytraria, Nelsonia.

SUBFAMILY IV. THUNBERGIOIDEAE— Vines or

shrubs. Cystoliths lacking in stems and leaves.

Flowers with an annular or toothed calyx; corollas

usually conspicuous; stamens four with two-celled

anthers. Fruit a beaked capsule with pulvinate fu-

nicles. The genera of this subfamily are all of Pa-

leotropical origin, and Neotropical species are cul-

tivated or escaped: Thunbergia.

FIELDIANA: BOTANY

ACANTHUS

mollis

2cm

APHELANDRA

aurantiaca

var. aurantiaca

5cm

APHELANDRA

aurantiaca

var. stenophylla

1cm

APHELANDRA

campanensis

APHELANDRA

scabra

APHELANDRA

dolichantha

FIG. 1. Acanthaceae: species of Acanthus and Aphelandra (in part) in Costa Rica. The 5-cm scale refers to foliage

and inflorescences, but note that Acanthus mollis is at a different scale.

BURGER: FLORA COSTARICENSIS

golfodulcensis

5cm

seibertii

sinclairiana

FIG. 2. Acanthaceae: species ofAphelandra (second part) in Costa Rica.

FIELDIANA: BOTANY

APHELANDRA

tonduzii

APHELANDRA

tridentata

ASYSTASIA

gangetica

BARLERIA

micans

BLECHUM

costaricense

BLECHUM

brownei

5cm

FIG. 3. Acanthaceae: species of Aphelandra (concluding part), Asystasia, Barleria, and Blechum in Costa Rica.

BURGER: FLORA COSTARICENSIS

BRAVAISIA

integerrima

BUCERAGENIA

glandulosa

CHAETOTHYLAX

eucanthus

CHAMAERANTHMUM

durandii

DICLIPTERA

5cm

imbricata

FIG. 4. Acanthaceae: species of Bravaisia, Buceragenia, Chaetothylax, Chamaeranthemum, and Dicliptera (first

part) in Costa Rica.

DICLIPTERA

iopus

FIELDIANA: BOTANY

DICLIPTERA

pallida

5cm

DICLIPTERA

podocephala

DICLIPTERA

trifurca

DICLIPTERA

unguiculata

DYSCHORISTE

valeriana

ELYTRARIA

imbricata

m

/

FIG. 5. Acanthaceae: species of Dicliptera (concluding part), Dyschoriste, and Elytraria in Costa Rica.

BURGER: FLORA COSTARICENSIS

GRAPTOPHYLLUM

pictum

HANSTEINIA

blepharorhachis

HANSTEINIA

sessilifolia

-

HANSTEINIA

stricta

HANSTEINIA

ventricosa

FIG. 6. Acanthaceae: species of Graptophyllum, Habracanthus, and Hansteinia in Costa Rica.

FIELDIANA: BOTANY

HENRYA

scorpioides

HERPETACANTHUS

panamensis

HYGROPHILA

costata

HYPOESTES

phyllostachya

JUSTICIA

angustibracteata

FIG. 7. Acanthaceae: species ofHenrya, Herpetacanthus, Hygrophilia, Hypoestes, and Justicia (first part) in Costa

Rica.

BURGER: FLORA COSTARICENSIS

JUSTICIA brenesii

5cm

candelariae

carthaginensis

comata

costaricana

FIG. 8. Acanthaceae: species ofJusticia (second part) in Costa Rica.

10

FIELDIANA: BOTANY

JUSTICIA \\ crenata

scm

•

metallica

oerstedii

orosiensis

FIG. 9. Acanthaceae: species ofJusticia (third part) in Costa Rica.

BURGER: FLORA COSTARICENSIS

11

JUSTICIA „„ parvibracteata

5cm

pectoralis

pittieri

sarapiquensis

FIG. 10. Acanthaceae: species ofJusticia (fourth part) in Costa Rica.

12

FIELDIANA: BOTANY

JUSTICIA

spicigera

5cm

tinctoria

tonduzii

urophylla

valerii

FIG. 1 1. Acanthaceae: species ofJusticia (concluding part) in Costa Rica.

BURGER: FLORA COSTARICENSIS

13

LOUTERIDIUM

costaricense

MENDONCIA

5cm

MENDONCIA

costaricana

MENDONCIA

lindavii

MENDONCIA

tonduzii

FIG. 12. Acanthaceae: species of Louteridium and Mendoncia in Costa Rica.

14

FIELDIANA: BOTANY

MEGASKEFttSMl

5cm

erythrochlamys

NELSONIA

canescens

1cm

POIKILACANTHUS

macranthus

1cm

PSEUDERANTHEMUM

cuspidatum

FIG. 13. Acanthaceae: species of Megaskepasma, Nelsonia, Odontonema, Poikilanthus, and P sender anthemum

(first part) in Costa Rica.

BURGER: FLORA COSTARICENSIS

15

PSEUDERANTHEMUM

pittieri

2 cm

PSEUDERANTHEMUM

praecox

5cm

PSEUDERANTHEMUM

standleyi

citrina

1 cm

RAZISEA

spicata

RAZISEA

wilburii

FIG. 14. Acanthaceae: species of Pseuderanthemum (concluding part) and Razisea in Costa Rica.

16

FIELDIANA: BOTANY

RUELLIA

biolleyi

y

1cm

5cm

geminiflora

1cm

golfodulcensis

inundata

jussieuoides

FIG. 15. Acanthaceae: species of Ruellia (first part) in Costa Rica.

BURGER: FLORA COSTARICENSIS

17

RUELLIA

metallica

5cm

nudiflora

palustris

paniculate

1cm

pittieri

praeclara

FIG. 16. Acanthaceae: species of Ruellia (second part) in Costa Rica.

18

FIELDIANA: BOTANY

5cm

RUELLIA stemonacanthoides

RUELLIA

tonduzii

RUELLIA

tubiflora

var. hirsuta

RUELLIA

tubiflora

var. tetrastichantha

1cm

STREBLACANTHUS

monospermus

FIG. 17. Acanthaceae: species of Ruellia (concluding part), Spathacanthus, and Streblacanthus in Costa Rica.

BURGER: FLORA COSTARICENSIS

19

1cm

TELIOSTACHYA

alopecuroidea

5cm

TETRAMERIUM

nervosum

TRICHANTHERA

gigantea

THUNBERGIA

alata

FIG. 18. Acanthaceae: species of Teliostachya, Tetramerium, Trichanthera, and Thunbergia in Costa Rica.

20

FIELDIANA: BOTANY

Key to Genera of Acanthaceae in Costa Rica

1 a. Fertile stamens 4 2a

Ib. Fertile stamens 2 1 7a

2a. Plants usually scandent; seeds not borne on hooklike funicles 3a

2b. Plants not scandent, usually erect herbs or shrubs; seeds borne on hooklike funicles or, if on

papilliform funicles, capsules not beaked 4a

3a. Fruit a drupe Mendoncia

3b. Fruit a beaked capsule with 2-4 seeds Thunbergia

4a. Corolla 1 -lipped; stamen filaments thick and prominent; leaves sinuate pinnatifid

Acanthus

4b. Corolla 2-lipped, subequal or regular; stamen filaments narrow; leaves ovate, obovate, elliptic,

or oblong, but not sinuate-pinnatifid 5a

5a. Calyx spathaceous, 2-lobed; fruits 6.5-8 cm long Spathacanthus

5b. Calyx 4- or 5-parted, not spathaceous; fruits less than 4 cm long 6a

6a. Anthers 1 -celled; cystoliths lacking in the leaves Aphelandra

6b. Anthers 2-celled in at least 1 pair of stamens; cystoliths present in the leaves 7a

7a. Calyx segments 4, unequal; bracts with spiny margins Barleria

7b. Calyx segments 5, either equal or unequal; bracts lacking spiny margins 8a

8a. One pair of stamens with 2-celled anthers, the other pair of stamens with 1 -celled anthers

or sterile 9a

8b. Both pairs of stamens with 2-celled anthers lOa

9a. Corolla bilabiate; bracts loosely imbricate and conspicuous, 9-14 mm long

Herpetacanthus

9b. Corolla nearly regular; bracts lax and inconspicuous, 1.5-2 mm long

Chamaeranthemum

lOa. Shrubs or trees; calyx segments often apically rounded or obtuse 1 la

1 Ob. Herbs, often more or less suffrutescent (rarely shrubs); calyx segments subulate or lanceolate,

usually apically acute or acuminate 1 2a

1 la. Anthers basally rounded, not spurred; capsules 4-seeded, sericeous Tricanthera

lib. Anthers basally spurred; capsules 8-seeded, glabrous Bravaisia

1 2a. Corolla conspicuously bilabiate, 1 cm long or less; capsules sessile (bilocular from base) . .

13a

12b. Corolla regular or nearly so; capsule stipitate (basal portion solid for at least 2 mm) . . 14a

1 3a. Inflorescence a dense spike; anther cells unequal; seeds 4 per capsule . . Teliostachya

13b. Inflorescence fasciculate in the leaf axils; anther cells equal; seeds 12-16 per capsule

Hygrophila

1 4a. Inflorescences in 1 -sided racemes; corollas yellow with purplish throats Asystasia

1 4b. Inflorescences in cymes, heads, panicles, spikes, or flowers axillary; corollas variously colored,

not yellow with purplish throats 1 5a

1 5a. Flowers in closely imbricate bracted, 4-sided spikes; bracts conspicuous, ovate . . Blechum

1 5b. Flowers otherwise; bracts mostly inconspicuous and linear 1 6a

1 6a. Calyx segments fused for '/j-'/i their length; anther cells basally spurred Dyschoriste

\ 6b. Calyx segments parted nearly to base; anther cells basally rounded Ruellia

17a. Anthers 1 -celled, or if 2, one always much smaller than the other and usually abortive 18a

1 7b. Anthers 2-celled, the cells about equal in size 25a

1 8a. Corolla less than 5 mm long; staminodes present Buceragenia

1 8b. Corolla more than 5 mm long; staminodes absent 1 9a

1 9a. Inflorescences condensed, headlike, bracts white-ciliate, linear subulate 20a

1 9b. Inflorescences open, bracts otherwise 2 la

20a. Calyx segments 4; corolla ca. 1 5 mm long Chaetothylax

20b. Calyx segments 5; corolla more than 20 mm long Chaetochlamys

2 la. Corolla blue; inflorescence a loose, terminal thyrse (cymose panicles) Habracanthus

BURGER: FLORA COSTARICENSIS 21

2 1 b. Corolla white, red, magenta, orange, or yellow; inflorescence of single flowers, spikes, racemes,

compact thyrses, or racemose panicles 22a

22a. Leaves pink-dotted; corolla with 3-lobed upper lip and 1 -lobed lower lip Hypoestes

22b. Leaves lacking pink dots; corolla with 1 - or 2-lobed upper lip and 3-lobed lower lip . . 23a

23a. Inflorescences paniculate; corolla tube usually abruptly enlarged above the ovary

Hansteinia

23b. Inflorescences a spike or narrow spikelike thyrse; corolla tube expanding gradually from the

base 24a

24a. Inflorescences spicate; calyx 4-parted Streblacanthus

24b. Inflorescences a narrow spikelike thyrse; calyx 5-parted Razisea

25a. Inflorescences condensed, headlike; bracts white-ciliate, linear subulate 26a

25b. Inflorescences open, bracts otherwise 27a

26a. Calyx segments 4; corolla ca. 1 5 mm long Chaetothylax

26b. Calyx segments 5; corolla more than 20 mm long Chaetochlamys

27a. Cystoliths lacking in the leaves; seeds not borne on hooklike funicles 28a

27b. Cystoliths present in the leaves; seeds borne on hooklike funicles 30a

28a. Bracts large and showy, more than 20 mm long; corolla more than 30 mm long

Megaskepasma

28b. Bracts small and inconspicuous, less than 10 mm long; corolla less than 15 mm long . 29a

29a. Bracts stiff and awn-tipped; leaves mostly more than 10 cm long Elytraria

29b. Bracts flaccid and awnless; leaves mostly less than 8 cm long Nelsonia

30a. Staminodes present 3 la

30b. Staminodes absent 35a

3 la. Anther cells basally spurred; flowers in fascicles of 3-8, loosely enclosed by bracts

Sanchezia

31b. Anther cells not basally spurred; flowers not in fascicles enclosed by bracts 32a

32a. Bracts conspicuous with spiny margins, more than 2 cm long; calyx 4-parted, segments

unequal Barleria

32b. Bracts inconspicuous, without spiny margins, less than 1 cm long; calyx 5-parted, segments

equal 33a

33a. Corolla limb subequally 5-parted, tube narrowly cylindrical, limb spreading; stamens included

(except Pseuderanthemum atropurpureum) Psenderanthemum

33b. Corolla limb 2-lipped, tube expanding; stamens exserted 34a

34a. Leaf blades mottled; corollas 4.5-8 cm long; plants cultivated Graptophyllum

34b. Leaf blades green; corollas 1.5-3 cm long; plants mostly wild Odontonema

35a. Flowers in dense spikes; bracts imbricate, often 4-ranked 36a

35b. Flowers borne in lax spikes, panicles, cymes, or thyrses; bracts not imbricate and not 4-ranked

38a

36a. Bracts more than 2 cm long; calyx segments widely dissimilar Barleria

36b. Bracts less than 2 cm long; calyx segments similar 37a

37a. Bracts 4-ranked; floral bracts not uniting to form an involucre Tetramerium

37b. Bracts not 4-ranked; floral bracts uniting almost to apex forming an involucre .... Henrya

38a. Stems 6-angled in cross section; calyx subtended by 2 partially united bracts Dicliptera

38b. Stems rounded to 4-angled; bracts distinct 39a

39a. Corolla lobes 4; mature dried capsules black or dark purple Carlowrightia

39b. Corolla lobes 5; mature dried capsules yellowish brown to dark brown 40a

40a. Calyx 3-parted Louteridium

40b. Calyx 5-parted 41a

4 1 a. Bracts large, to 44 mm long Megaskepasma

4 1 b. Bracts small, less than 20 mm long 42a

42a. Corolla more than 6 cm long; leaf pairs unequal (excluding J. trichotoma which has unequal leaf

pairs); pollen polyporate Poikilacanthus

42b. Corolla less than 6 cm long (all species except /. aurea are less than 5 cm long); leaf pairs equal,

with the exception of/, trichotoma; pollen mostly 2-porate, some 3- or 4-porate Justicia

FIELDIANA: BOTANY

Acanthus L.

REFERENCE— T. H. Everett, The New York Bo-

tanical Garden Illustrated Encyclopedia of Hor-

ticulture 1: 21. 1980.

Herbs or small shrubs. Leaves petiolate; laminae broad

and pinnatifid. Inflorescences in terminal spikes; flowers

sessile, borne singly in the axils of the bracts; bracteoles

2 per flower; calyx 4-merous. segments unequal; corolla

1 -lipped, the lip 3-lobed; stamens 4, filaments thick and

prominent, anthers 1 -celled, wooly. Fruit a capsule.

This is a genus of about 20 species in subtropical

and tropical regions of the Mediterranean, Asia,

and Africa. Two species are often cultivated for

their appearance, A. montanus T. Anders and A.

mollis L.

Acanthus mollis L., Sp. pi. 639. 1753. Figure 1.

Herbs to 70 cm tall, erect. Leaves mostly basal, pet-

ioles to 2 1 cm long, subquadrangular, 3 mm wide, spar-

ingly pilose; laminae to 60 cm long, 30 cm broad, cor-

date, smuatc-pmnatifid. dentate, sparingly pilose, more

so on the veins. Inflorescences in large, terminal spikes

to 65 cm tall; peduncle quadrangular to 35 cm long, 5

mm thick, sparingly pilose, rachis similar; floral bracts

ovate, to 3.5 cm long, 1.5 cm broad, apically acute, spine-

tipped, puberulous, margins with 6-8 spreading, spiny

teeth to 5 mm long; bracteoles linear, 15-20 mm long,

1.5-2 mm broad, spine-tipped, puberulous; upper calyx

segment obovate, 4.5 cm long, 2 cm broad, apically

2-lobed, lobes 25 mm long apically acuminate, lower

segment obovate, 3.5 cm long, 1.2 cm broad, apically

2-lobed, lobes 22 mm long, apically acuminate, lateral

lobes orbicular, 50 mm in diameter, all segments puber-

ulous; corolla whitish, rose or lilac, 1 -lipped, to 4 cm

long, the lip 3-lobed, lobes suborbicular, 4 mm long,

middle lobe to 1 9 mm broad, lateral lobes 1 4 mm broad,

glabrous, tube short, 4 mm long, glabrous with a circle

of hairs 2 mm long at point of attachment of stamens;

stamens included within upper calyx lobe, filaments sig-

moid, to ca. 20 mm long, 2 mm thick, prominent, in-

durate, glabrous; anthers 10 mm long, 2 mm broad.

Fruits not seen; seeds 4.

A common planting in yards and flower gardens

of the Meseta Central. This species is a native of

Italy.

Aphelandra R. Br.

REFERENCES— D. C. Wasshausen, The Genus

Aphelandra, Smithsonian Contr. Bot. No. 18. 1975.

L. A. McDade, Systematics and reproductive bi-

ology of the Central American species of the Aphe-

landra pulcherrima complex (Acanthaceae). Ann.

Missouri Bot. Card. 71: 104-165. 1984.

Shrubs or suffrutescent herbs. Leaves opposite; lam-

inae usually large, oblong to elliptic, margins toothed,

lobed, crenate or entire. Inflorescences of terminal or

axillary spikes, spikes often conspicuous with showy

bracts and flowers; bracts most often imbricate, certain

species bearing dorsal clusters of submarginal glands (ex-

tra-floral nectaries) which may be conspicuous, margins

of bracts entire, toothed or ciliate; bracteoles various,

rudimentary in a few species; flowers with 5-merous ca-

lyx, segments separate nearly to base, frequently lanceo-

late and striate-veined, posterior segment usually broad-

er than others; corolla straight or curved, the limb usually

bilabiate, upper lip erect, 2-lobed or entire, lower lip

reflexed or spreading, 3-lobed, the middle lobe often

larger than the laterals; stamens 4, usually exserted, but

usually not beyond the upper lip of the corolla; anthers

narrow, 1 -celled, often pilose dorsally and held together

at apices by intertwining hairs. Fruits a capsule, usually

clavate, 4-seeded.

Aphelandra is one of the larger genera of the

family and is endemic to tropical America. Species

of the genus are often quite showy and yet they

are seldom cultivated because of difficulties in cul-

tivation. In Costa Rica the genus is represented in

a variety of forested habitats from sea level to

elevations of 1 500 m, including the dry forests of

the Pacific watershed, cloud forests of higher ele-

vations, and in the rain forests of the Caribbean

watershed. Aphelandra scabra of the Pacific dry

forests is probably the most abundant and wide-

spread species in the genus.

Key to Species of Aphelandra

la. Bracts toothed 2a

Ib. Bracts entire 8a

2a. Bracts with submarginal glands 3a

2b. Bracts without submarginal glands 4a

3a. Bracts more than 20 mm long; petioles 1 0-30 mm long A. storkii

3b. Bracts less than 1 6 mm long; petioles 0-5 mm long A. scabra

BURGER: FLORA COSTARICENSIS

23

4a. Bracts less than 1 5 mm long; corolla less than 2 cm long 5a

4b. Bracts more than 20 mm long; corolla more than 5 cm long 6a

5a. Leaves mostly less than 10 cm long; bracts more than 10 mm long A. seibertii

5b. Leaves more than 10 cm long; bracts less than 8 mm long A. tonduzii

6a. Bracts with 1-2 pairs of teeth above midpoint; peduncle more than 10 mm long

A. trident at a

6b. Bracts with spine-tipped, serrate margins; peduncle short, less than 5 mm long 7a

7a. Leaves elliptic to ovate, 3-8.5 cm broad A. aurantiaca var. aurantiaca

7b. Leaves lanceolate, 1.5-2.5 cm broad A. aurantiaca var. stenophylla

8a. Bracts with submarginal glands 9a

8b. Bracts without submarginal glands 1 3a

9a. Submarginal glands consisting of numerous, minute glands appearing as a dull, alveolar spot

lOa

9b. Bracts with submarginal glands represented by 1 or more well-defined, elliptical, shiny areas

12a

lOa. Petioles short, to ca. 1 cm long; corolla puberulous; capsules clavate . . A. lingua-bovis

lOb. Petioles 2-8 cm long; corolla papillose; capsules oblong or ovate 1 la

1 la. Bracts acuminate to acute, 7-9.5 mm long; calyx segments 9-1 1 mm long

A. leonardii

1 Ib. Bracts obtuse to acute, 10-1 1 mm long; calyx segments 15-17 mm long

A. campanensis

12a. Bracts 6-13 mm long, drying to light or dark brown color; younger stems strigose

A. golfodulcensis

1 2b. Bracts 1 6-20 mm long, drying to light orange color (orange-red when fresh); younger stems

velutinous A. sinclairiana

13a. Bracts more than 3 cm long; young stems quadrangular, puberulous to glabrous . A. dolichantha

1 3b. Bracts less than 8 mm long; stems terete, pilose A. tonduzii

Aphelandra aurantiaca (Scheidw.) Lindl. in Ed-

wards, Bot. Reg. 31: pi. 12. 1845. Hemisandra

aurantiaca Scheidw., Bull. Acad. Roy. Sci. Bru-

xelles9:22. 1842. Figure 1.

Suffrutescent herb to shrub, erect, 1 m tall; young steins

tetrangular, older stems terete, internodes between leaf-

bearing nodes 2-6 cm long, 2-4 mm thick, younger stems

glabrous to velutinous. Leaves petiolate; petioles to 1 cm

long, glabrous; laminae elliptic to ovate with undulate

to entire or sinuate-repand margins, frequently thick-

ened, 8-28 cm long, 2.5-12.5 cm broad, apically acu-

minate, basally attenuate, occasionally obliquely decur-

rent, glabrous except for occasional puberulence along

veins, costa and lateral veins prominent, reticulations

conspicuous on upper surface and obscure beneath. In-

florescences in mostly terminal, quadrifarious spikes to

1 7 cm long, 3 cm broad, excluding the corollas; peduncle

short, to 3 mm long, 2 mm thick, puberulous, rachis

similar, flattened at the nodes; bracts elliptic or ovate,

imbricate, 2.5-3 cm long, 0.75-1 cm broad, apically acute,

basally cuneate, puberulous-glandular on both surfaces,

reticulations prominent on inner surface, margins ser-

rate, serrations tipped with spines to 1.5 mm long; brac-

teoles narrowly lanceolate, to 10 mm long, 0.5 mm broad,

puberulous on both surfaces, striate-ridged. Flowers with

calyx segments 1 cm long, posterior segment oblong, 2.5

mm broad, with a 3-toothed apex, lateral and anterior

segments linear-lanceolate, 1 mm broad, all segments

puberulous with striate ridges culminating at the base in

callus tissue ca. 1 mm thick; corolla red, orange-red, or

scarlet, tube to 4 cm long, 3 mm broad, bilabiate, upper

lip to 2 cm long and 1 cm broad above base, oblong-

ovate, apiculate at apex, forming a partial hood over the

anthers, lower lip 3-lobed, elliptic middle lobe 2 cm long,

1.2 cm broad, apically acute, lateral lobes elliptic-ovate,

to 1 .2 cm long, 0.75 cm broad, apically acute, puberulous

on both surfaces; style to 4.6 cm long, sparingly puber-

ulous with ascending hairs, ovary puberulous at apex.

Fruits 1 .6 cm long, sparingly puberulous, seeds hispid-

ulous.

Found in dense forests from Mexico to Bolivia.

Aphelandra aurantiaca is recognized by its large,

spiny toothed bracts which lack submarginal

glands. It is one of the most easily recognized

species of the genus and does not closely resemble

any of the other Costa Rican species.

Aphelandra aurantiaca (Scheidw.) Lindl. var. au-

rantiaca

The characteristic features of this variety are elliptic

to ovate laminae 8-21 cm long, 3-8.5 cm broad with

undulate to entire margins that are usually thickened.

24

FIELDIANA: BOTANY

This variety is found in dense forests from Mex-

ico to Bolivia. In Costa Rica it ranges from the

wet evergreen forests of the Caribbean watershed

to the moist evergreen forests around the Meseta

Central and into Guanacaste to Liberia, at ele-

vations near sea level to above 1 500 m. Flowering

collections have been made from August through

January.

Aphelandra aurantiaca (Scheidw.) Lindley var.

stenophylla Standley, Publ. Field Columbian

Mus., Hot. Ser. 4: 324. 1929; Field Mus. Nat.

Hist., Bot. Ser. 10: 59. 1931. Aphelandra re-

panda Nees in DC., Prodr. 11: 728. 1847.

The characteristic features of this variety are lanceo-

late leaves, laminae 20-28 cm long, 1.5-2.5 cm broad,

gradually narrowed apically and basally, margins sin-

uate-repand.

Commonly found in wet forests from Central

America to Bolivia. Collected mostly in Costa Rica

in the Meseta Central region from an altitude of

1 300 m down to elevations of 700 m near Tilaran,

Guanacaste.

Aphelandra campanensis Durkee, Ann. Missouri

Bot. Gard. 65: 162. 1978. Figure 1.

Shrub to 3.5 m tall; internodes between leaf-bearing

nodes 2.5-8 cm long, 3.5-7.5 mm thick, tetrangular,

sparingly to moderately strigose, lenticels corky ca. 1 mm

long. Leaves with strigose petioles to 5 cm long; laminae

elliptic, 14-30 cm long, 7-12.5 cm broad, apically acu-

minate, basally attenuate, margins entire or undulate,

glabrous to sparingly strigose above, strigose beneath,

more so on the costa and veins. Inflorescences in ter-

minal spikes borne singly or in clusters, to 24 cm long,

1-1.5 cm broad excluding corollas; peduncles short, 5-

13 mm long, sparingly strigose; rachis glabrate; bracts

rhombic-ovate, 10-11 mm long, 7-8 mm broad, apically

obtuse to acute, apiculate, glabrate and minutely rough-

ened with strigose hairs in some, the nerves prominent

except toward the apex, with 2 submarginal glandular

areas at about the midpoint, each consisting of a compact

cluster of numerous, minute, dull brown glands making

an elliptic, alveolar spot 1.5-2.0 mm long and 0.5-0.75

mm broad, bract margins subhyaline, entire; bracteoles

falcate-lanceolate, 9-10 mm long, 2.5-3.25 mm broad,

keeled, striate-nerved, densely hirsute along the keel,

marginal area glabrous, subhyaline, sparingly papillose.

Flowers with the calyx segments conspicuously longer

than the bracts, segments 15-17 mm long, apically ob-

tuse, apiculate, striate-nerved, glabrous outside, anterior

segments lanceolate, 3.5 mm broad, posterior segment

ovate, 7 mm broad; corolla orange-red, bilabiate, densely

papillate outside and inside, 5-7 cm long, the tube to 4

cm long, 5 mm broad basally, constricted to 3.5 mm

broad just above the apex of the ovary, expanding to 1 1

mm broad at the mouth, the upper lip ovate, ca. 1 9 mm

long, 9 mm broad, apically with 2 acuminate lobes ca.

6 mm long, the lower lip spreading at maturity, the mid-

dle lobe ovate, to 23 mm long, 1 1 mm broad, apically

acuminate, the tip keeled, the lateral lobes triangular,

the free portion 1.5 mm long, 2 mm broad, fused to the

upper lip for most of their length, the tip rounded; sta-

mens extending to the tip of the upper lip, the anthers 9

mm long, 1.5 mm broad, acute at both ends; style and

ovary glabrous. Fruits ovate, ca. 20 mm long, 6 mm

broad, the tip rounded, the surface shiny and punctic-

ulate; seeds suborbicular, 4 mm in diameter, the surface

minutely roughened.

This species occurs in lowland to mid-elevation

( 1 000 m) rain forests mainly in western provinces

bordering on the Caribbean in Panama, but ex-

tends into Costa Rica. Only one Costa Rican col-

lection, McDade 242 (DUKE), has been observed.

Flowering collections have been made from Au-

gust through November.

Aphelandra campanensis is recognized by its

shrubby habit; compact terminal inflorescences

with imbricate bracts that lack marginal teeth, are

apically obtuse to acute, and have two submarginal

glandular areas, each consisting of numerous mi-

nute, dull, alveolar-appearing glands; its calyx lobes

conspicuously longer than the bracts; and its 5- to

7-cm long corollas that are orange-red and are

externally papillate. It may be confused with A.

leonardii, which differs in its shorter bracts (7 to

9.5 mm long vs. 10 to 1 1 mm), which are apically

acute to acuminate, its shorter calyx segments (9

to 1 1 mm long vs. 1 5 to 17 mm), and its bright

red corollas. It may also be confused with A. lin-

gua-bovis, which can be distinguished by its cla-

vate capsules, bright red bracts, and puberulous

corollas.

Aphelandra dolichantha Donnell Smith, Bot. Gaz.

(Crawfordsville) 27: 438. 1899. Figure 1.

Shrubs or suffrutescent herbs ca. 1 m tall; young stems

quadrangular, internodes between leaf- bearing nodes 1-

5 cm long, 2.5-4 mm thick, puberulous to glabrous, hairs

brownish. Leaves petiolate, petioles 0.5-2 cm long, mi-

nutely strigose; laminae mostly elliptic, to 24 cm long,

10.5 cm broad, apically acuminate, basally attenuate,

margins entire, glabrous above, glabrous to finely strigose

below, when present, hairs more dense on costa and

lateral veins, drying to a lighter green than the upper

surface, costa and lateral veins prominent. Inflorescences

of terminal, sessile spikes, borne singly or occasionally

in small clusters, to 13 cm long, 1.5 cm broad excluding

corollas; rachis puberulous; bracts closely imbricate at

anthesis, spreading at time of fruiting, green when fresh,

BURGER: FLORA COSTARICENSIS

25

drying reddish brown, ovate-lanceolate to oblanceolate,

to 3.5 cm long, 1 cm broad, apically acuminate to sub-

acute, glabrous, margins ciliolate; bracteoles linear, to 8

mm long, 0.5 mm broad, with appressed pilose hairs,

0.5 mm long plus smaller gland-tipped hairs. Flowers

6.5-7.2 cm long; calyx minute, segments linear-subulate,

ca. 1.5 mm long, 0.25 mm broad, ciliolate; corolla white,

the tube slender, to 5 cm long, nearly cylindrical, basally

2 mm broad, 3.5 mm broad at throat, glandular-puber-

ulous, the upper lip erect, oblong, to 1 2 mm long, 6 mm

broad, 2-lobed, the lobes 5 mm long, 3 mm broad at

base, apically rounded, the lateral lobes oblong to 17

mm long, 6 mm broad, apically rounded, the middle

lobe of the lower lip to 22 mm long, 9 mm broad, apically

rounded; stamens included, filaments ca. 6 mm long,

affixed just below the throat, anthers 2.5 mm long, ba-

sally apiculate, adhering by terminal hairs; style includ-

ed. Fruits clavate, 17 mm long, 4 mm broad, 3 mm

thick, glabrous, minutely punctate, perianth persistent

except early deciduous corolla; seeds flattened, ovoid to

obliquely ovoid, to 5 mm long, 4 mm broad, 1.5 mm

thick dark brown, smooth.

Found in lowland wet evergreen forests on the

Caribbean side of Costa Rica, in higher wet ev-

ergreen forests of Bajo La Hondura to an altitude

of 1400 m, and at 750 m near Tilaran in Guana-

caste. Flowering collections have been made from

April through October. The species is also found

in similar habitats of Panama and Colombia.

Aphelandra dolichantha is recognized by its long

(more than 3 cm), narrow (1 cm) bracts, which

lack submarginal glands, and its large (more than

5 cm long), narrow, white flowers. It slightly re-

sembles A. storkii, but this species has bracts that

are shorter, with toothed margins and submarginal

glands, orange flowers, and apparently a much more

restricted habitat.

Aphelandra golfodulcensis McDade, Ann. Mis-

souri Bot. Card. 69: 405. 1982. Figure 2.

Shrubs to small trees 1-6 m tall; internodes between

leaf-bearing nodes 1-4 cm long, 3-6 mm thick, terete,

densely retrorse. Leaves on moderately strigose petioles

to 1 cm long; laminae elliptic to oblanceolate, 8.5-30

(45) cm long, 2-8 ( 1 5) cm broad, apically acute to acu-

minate, basally attenuate, margins entire or slightly un-

dulate, sparsely strigose on the veins above, moderately

strigose beneath. Inflorescences borne in numerous ter-

minal spikes to 1 5 cm long, 1 cm broad excluding co-

rollas; peduncles to 10 cm long, moderately pubescent;

rachis glabrous to minutely puberulous; bracts imbricate,

rhombic-ovate, apically acute, 6-10 mm long, 4-7 mm

broad, green to dull brown-orange, margins ciliolate, with

2 clusters of submarginal glands just below the middle

of the dorsal surface, glands ca. 0.75 mm long, 0.5 mm

broad, 1-7 glands per cluster; bracteoles narrowly ovate,

apically attenuate, 4-6.5 mm long, 2-4 mm broad, green,

moderately puberulous. Flowers with calyx segments 6-

9 mm long, apically acute, green, finely striate, minutely

puberulous, posterior segment narrowly ovate, 3-4 mm

broad, anterior segments broadly lanceolate, 2-2.5 mm

broad, lateral pair narrowly lanceolate, ca. 1 .5 mm broad;

corolla orange to red, 6.0-7.3 cm long, puberulous, tube

3.8—4.0 cm long, 2-3 mm broad basally, expanding to

6-8 mm broad at the mouth, the upper lip erect, elliptic,

17-19 mm long, 7-11 mm broad, 2-lobed, the lobes

triangular, acuminate, 6-10 mm long, the lateral lobes

of the lower lip triangular, 3 mm long, 5-7 mm broad,

middle lobe broadly lanceolate, spreading, 22-26 mm

long, 6-9 mm broad, apically acuminate; stamens exsert-

ed to just below the tip of the upper lip and enclosed by

it. Fruits clavate, terete, 19-23 mm long, 5-8 mm broad,

5.5-7 mm thick, glabrous; seeds 4, slightly flattened,

orbicular, 4-6 mm in diameter.

According to McDade this species is found pri-

marily in the wet lowlands of the Golfo Dulce

region in Puntarenas Province. It extends into the

Burica Peninsula of Panama, to mid-elevations

above the Golfo Dulce region, and to the north in

Alajuela and Guanacaste, where local conditions

provide similar moist habitats. Flowering collec-

tions have been made from December through

March.

Aphelandra golfodulcensis is recognized by its

imbricate, rhombic-ovate bracts (6 to 10 mm long,

4 to 7 mm broad), which are green or dull brown-

orange and minutely puberulent, its minutely pu-

berulent corolla, and its glabrous green fruits. It

most closely resembles A. sinclairiana, which dif-

fers in its larger obovate-rhombic bracts ( 1 6 to 2 1

mm long, 14 to 20 mm broad), which are bright

orange, its corollas, which are short velutinous,

and its black fruits, which are pubescent. Accord-

ing to McDade these two species are closely re-

lated, as observed in experimental hybridizations

and several collections of putative hybrids from

northeastern Costa Rica. She reports further that

these hybrids are morphologically intermediate

between the two parental species and are sterile.

Aphelandra leonardii McDade, Ann. Missouri Bot.

Gard. 69: 408. 1982. Figure 2.

Shrubs 1-5 m tall; younger stems quadrangular, older

stems terete, internodes between leaf-bearing nodes 2-5

cm long, 1.5-2.5 mm thick, strigose, lenticels scattered,

oval, ca. 0.5 mm long, corky. Leaves petiolate, petioles

ca. 3 cm long, pubescence that of the costa; laminae

elliptic to elliptic-obovate, 10-20 (30) cm long, 4-10 cm

broad, apically acute to acuminate, basally attenuate,

margins shallowly crenate to slightly undulate, glabrous

above, sparingly strigose to glabrous below. Inflores-

cences in terminal spikes, quadrangular, to 8 cm long, 1

26

FIELDIANA: BOTANY

cm broad excluding corollas; peduncle subsessile, rachis

glabrous to sparingly puberulous; bracts rhombic-ovate,

imbricate, green to bright orange, 7-9.5 mm long, 5-6

mm broad, apically acute to acuminate, tip apiculate,

keeled and stria te-nerved, minutely puberulous on upper

bracts, lower bracts increasingly strigose mostly over the

keel, with submarginal glandular areas just below mid-

point consisting of numerous, minute (0.2 mm), often

shiny brown to dark brown glands making an elliptic,

alveolar spot 1-1.5 mm long, 0.5-1 mm broad, margins

ciliolate; bracteoles falcate, keeled, to 8 mm long, 2.5

mm broad, striate-nerved, finely puberulous outside ex-

cept wooly keel, glabrous inside. Flowers 6.5-7.5 cm

long; calyx segments conspicuously longer than the bracts,

9-1 1 mm long, glabrous, anterior segments oblong, all

others lanceolate, apically acute, anterior segments 3 mm

broad, lateral segments 2.5 mm broad, posterior segment

5 mm broad, all striate-nerved; corolla bright red, bi-

labiate, papillate outside and inside, tube to ca. 5 cm

long, 4 mm broad basally, constricted to ca. 2 mm broad

just above the ovary, expanding to 9 mm broad at the

throat, upper lip erect, ca. 20 mm long, basally 9 mm

broad, consisting of 2 acuminate lobes ca. 1 2 mm long,

the lower lip spreading at maturity, with 3 lobes, the

middle lobe lanceolate, ca. 26 mm long, 8 mm broad,

the lateral lobes ca. 6 mm long, acute, connate to the

upper lip for most of their length; stamens extending to

ca. the length of the upper lip. Fruits oblong, flattened,

glabrous, 17.5-19 mm long, ca. 5 mm broad, 3.5-4 mm

thick; seeds irregularly orbicular, strongly flattened, 3.5-

6.5 mm in diameter.

A shrub of the forest understory, this species is

found in lowland and premontane forests in east-

ern Panama and in Costa Rica from along the

Pacific in Guanacaste to the Meseta Central and

the Cordillera Talamanca north of San Isidro to

elevations of 1 300 m. Flowering collections have

been made in December only.

Aphelandra leonardii can be recognized by its

calyx, which is conspicously longer than the bracts,

its short (7 to 9.5 mm long), entire bracts with

submarginal alveolate glands, and its long (ca. 7

cm), bright red corollas, which are externally pa-

pillate. It resembles A. campanensis, a largely Pan-

amanian species that has been collected in adja-

cent areas of Costa Rica. (See the description of

that species for a summary of differences.) It also

resembles A. lingua-bovis, and these differences

are summarized in descriptions of both this species

and A. campanensis.

Aphelandra lingua-bovis Leonard, Contr. U.S.

Natl. Herb. 31: 268. 1953. Figure 2.

Shrubs, 1-3.5 m tall; stems terete or subquadrangular

toward tips, internodes between leaf-bearing nodes 2-9

cm long, 3-5 mm thick, moderately and minutely stri-

gose, the hairs upwardly appressed. Leaves petiolate, pet-

ioles to ca. 1 cm long, strigose; laminae oblanceolate to

narrowly elliptic, to 38 cm long, 1 2 cm broad, apically

acuminate and often curved to one side, basally atten-

uate, margins entire or undulate, glabrous to sparingly

strigose above, hairs densest along the prominent costa

and lateral veins, strigose to sparingly strigose below,

lower surface drying to a lighter green than the upper.

Inflorescences in terminal spikes, mostly solitary, spikes

to 38 cm long, 1.2 cm broad; peduncle to 1 cm long,

sericeous; rachis sparingly sericeous; bracts bright red

when fresh, drying to reddish brown, rhombic-ovate, 10-

14 mm long, 8-10 mm broad, apically acute to slightly

obtuse, glabrous to puberulous with upwardly appressed

hairs along the prominent costa, lateral veins also prom-

inent, with 2 dull, elliptic, alveolate, submarginal glan-

dular areas about midway from the base, to 3 mm long,

1 mm broad, margins ciliolate; bracteoles linear-lanceo-

late, to 1 5 mm long, 2 mm broad, apically acuminate,

keeled, the keel densely hirsute with ascending hairs ca.

1 mm long. Flowers with the calyx ca. 1 7 mm long, the

segments lanceolate, increasingly puberulous toward the

apex, striate-nerved, the posterior segments 8 mm broad,

the lateral segments 1 .7 mm broad, the anterior segments

2.2 mm broad; corolla bright red, or reddish yellow, 5.5-

6 cm long, puberulous, the tube subcylindric, slightly

curved, 3.5 mm broad at the base, 6 mm broad at the

throat, the upper lip erect, elliptic, 16 mm long, 7 mm

broad, 2-lobed, the lobes apically acute, 4 mm long, 3

mm broad basally, the middle lobe of the lower lip lan-

ceolate, 1 8 mm long, 6 mm broad, acute, the lateral lobes

4 mm long, partly coalescent to the upper lip, the free

portions triangular, ca. 1.5 mm long, 1 mm broad, api-

cally acute; stamens slightly exserted, anthers apiculate

at both ends, webby-pilose at tip; ovary densely strigose,

hairs ascending. Fruits clavate, 1 8-20 mm long, 4 mm

broad, 3.5 mm thick, glabrous except for ascending stri-

gose hairs toward the round apex, perianth persistent

except the early deciduous corolla; seeds 4, dark brown,

deltoid-shaped, ca. 5 mm long, 4 mm broad, margins

sparingly tuberculate.

Found in mature rain forest of lowland areas in

southwestern Costa Rica. Most collections are from

the Golfo Dulce area at low elevation, although

one collection, Raven 22017 (F), has been made

at Wilson's finca south of San Vito (San Vito de

Java) at ca. 1300 m elevation. Flowering collec-

tions made from June through March. This species

is also found in Panama and Colombia.

Aphelandra lingua-bovis is recognized by its long

(to 38 cm), compact inflorescences with entire,

bright red bracts (when fresh) with alveolate sub-

marginal glands and large (ca. 6 cm long), pu-

berulous, bright red corollas. It resembles A. cam-

panensis and A. leonardii, which differ in their

smaller inflorescences, papillose corollas, and either

oblong or ovate (vs. clavate) capsules.

Aphelandra sea bra (Vahl) Sm. in Rees, Cyclopae-

dia 39, Aphelandra n. 3( 1 8 1 8), based on Justicia

BURGER: FLORA COSTARICENSIS

27

scabra Vahl, Enum. 1: 120. 1804. Aphelandra

deppeana Schlecht. & Cham., Linnaea 5: 96.

1830. A. pulcherrima sensu H.B.K., Nov. gen.

sp. 2: 236. 1817. A. pectinata Willd. ex Nees,

DC., Prodr. 1 1: 298. 1847. A. haenkeana Nees,

DC., Prodr. 11: 298. 1847. Figure 1.

Shrubs 1-3.5 m tall; young steins subquadrangular to

terete, glabrous to velutinous, leafy inter nodes 3-10 cm

long, 2.5 mm thick, lenticels oval to linear, mostly less

than 1 mm long. Leaves petiolate, petioles sessile to 5

mm long, strigose; laminae elliptic to 25 cm long, 8 cm

broad, apically acuminate, basal ly attenuate, margins en-

tire to undulate, glabrous to scabridulous above, strigose

to velutinous beneath, especially on costa and lateral

veins. Inflorescences in dense terminal spikes to 1 9 cm

long, 1 cm broad excluding corollas; peduncle subsessile;

rachis pilose; bracts imbricate to 15 mm long, 6 mm

broad, apically acuminate, basally cuneate, strigose out-

side, puberulous inside, with 2 clusters of submarginal,

mid-dorsal glandular areas with 1-10 ovular glands per

area, each gland ca. 0.5 mm long, outside of bract rather

strongly 5-nerved, with 4-8 marginal teeth to 1.5 mm

long, margins ciliate; bracteoles lanceolate, to 6 mm long,

1.5 mm broad, keeled, velutinous along the keel. Flowers

to 3 cm long, calyx segments lanceolate, to ca. 8 mm

long, anterior segments ca. 1.5 mm broad, lateral seg-

ments ca. 1 mm broad, posterior segment ca. 3 mm

broad, all segments puberulous toward apex; corolla pink,

purplish red, scarlet or red, bilabiate, to 3 cm long, grad-

ually expanding to 4 mm broad at the mouth, constricted

to 2 mm broad just above the ovary, puberulous outside,

glabrous inside, lips ovate, upper lip 2-lobed, to 1 cm

long, each lobe acuminate, 3 mm long, lower lip slightly

longer, acuminate at apex; stamens with hispid filaments.

exserted. Fruits elliptic, to 1 5 mm long, 6 mm broad, 3

mm thick, apex acute, glabrous to sparingly puberulous;

seeds 2 per locule, flattened, irregular in shape, ca. 3 mm

broad, 1 mm thick, surface minutely roughened, dark

brown.

This is a common and wide-ranging plant in

shaded areas within mature tropical dry forests or

old secondary forests, generally at lower elevations

of the western coastal areas in Costa Rica, but also

found in the Meseta Central region at elevations

as high as 1 100 m. This species grows from south-

ern Mexico to northern South America. Flowering

observed year round.

Aphelandra scabra is recognized by its pale green

(when fresh) toothed bracts with two clusters of

submarginal glands, smallish bracts (less than 1 6

mm long), short, reddish corollas (to 3 cm long),

and its subsessile leaves. It bears a slight resem-

blance to A. storkii, which is easily distinguishable,

with its longer bracts, longer orange corolla, and

much longer petioles.

Aphelandra seibertii Leonard, Ann. Missouri Bot.

Card. 24: 205. 1937. Figure 2.

Herbs to 19 cm tall; stems quadrangular, internodes

between leaf-bearing nodes 0.5-1 cm long, 2-2.5 mm

thick, pilose hairs to 1 mm long, lower stem rooting at

the nodes. Leaves petiolate, petioles to 1.6 cm long,

densely pilose; laminae elliptic, 4.5-9.5 cm long, 1.5-

4.0 cm broad, apically acute, basally acute, margins en-

tire to slightly undulate, pilose hairs to 1 mm long above

and beneath, very dense along costa and lateral veins

beneath. Inflorescences in 1-3 terminal spikes to 13 cm

long, 1 cm broad excluding corollas; peduncles 3-5 cm

long, densely yellowish pilose; rachisdensely pilose; bracts

closely imbricate, spreading at maturity, rhombic, to 14

mm long, 6 mm broad, tips often purplish, margins above

midpoint dentate with 2-3 erect teeth per side, each ca.

1 mm long; bracteoles lanceolate, to 9.5 mm long, 2 mm

broad, keeled, translucent, pilose along the keel. Flowers

15-20 mm long, falling away at maturity when bracts

are expanded; calyx segments lanceolate, subequal, ca.

5 mm long, translucent, with small glandular hairs plus

pilose hairs, posterior segment 1.4 mm broad, lateral

segments 0.5 mm broad, anterior segments 0.7 mm broad;

corolla yellow at base, lobes lavender-tipped, puberu-

lous-glandular outside, glabrous inside, bilabiate, tube

funnelform, 1 1 mm long, 2 mm broad at base expanding

to 4 mm broad at mouth, upper lip 6.5 mm long, 5.5

mm broad, apex with 2 rounded lobes 2 mm long, lower

lip 3-lobed, middle lobe retuse, 8 mm long, 5 mm broad,

lateral lobes apically rounded, 6 mm long, 2 mm broad;

stamens slightly exserted, filaments pilose. Fruits oblan-

ceolate, ca. 8 mm long, 3 mm broad, 1.5 mm thick,

reddish brown; seeds 4, subtri angular. 3 mm long, 0.7

mm thick, puberulent.

Found in rain forests in deep shade in Costa

Rica and Panama. Found in the Caribbean slope

at lower elevations in Costa Rica. Flowers through

most of the year.

Aphelandra seibertii is recognized by its low her-

baceous habit, pilose stems and leaves, small leaves

(mostly less than 1 0 cm long), toothed bracts, and

corollas with yellow bases and purplish tips. This

may be confused with Justicia chamaephyton,

which can be distinguished from A. seibertii by its

basally obtuse or rounded leaves, two stamens, less

compact spikes, and smaller entire bracts.

Aphelandra sinclairiana Nees in Benth., Bot. voy.

Sulphur 146. pi. 47. 1844. Figure 2.

Shrubs or small trees to 6 m tall; younger stems terete

to subquadrangular at the apex, internodes between leaf-

bearing nodes 1—4 cm long, 2-8 mm thick, velutinous.

Leaves petiolate, petioles 1-2 cm long, pilose; laminae

narrowly elliptic to near-oblanceolate, to 30 cm long, 8.5

cm broad, apically acuminate, basally attenuate, margins

entire to shallowly crenate, sparingly pilose above except

at costa, pilose below. Inflorescences borne in terminal

and subterminal spikes in clusters of as many as 1 0 spikes,

spikes to 15 cm long, 2 cm broad excluding corollas;

peduncles subquadrangular, to 5.5 cm long, rachis ve-

lutinous; bracts orange-red, densely imbricate, obovate-

28

FIELDIANA: BOTANY

rhombic, 16-20 mm long, 14-20 mm broad, lowermost

bracts apically acuminate, upper bracts often obtuse, 5-

6 rather prominent nerves with branching toward apex

of bract, puberulous to velutinous dorsally, puberulous

ventrally, with 2 clusters of submarginal elliptic glands

at about the middle of the dorsal surface, glands ca. 0.5

mm long, 0.25-0.4 mm broad, 2-10 glands per cluster,

margins entire; bracteoles lanceolate to ovate-lanceolate,

6.5-7.5 mm long, 3-3.5 mm broad, striate, subcarinate,

puberulous to tomentose outside, puberulous inside.

Flowers 5.5-6.5 cm long; calyx segments lanceolate, the

posterior segment 7-9 mm long, 3 mm broad, lateral

and anterior segments 7-9 mm long, 1.5-2 mm broad,

puberulous outside, glabrous inside, striate-nerved; co-

rolla purplish red, bilabiate, short-velutinous outside,

tube funnelform, 4.5-5 cm long, 3.5 mm broad at base,

6.5 mm at mouth, upper lip erect, to 1 8 mm long, 9 mm

broad, 2-lobed, the lobes acuminate, keeled apically, to

7.5 mm long, the lower lip spreading, middle lobe lan-

ceolate, keeled, to 25 mm long, 8 mm broad, lateral lobes

triangular, connate to upper lip for most of their length,

acute, free portion to 1.3 mm long, 1.5 mm broad; sta-

mens exserted to just below the tip of the upper lip and

enclosed by it. Fruits clavate, black, to 1.8 cm long, 6

mm broad, 3 mm broad at base, puberulent, apically

obtuse; seeds 4, puberulent, flattened, ca. 3.5 mm in

diameter.

According to McDade this species grows mainly

in gaps in lowland rain forests extending to pre-

montane forest and is also a successful colonizer

in more extensively disturbed areas. In Costa Rica

it has been collected mostly from such habitats in

Limon Province, with a few older collections from

Cartago and Heredia provinces. Flowering collec-

tions have been made from September through

April.

Aphelandra sinclairiana is recognized by its large,

obovate-rhombic bracts ( 1 6 to 20 mm long), which

are orange-red and have two clusters of shiny, sub-

marginal glands, its large (ca. 6 cm long), purplish

red corollas, and its often relatively tall height (to

6 m). It is easily confused with A. golfodulcensis.

(See the description of that species for a discussion

of these differences.)

Aphelandra storkii Leonard, Publ. Field Mus. Nat.

Hist., Bot. Ser. 18: 1197. 1938. Figure 2.

Shrub to ca. 2 m tall; young stems quadrangular, terete

shortly below terminus, internodes between leaf-bearing

nodes 2-4.5 cm long, 2-3.5 mm thick, pilose toward the

terminus. Leaves petiolate, petioles 1-3 cm long, hirsute

to suhglahrous; laminae elliptic to elliptic-obovate, to

44.5 cm long, 1 3.5 cm broad, apically acuminate, basally

attenuate, margins serrulate to undulate, sparingly stri-

gose above, densely strigose below, the veins prominent.

Inflorescences in terminal spikes to 18 cm long, 2.5 cm

broad excluding corollas; peduncles quadrangular, to 2.5

cm long, hirsute, rachis quadrangular, hirtellous; bracts

oblong-ovate, to 25 mm long, 1 3 mm broad, pilosulous,

with 2 clusters of 1 5-20 submarginal elliptic glands just

below midpoint of dorsal surface, glands ca. 0.5 mm

long, 0.2 mm broad, margins ciliolate, with 2-3 pairs of

teeth to 0.9 mm long near midpoint, one of which is

frequently larger than the others; bracteoles lanceolate,

to 10 mm long, 2 mm broad, apically acuminate, striate-

nerved, pilosulous. Flowers to 7 cm long; calyx segments

subequal, 12-13 mm long, posterior segment 4 mm broad,

lateral and anterior segments 2 mm broad, all apically

attenuate, pilosulous, striate-nerved; corolla orange, pi-

lose, the tube funnelform, 3 mm broad at base, 7 mm

broad at throat, upper lip erect, to 20 mm long, 1 2 mm

broad, apically 2-lobed, lobes 1 1 mm long, apically keeled

and attenuate, the middle lobe of the lower lip lanceolate,

to 22 mm long, 9 mm broad, apically attenuate and

curved back, lateral lobes triangular, connate to upper

lip for most of their length, acute, free portion to 0.8

mm long, 0.9 mm broad, stamens exserted to just below

the tip of the upper lip and enclosed by it; anthers to 8

mm long, 1 mm broad, apically adherent, basally acute;

filaments glabrous. Fruits clavate, 25-28 mm long, 7-8

mm broad, 4-5 mm thick, glabrous; seeds 4, ovate, ca.

6 mm long, 5 mm broad, glabrous.

Found in lowland rain forest. Only one collec-

tion, Opler 988 (F, NY, MO) from La Selva near

Puerto Viejo of Heredia Province, has been seen

other than the type collection made at Livingston

along the Rio Reventazon in Limon Province.

Flowering collections have been made in July and

August.

Aphelandra storkii is recognized by its long (more

than 2 cm) gland-bearing, toothed (two to three

pairs) bracts, its long orange corolla (7 cm), and

its relatively large, petiolate leaves. It bears a slight

resemblance to A. scabra, which has shorter bracts,

corollas, and leaves; its leaves are subsessile and

its corollas are reddish, but not orange.

Aphelandra tonduzii Leonard, Publ. Field Mus.

Nat. Hist., Bot. Ser. 18: 1 198. 1938. Figure 3.

Herbs to 16 cm tall, stems terete, pilose, internodes

between leaf-bearing nodes 3-30 mm long, 2-4 mm thick.

Leaves on pilose petioles 1-3 cm long; laminae elliptic

to 18 cm long, 5.5 cm broad, apically acute to obtuse,

basally attenuate, margins entire, pilose on both surfaces,

more so on costa and lateral veins. Inflorescences of loose

axillary spikes to 20 cm long, 0.5-1 cm broad excluding

corollas; peduncle subquadrangular, pilose, plus occa-

sional glandular hairs, rachis subquadrangular, pilose with

numerous glandular hairs; bracts lanceolate, slightly

keeled, 6-8 mm long, 1.25-1.5 mm broad, puberulous

with occasional glandular hairs, margins ciliolate, often

with a pair of minute, marginal teeth, bracts not imbri-

cate at anthesis; bracteoles subulate, ca. 4.5 mm long,

0.6 mm broad, puberulous with occasional glandular

hairs. Flowers to 1 3 mm long; calyx segments to 5 mm

BURGER: FLORA COSTARICENSIS

29

long, posterior segment lanceolate, 1 mm broad, lateral

and anterior segments subulate, 0.5 mm broad, all striate-

nerved, puberulous toward tip; corolla white in bud, be-

coming purplish with age, bilabiate, tube to 8 mm long,

1 mm broad at base, 1.5 mm broad at mouth, glabrous

outside, puberulous within, the 2 lobes of the upper lip

obovate, 2.5 mm long, 1 mm broad, apically retuse,

middle lobe of the lower lip rhomboid-ovate, 5 mm long,

5 mm broad, apically retuse, lateral lobes obovate, 4 mm

long, 3 mm broad, apically retuse, minutely puberulous

inside; stamens included, filaments sparingly puberu-

lous, anthers bearing a tuft of hairs apically. Fruits cla-

vate, ca. 8 mm long, 3 mm broad, glabrous; seeds 4,

obovate, flattened, 3 mm long, 2.5 mm broad, 1 mm

thick, surface hispidulous.

red, puberulous, the tube 4 mm broad at base, narrowing

to 2 mm just above the ovary, gradually enlarging to 7

mm broad at the mouth, the upper lip erect, elliptic, to

25 mm long, 8 mm broad, apically keeled, lower lip

3-lobed, middle lobe ovate, to 25 mm long, 17 mm

broad, apically obtuse, lateral lobes oblong-obovate, to

15 mm long, 6 mm broad, apically rounded; stamens

exserted to just below tips of upper lip, anthers lemon

yellow, 6.5 mm long, 1 mm broad, apically acuminate

with pilose hairs, basally acute; filaments sparingly pu-

berulous. Fruits ovate, ca. 15 mm long, 6.5 mm broad,

apically apiculate, glabrous, shiny, perianth except co-

rolla persistent; seeds orbicular, ca. 4 mm broad, surface

covered with tubercles.

Found in cloud forest at and above 1000 m in

Costa Rica and adjacent Panama. It has been col-

lected over a wider range in Costa Rica, where

most collections are from the Cordillera Central,

and others are from the vicinity of San Isidro.

Flowering collections have been made from Oc-

tober through February.

Aphelandra tonduzii is recognized by its small

size (to 16 cm tall), its small (less than 8 mm),

toothed, eglandular bracts, and its small flowers

(to 1 3 mm long). Superficially, it can be confused

with some forms of Elytraria imbricata, which

differs in its smaller, entire, and more imbricate

bracts, its smaller flowers (to 8 mm long), and its

slenderly conical capsules, which lack hooklike fu-

nicles. Aphelandra tonduzii also resembles A. ar-

noldii of Colombia and Panama, which differs with

its eglandular rachis and red flowers.

Found in cloud forests and forest clearings from

the San Jose area to Monteverde in Guanacaste at

elevations of 850-1600 m. Flowering collections

have been made from September through Decem-

ber.

Aphelandra tridentata is recognized by its long

(2.5 to 3 cm), eglandular bracts with one or two

pairs of teeth and its long (5 to 6.5 cm), bright red

corollas; it is found at higher elevations (above

850 m). Older inflorescences lacking flowers could

be confused with A. dolichantha, which has less

compact inflorescences and also differs in its lack

of toothed bracts and its longer (6.5 to 7.2 cm),

white corollas; it occurs between sea level and

1000m.

Asystasia Blume

Aphelandra tridentata Hemsl., Biol. cent.-amer.,

Hot. 2: 513. 1882. Figure 3.

Herb to subshrub. erect, to 2 m tall; young stems qua-

drangular, internodes between leafy nodes 1-6 cm long,

2-5 mm thick, pilose to wooly. Leaves petiolate, petioles

to 3.0 cm long, pilose to strigose; laminae elliptic to

elliptic-oblong, 10-18 cm long, 2.5-3 cm broad, apically

acuminate, basally attenuate, margins entire to undulate,

strigose to glabrous above, strigose below, upper surface

drying to pale green, lower surface to dark green. Inflo-

rescences in solitary, dense terminal spikes to 1 1 cm long,

3 cm broad excluding corollas; peduncle quadrangular,

to 1 3 mm long, wooly, rachis quadrangular, wooly; bracts

oblong-lanceolate, green, 3.5-4 cm long, 0.5-1.5 cm

broad, sericeous, margins entire with 1-2 pairs of teeth

above the midpoint, one conspicuously larger than the

rest, to 3.5 mm long; bracteoles linear, to 13 mm long,

1.3 mm broad, apically acuminate, striate-nerved, se-

riceous. Flowers 5-6.5 cm long; calyx segments lanceo-

late, to 1 1 mm long, puberulous, striate-nerved, poste-

rior segment 3.5 mm broad, lateral segments 1.5 mm

broad, anterior segments 2.5 mm broad; corolla bright

Herbs or shrubs. Leaves entire, petiolate, with abun-

dant cystoliths when mature. Inflorescences of secund

spikes or racemes, these loose or compact, simple or

branched. Flowers solitary on short pedicels, subtended

by 2 small bracts and 2 bracteoles of ca. equal size; calyx

5-merous, segments linear or lanceolate; corolla white,

blue, purple, rose, or yellow, tube short to funnelform,

5 lobes, subequal; stamens 4, didynamous; anthers ob-

long with 2 parallel cells, calcarate or muticous at base;

ovary 4-ovulate, pubescent; stigma minutely 2-parted or

subcapitate; capsules elliptic, 2-4 seeded; seeds com-

pressed, orbicular or irregularly angled, glabrous.

This genus is not native to the Neotropics. Ac-

cording to Leonard (1951) about 70 species have

been described which are mostly from tropical Af-

rica, Asia, and the East Indies.

Asystasia resembles the genus Dyschoriste with

which it shares most characteristics including the

few pollen grains that have been examined. They

differ in inflorescences, with Dyschoriste having

30

FIELDIANA: BOTANY

flowers borne in cymose fascicles or singly at the

axils.

More than 1 00 species of this genus have been

described, and nearly all of them occur in the Old

World Tropics.

Asystasia gangetica (L.) T. Anders in Thwaites,

Enumeratio plantarum zeylaniae 235. 1859-

1864. Justicia gangetica L., Amoen. acad. 4:

299. 1759. Figure 3.

Barleria micans Nees in Benth., Bot. voy. Sulphur

146. 1844. B. discolor Nees in Benth., loc. cit.

Figure 3.

Herbs, ascending, to ca. 30 cm tall; stems quadran-

gular, ink-modes between leaf-bearing nodes 4.5-7.5 cm

long, 1-2 mm thick, pilose. Leaves petiolate, petioles to

1 1 mm long, sparingly strigose; laminae ovate to deltoid,

3.5-5 cm long, 2-2.5 cm broad, apically acuminate, ba-

sally obtuse to truncate, margins crenulate to entire,

subglabrous to sparingly pilose especially on the veins

above and below, numerous cystoliths visible only on

the upper surface. Inflorescences in terminal, mostly se-

cund racemes, to 1 3.5 cm long; flowers on short pedicels

to 2 mm long; 4 bracteoles in 2 series subtending each

flower, lanceolate, to 2 mm long, pilose. Flowers to 4

cm long; calyx and pedicel pilose with numerous glan-

dular hairs, calyx lobes lanceolate, 5-7 mm long; corolla

yellow with purplish throat, funnelform, 3.5 mm broad

at base, 10 mm broad at throat, lobes semiorbiculate,

ca. 10 mm broad with numerous minute glandular hairs

outside, glabrous within, stamens included; anther cells

each mucronulate apically and basally. 3 mm long, 1

mm broad; ovary 3.5 mm long, style 21 mm long, ve-

lutinous, stigma 2-lohed. Fruit oblong, to 1 3 mm long,

2 mm broad; perianth persistent except early deciduous

corolla; with 2-4 flat, oval seeds ca. 1 mm long, 0.75

mm broad, oblique at point of attachment, edged with

dense mucilaginous hairs when wet.

Found in cultivation or as an escape in Costa

Rica and much of tropical America. Native to

tropical Africa, Asia, and Malaya. In Costa Rica

and Panama, it is found from near sea level to ca.

1000 m altitude. Flowering collections have been

made in March, April, and May.

Asystasia gangetica is recognized by its one-sid-

ed racemes with large (to 4 cm long), funnelform

corollas, which are yellow with purplish throats,

and its minute bracteoles (2 mm long).

Barleria L.

Shrubs or herbs, erect, sometimes spiny. Leaves op-

posite, petiolate to near sessile; laminae entire, cystoliths

generally prominent and dense on both surfaces. Flowers

axillary or borne in dense terminal bracted spikes; calyx

deeply 4-lobed, lobes unequal; corolla 5-merous, the

broad, spreading lobes mostly subequal; stamens 4, di-

dynamous, adnate to base of corolla tube, generally all

stamens perfect, but American species with only 2 per-

fect stamens plus 2 staminodes. Fruits ovate to oblong,

flattened, with 2 seeds per locule; seeds flattened, ovate

to suborbicular.

Herbs with erect stems to 75 cm tall; internodes be-

tween leaf-bearing nodes 1.6-6.2 cm long, 1.5-4 mm

thick, stems strigose, hairs appressed, to 1 mm long,

cystoliths prominent and dense, 0.3 mm long. Leaves

petiolate, petioles to 1.5 cm long, strigose; laminae lan-

ceolate to oblanceolate, 5.5-23 cm long, 1 .5-5.2 cm broad,

apically acuminate, basally attenuate, margins entire,

sparsely strigose above with cystoliths, strigose below,

cystoliths few, trichomes densest on veins and margins,

bluish along costa upon drying. Inflorescences in dense

terminal spikes to 7 cm long, 3 cm broad excluding

corollas, subsessile; bracts ovate, to 3.5 cm long, 1.5 cm

broad, apically acuminate, basally acute, chartaceous,

costa prominent, sericeous inside, strigose outside plus

occasional glandular hairs, midvein and base of each side

velutinous, often bluish upon drying, margins of spiny

hairs to 2 mm long; bracteoles oblanceolate, to 2 cm

long, sericeous with simple and glandular hairs, margins

ciliate. Flowers sessile, to 4.5 cm long; calyx segments

unequal, anterior segment elliptic, to 2.5 cm long, 1 cm

broad, apically bidentate, margin with spines to 2 mm

long, lateral segments lanceolate, to 2 cm long, 2 mm

broad, apically aristate, sericeous, posterior segment

slightly longer and narrower than the anterior segment,

apically acuminate, margin with spines to 2 mm long;

corolla yellow, drying purple, bilabiate, the tube to 3 cm

long, slender, 4 mm broad at base, lobes subequal, ob-

ovate, ca. 1.5 cm long, 6 mm broad, apically rounded,

upper lip of 1 lobe, lower lip of 4 lobes; perfect stamens

2, exserted, anthers 2 -celled, 5 mm long, filaments gla-

brous, to 4 cm long, imperfect stamens reduced to stami-

nodes ca. 0.2 mm long, staminodes with a centrally lo-

cated spur; ovary pandurate, ca. 5 mm long, style ca.

4.2-4.5 cm long, glabrous, stigma cylindrical, to 1 mm

long. Fruits elliptic to 2 cm long, 1.6 cm broad, 2 mm

thick, apically acuminate, dark purple-brown when dry,

the 4 seeds irregularly ovate, ca. 5 mm long, 0.3 mm

thick, velutinous.

Found in dry forest regions of all provinces in

the Pacific watershed from near sea level to about

600 m of the western Meseta Central. Flowering

collections made from December to April. This

species ranges from west-central Mexico to Co-

lombia.

Barleria micans is recognized by its dense ter-

minal spikes with large, conspicuous, overlapping

bracts with spiny margins, these often turning dark

bluish upon drying, and its conspicuous yellow

(drying to purple), subequal flowers to 4.5 cm long.

This species is slightly similar to Aphelandra au-

BURGER: FLORA COSTARICENSIS

rantiaca, but close examination will show all

Aphelandra species to have four fertile stamens

instead of two and bilabiate flowers.

Blechum P. Browne

Perennial herbs or shrubs. Leaves petiolate; laminae

lanceolate to ovate, entire, undulate, crenate, or serru-

late, containing numerous cystoliths on both surfaces.

Inflorescences in dense terminal, quadrifarious spikes.

Flowers 5-merous, calyx segments subequal, linear-su-

bulate; corolla white, blue, lavender or purplish, fun-

nelform with narrow tube, lobes subequal, bracts 1 per

flower, imbricate, ovate-oblong to ovate-deltoid; brac-

teoles 2, linear, oblanceolate or lanceolate; stamens 4,

didynamous, cells of anthers parallel and equal. Fruits

of elliptic capsules, flattened, dissepiment breaks away

from capsule wall at maturity, seeds few to 16, reddish

brown, flattened, ca. 2 mm in diameter, edges with a

band of sticky, dense, mucilaginous hairs when wet, gla-

brous when dry.

Native to tropical America, although species such

as B. brownei are reported from other tropical areas

as well. Leonard (195 1) states that this genus con-

sists of about six tenable species and that there is

a close relationship between Blechum and Ruellia,

but Blechum can be recognized by its dense, four-

sided spikes and very small flowers.

Key to Species of Blechum

la. Corolla small, equally or barely exceeding the bracts; bracts acute at apex B. brownei

Ib. Corolla relatively large, twice as long as the bracts or longer; bracts acuminate at apex

B. costaricense

Blechum brownei Juss., Ann. Mus. Natl. Hist. Nat.

9: 270. 1807. 1918. Ruellia blechum L., Syst.

nat. 10, 2: 1120. 1759. Barleria pyramidata

Lam., Encycl. 1: 380. 1783. Blechum pyrami-

datum (Lam.) Urb. in Repert. Spec. Nov. Regni

Veg. 15: 323. 1918. Blechum brownei f. puber-

ulum Leonard, J. Wash. Acad. Sci. 32: 942. Fig-

ure 3.

Herbs with erect to declining stems to 50 cm tall;

younger branches quadrangular, appressed-puberulent,

internodes between leaf-bearing nodes 1 .2-7 cm long, 1-

2 mm thick. Leaves petiolate, petioles to 14 mm long,

puberulent; laminae ovate to ovate-lanceolate, 2-6.5 cm

long, 1-3 cm broad, apically acute, basally cuneate to

obtuse, margins crenate to entire, both surfaces sparsely

pilose to glabrous, numerous cystoliths. Inflorescences

in terminal spikes, 4-sided, 1.5-7 cm long, peduncles to

1.5 cm long, appressed-puberulent; bracts ovate, 7.5-14

mm long, 6-1 1 mm broad, apically acute, basally round-

ed, loosely strigose and densely puberulous, margins cil-

iate; bracteoles lanceolate, 5 mm long, 1 mm broad,

ciliate; calyx segments 3 mm long, 0.5 mm broad, cil-

iolate; corolla white turning pale pink and lavender with

age, 10-15 mm long, slightly longer than the subtending

bract. Fruit obovate, ca. 6 mm long, 3 mm broad, 1 mm

thick, apically acuminate, puberulous, perianth persis-

tent except the early deciduous corolla; seeds 16, sub-

orbicular, 1 mm in diameter.

Found as a weed in disturbed areas along riv-

erbanks, roadsides, lawns, and pastures; flowering

year-round, it occurs in all parts of Costa Rica

below 1400 m elevation. Ranging from eastern

and west-central Mexico through Central and South

America to the West Indies, Guam, Taiwan, and

the Phillipine and Caroline islands.

Blechum brownei is recognized by its four-sided,

terminal spikes, its apically acute bracts, and its

small flowers, which barely extend beyond the

bracts. Closely resembling B. costaricense, it can

be distinguished from it by its shorter flowers and

acute bracts; it usually grows in a more disturbed,

open habitat. Commonly called Sornia, it is used

as a folk remedy in Costa Rica and Panama in the

form of a tea to relieve amoebic and probably

other types of dysentery.

Blechum costaricense Oerst., Vidensk. Meddel.

Dansk Naturhist. Foren. Kjobenhavn 1854: 168.

1855. Blechum dariense Lindau, Repert. Spec.

Nov. Regni Veg. 12: 423. 1913. Figure 3.

Herbs to suflrutescent at the base in older forms, to

l.lm tall; young stems quadrangular, pilose, internodes

between leaf-bearing nodes 2.2-6.5 cm long, 1-2 mm

thick. Leaves with petioles to 2 cm long, pilose; laminae

ovate to ovate-lanceolate, 2-11 cm long, 1-5 cm broad,

apically acuminate, basally attenuate, margins undulate

or serrulate, both surfaces pilose to sparingly pilose, nu-

merous cystoliths visible on both surfaces. Inflorescences

in terminal spikes, 4-sided, to 5.5 cm long, peduncles to

2 mm long, pilose; bracts ovate-deltoid, apically cuspi-

date or acuminate, basally rounded, to 2 cm long, 1.5

32

F1ELDIANA: BOTANY

cm broad, sparingly pilose outside and inside, margins

ciliate; bracteoles linear to oblanceolate, 7-10 mm long,

0.5-1 mm broad, pilose margins, ciliate; calyx segments

equal, ca. 5 mm long, 0.75 mm broad at base, ciliate;

corolla 2-2.5 cm long, pale lavender to purple, extending

well beyond the subtending bract. Fruit elliptic, ca. 8

mm long, 5 mm thick, apically acuminate, perianth per-

sistent except the soon deciduous corolla; seeds subor-

bicular, ca. 2 mm in diameter.

Found in Costa Rica in rain forests of the Ca-

ribbean watershed area, the Pacific slopes of Valle

del General area to the Cordillera Tilaran, at el-

evations of 200 to 1000 m. This species is also

found rather commonly in adjacent areas of Pan-

ama to sea level and in lowland areas in the Canal

Zone. Flowering collections have been made from

September through March.

Blechum costaricense is recognized by its four-

sided, terminal spikes, its broad, acuminate bracts,

and its corollas, which are twice as long as the

bracts. Closely resembling Blechum brownei, it can

be distinguished by its larger flowers and acumi-

nate bracts.

Bravaisia DC.

Shrubs or trees. Leaves petiolate; laminae ovate to

elliptic, apically acute to acuminate, margins entire to

slightly undulate. Inflorescences terminal in compact,

leafy panicles. Flowers subtended by a pair of bracts;

calyx 5-merous, subcoriaceous; corolla white, yellow, or

purplish with a short tube and campanulate throat, 5

lobes, subequal; stamens 4, anther cells 2, contiguous,

spurred at the base. Fruit a capsule with 4-8 seeds.

A genus of the Neotropics, Bravaisia has five

species variously reported from Mexico, Central

America, Colombia, Venezuela, Peru, and the West

Indies.

Bravaisia integerrima (Spreng.) Standley, Contr.

U.S. Natl. Herb. 23: 1335. 1926. Amasonia in-

tegerrima Spreng., Syst. veg. ed. 16. 2: 765. 1825.

OnchyacanthusspeciosusOersl., Vidensk. Med-

del. Dansk Naturhist. Foren. Kj0benhavn 1854:

131. 1855. Figure 4.

Tree to 20 m tall; branchlets quadrangular, internodes

between leaf-bearing nodes 2.5-7 cm long, 2.5-5 mm

thick, glabrous to tomentose on youngest portions. Leaves

with petioles to 4 cm long, pubescence of petioles like

that of the stem; laminae ovate to ovate-elliptic, 3.5-

28.5 cm long, 1-1 1.8 cm broad, apically acute or acu-

minate, hasally acute or oblique, margins entire to slight-

ly undulate, subcoriaceous, surfaces glabrous, sparingly

pilose at costa and lateral veins. Inflorescences in leafy,

compact, terminal panicles, to 13 cm long, 25 cm wide,

peduncles quadrangular. Flowers subtended , - oval

bracteoles which are ca. 2 mm long, apically obtuse,

puberulent and ciliolate; calyx segments subequal, spat-

ulate, 5-6 mm long, 2-3 mm broad, puberulous and

ciliolate; corolla white or yellow with a purple spot in

the throat, 1 .5-2.0 cm long, glabrous, tube campanulate,

ca. 8 mm broad at the mouth, the limb ca. 2 cm across,

lobes rounded or emarginate, ca. 10 mm long, 6-8 mm

broad. Fruits clavate, flattened, 10-12 mm long, 4-5 mm

broad, 3 mm thick, glabrous, perianth persistent; seeds

8, orbicular, 3.5 mm in diameter.

Plants of evergreen and deciduous forest for-

mations, ranging from 100 to 600 m in the Pacific

coastal regions to the Meseta Central and the Llan-

ura de San Carlos. Flowering collections have been

made from December through May. The species

ranges from Peru, Colombia, Venezuela, and north

to Oaxaca and Tamaulipas, Mexico.

Bravaisia integerrima is exceptional among the

Acanthaceae for its large size. In addition to its

large size, it is recognized by its flowers with four

stamens having anthers with basal spurs. It also

has eight-seeded capsules. It most closely resem-

bles another woody species, Trichanthera gigan-

tea, which has anthers with rounded bases and

four-seeded capsules. Commonly called palo de

agua in Costa Rica, it has a light-colored wood

that is suitable for packing boxes and food pails.

Numerous aerial roots similar to mangroves are

often produced at the bases of the trunks (Leonard,

1951).

Buceragenia Green man

Herbs; stems erect. Leaves petiolate; laminae usually

ovate or elliptic. Inflorescences spicate or spicate pani-

cles, the flowers solitary or in clusters; bracts and brac-

teoles minute; calyx 5-merous, segments linear, sub-

equal; corolla tubular, shallowly bilabiate, posterior lip

in-curved, emarginate or 2-lobed. anterior lip erect or

slightly spreading, 3-lobed; stamens 2, inserted at middle

of corolla tube, included; anthers usually 1 -celled, the

lower cell aborted or vestigial and sterile, if fertile, always

considerably smaller; staminodes 2; fruits a clavate cap-

sule, stipe narrow; seeds 4, flattened, suborbicular.

This is a genus of four species which extend from

Mexico to Costa Rica.

BURGER: FLORA COSTARICENSIS

33

Buceragenia glandulosa Leonard, Publ. Field Mus.

Nat. Hist., Bot. Ser. 18: 1205. 1938. Figure 4.

Herbs to 1.5 m tall, stem erect, branching, subqua-

drangular, glabrous to sparingly puberulous; internodes

between leaf-bearing nodes 1.5-4 cm long, 2.5-3 mm

thick. Leaves with petioles to 3 cm long, those subtending

inflorescences subsessile, pubescence like that of the stem;

laminae ovate to ovate-oblong, 5.5-18 cm long, 2-7.5

cm broad, apically acuminate, basally attenuate except

the leaves immediately subtending the inflorescence

which are basally rounded to cordate, margins entire,

costa often reddish purple, glabrous. Inflorescences in

terminal and axillary spikes or spicate panicles to 20 cm

long, flowers sessile or nearly so, borne in fascicles of 2-

4 or occasionally singly; bracts subulate, 1.5 mm long,

0.5 mm broad, glabrous, bracteoles similar, but smaller;

rachis glandular-puberulous; calyx 5-merous, 3 mm long,

lobes subulate, 2 mm long, 0.5 mm broad at base, gla-

brous; corolla small, tubular, glabrous, 3-3.5 mm long,

1 mm broad, lobes 5, more or less equal, suborbicular,

0.5 mm long, 0.25 mm broad, stamens 2, included, 1

mm long, anthers usually 1 -celled, other cell usually

aborted or vestigial and sterile, if fertile, always much

smaller, 1 or both cells mucronate. Fruits clavate, 1 3-

19 mm long, glabrous, stipe 6.5-10 mm long, 1.5 mm

broad, expanded portion 6.5-9 mm, 2.5 mm broad, seeds

flat, oval, 3 mm long, 2 mm broad, roughened.

Found in cloud forests at 1200 to 1800 m, this

species is reported from Mexico, Guatemala, Nic-

aragua, and Costa Rica. Only three collections have

been observed from Costa Rica, all from Cartage

Province: Holway 400 (us, holotype), Standley

33686 (us), and Standley 35503 (us). All collec-

tions were made in January or February, and all

were in both fruit and flower.

Buceragenia glandulosa is recognized by its her-

baceous habit with terminal and axillary spikes or

spicate panicles, glandular-puberulous rachises,

inconspicuous flowers in clusters of two to four or

occasionally single, small, five-merous, mostly

equal corollas 3 to 3.5 mm long with two stamens

and two staminodes, anthers two-celled, with one

cell usually aborted or vestigial, sterile, and much

smaller; leaves immediately subtending the inflo-

rescence sessile with ovate laminae and rounded

to cordate bases. It resembles Pseuderanthemum

cordatum, but is distinguished from it by its mi-

nute flowers and inflorescences with glandular-

puberulous rachises (vs. puberulous). It also re-

sembles P. cuspidatum, but may be distinguished

from it by its minute flowers which occur in fas-

cicles (vs. solitary flowers which are much larger).

Carlowrightia Gray

REFERENCES— A. Gray, Contributions to the

botany of North America. Proc. Amer. Acad. Arts

13: 361-374. 1878. T. F. Daniel, Carlowrightia

(Acanthaceae). Flora Neotropica 34: 1-116. 1983.

Subshrubs and suffrutescent herbs. Leaves opposite,

sessile to petiolate; laminae linear to ovate, entire. Flow-

ers in axillary clusters or more commonly in spicate

panicles. Flowers with a 5-merous calyx; corollas 4-part-

ed, ranging from nearly regular to strongly zygomorphic,

the lower lobe somewhat keeled and containing the an-

thers and style in bud, upper lobe spatulate and banner-

like, often marked with a distinctively colored eye and/

or radiating nerves; stamens 2, these follow the lower lip

out the corolla tube, anther 2-celled with the cells equal

and nearly parallel, opening toward the upper lip. Fruits

with flattened stipes, compressed-ovoid, acuminate heads;

seeds usually 4, ovoid, disklike, smooth, muricate to

tuberculate, apically rounded and basally obliquely

notched.

This is a genus endemic to the New World, with

20 species ranging from the southwestern United

States to Costa Rica.

Carlowrightia arizonica A. Gray, Proc. Amer.

Acad. Arts 13: 361-374. 1878. Carlowrightia

costaricana Leonard, Publ. Field Mus. Nat. Hist.,

Bot. Ser. 18: 1206. 1938.

Erect, suffrutescent plants to ca. 60 cm tall; internodes

between leaf-bearing nodes 2-4 cm long, 3.5-4.5 mm

thick, terete or subterete, retrorse pubescence. Leaves

with short, puberulous petioles 2-6 mm long; laminae

ovate, 3-4.5 cm long, 1.1-2.3 cm broad, apically acu-

minate, basally obtuse to rounded, margins entire to

undulate, puberulous above and beneath, cystoliths vis-

ible on both surfaces, more obscure below. Inflores-

cences in lax terminal and subterminal spicate panicles,

rachis puberulous; flowers sessile, mostly 1 per node, but

occasionally 2 per node and opposite or 2 per bract;

bracts and bracteoles subulate, 2-2.5 mm long, 0.5 mm

broad, puberulous. Flowers with subulate, puberulous

calyx segments, 1.2-1.5 mm long, ca. 0.5 mm broad;

corolla bilabiate, puberulous, 9-15 mm long, ca. 1 mm

broad basally, drying purplish, white with a yellowish

eye bordered by purplish radiating lines on the upper

lobe, narrowly oblong lobes twice the length of the tube;

stamens exserted. Fruits clavate, 9-10 mm long, ca. 3

mm broad, 1 mm thick, apically attenuate, the slender

stipe ca. half the length of the fruit, capsule dark purple

or black when dry, glabrous; seeds 4, cordate, ca. 3 mm

long, 2.8 mm broad, drying purplish, papillose.

This species extends from Texas and Arizona

to Costa Rica. Only one collection from Costa

Rica has been observed, Standley & Valeria 44890

(us). This was in dry forest in the vicinity of Li-

bano, Guanacaste, at an elevation of 260 to 360

m. Flowering during January. In other areas it is

34

FIELDIANA: BOTANY

often found in rocky thickets with dry soil at sim-

ilar elevations.

Carlowrightia arizonica is recognized by lax spi-

cate panicles, bilabiate corollas with exserted sta-

mens that have near equally attached anther cells,

dark purple or black capsules when dry that have

a narrow stipe nearly equal to half the capsule

length, and short-petioled leaves. It may be con-

fused with either Justicia comata or Justicia pec-

toralis, but these differ in their superposed anther

cells and inflorescences with glandular puberulous

rachises.

Chaetochlamys Lindau

Plants of deciduous forest formations in the Pa-

cific coastal area at elevations of near sea level to

1 000 m in Tilaran and San Ramon and one col-

lection from near the Panamanian border in Li-

mon Province, Hazlett 5207 (F). Also found in

similar habitats of Panama. Flowering collections

have been made from December through March.

Chaetochlamys panamensis can be recognized

most easily by its inflorescences in sessile, terminal

heads with conspicuous ciliate, linear bracts. It

most closely resembles Chaetothylax leucanthus,

but can be distinguished from it by its longer co-

rolla (more than 20 mm long vs. 15 mm), five-

parted calyx (vs. four-parted), and longer capsules

(11 to 12 mm long vs. 6 mm).

Herbs to suffrutescent plants. Leaves petiolate; lami-

nae ovate to lanceolate. Flowers borne in short spikes

that are crowded into a terminal, congested head; bracts

and bracteoles narrow; calyx segments 5, equal, narrow;

corolla salverform, bilabiate, upper lip 2-lobed, lower lip

3-lobed; stamens 2, exserted, anthers 2-celled, cells at-

tached at different heights, the lower cells spurred. Fruits

a clavate capsule; seeds 4.

Chaetochlamys is native to the Neotropics, with

at least seven species known from South America

and Central America.

Chaetochlamys panamensis Lindau, Report . Spec.

Nov. Regni. Veg. 11: 124. 1912.

Herbs to 60 cm tall; steins branching, erect or decum-

bent, sometimes rooting at lower nodes, internodes be-

tween leaf-bearing nodes 2-4 cm long, 1-1.5 mm thick,

terete, glabrous to puberulous along 2 lines. Leaves with

petioles subsessile to 3 mm long, petioles puberulous to

glabrous; laminae lanceolate-elliptic, 2.5-10.5 cm long,

1. 1-3.2 cm broad, apically acuminate, basally attenuate,

margins entire to undulate, cystoliths dense, prominent

on both surfaces, glabrous to sparingly puberulous along

costa and veins. Inflorescences in terminal heads, sessile;

bracts linear, to 22 mm long, 0.75 mm wide, ciliate, the

hairs to 1 .5 mm long; bracteoles similar, but shorter and

narrower. Flowers sessile; calyx segments lanceolate, 10

mm long, 1 .5 mm wide, ciliate; corolla pink or purplish,

sparingly glandular-puberulous, to ca. 4 cm long, the tube

cylindrical ca. 22 mm long, 4 mm broad, the upper lip

oblong, 10 mm long, 3 mm broad, apically 2-lobed, the

middle lobe orbicular, 10 mm in diameter, the lateral

lobes elliptic, 1 2 mm long, 8mm broad; stamens exserted

to just above the mouth, anther cells unequal, inserted

at different heights, the upper cell 2 mm long, basally

acute, lower cells 2.5 mm long, basally subcalcarate, the

connective ca. 1 mm broad. Fruits stout-clavate, 11-12

mm long, 4.5-5 mm broad, 2 mm thick, glabrous, peri-

anth persistent except the early deciduous corolla; seeds

4, suborbicular, not flattened, 2-2.5 mm in diameter.

Chaetothylax Nees

Herbs. Leaves petiolate; laminae ovate to elliptic.

Flowers more or less secund, borne in short, dense ter-

minal and axillary spikes which often form panicles;

bracts narrow with filiform tips, puberulous with ciliate

margins; calyx 4-merous, segments resembling the bracts;

corollas white to purplish, puberulous, bilabiate, lips

suhcqual, upper lip narrowly ovate, lower lip deeply

3-lobed, the lobes rounded; stamens 2, slightly exserted,

not exceeding the corolla lips, anthers 2-celled, upper

cell oblong, lower cell reduced or vestigial or, as in the

type species, hairlike, the lobes separated by a slender

filiform connective. Fruit a clavate capsule; seeds 4, pa-

pillose or puberulous.

Chaetothylax is native to the Neotropics, oc-

curring in Central and South America. There are

at least eight species known.

Chaetothylax leucanthus Leonard, Contr. U.S.

Natl. Herb. 31: 663. 1958. Figure 4.

Herbs to 60 cm tall; stems erect, terete with retrorse

hairs along 2 lines, internodes between leaf-bearing nodes

1.5-3 cm long, 1-1.5 mm thick. Leaves with petioles

short, to 5 mm long, strigose; laminae ovate-elliptic to

elliptic, 2-10 cm long, 0.8-2.8 cm broad, apically cus-

pidate, basally attenuate, margins entire to crenate, gla-

brous above, cystoliths dense and prominent above. In-

florescences in compact, headlike terminal and axillary

spicate panicles, spikes to 4 cm long, 1 cm wide, spikes

dense, more or less 1 -sided; panicles subtended by small

leaves; bracts narrowly subulate, to 9 mm long, 0.5 mm

broad, ciliate and moderately puberulous. Flowers with

4-merous calyx, segments narrowly subulate, to 1 1 mm

long, ca. 0.4 mm wide, ciliate and minutely glandular-

puberulous; corolla white, ca. 14 mm long, puberulous

except at base, the tube cylindrical, 9 mm long, 0.5 mm

broad, the upper lip ovate-oblong, 3.5 mm long, 1 mm

BURGER: FLORA COSTARICENSIS

35

broad, apically emarginate, the lower lip 3-lobed, ca.

3.25 mm long, the lobes oblong, 2 mm long, the middle

lobe 1.5 mm broad, the lateral lobes 1.25 mm broad, all

apically obtuse; stamens exserted ca. half the length of

the upper lip, the upper anther cell 0.75 mm long, lower

cell attached 0.5 mm below the upper cell, reduced, ster-

ile, filaments glabrous. Fruit clavate, 6 mm long, 1 mm

broad, 2 mm thick, puberulous, perianth persistent; seeds

4, orbicular, slightly flattened, 1 mm in diameter, 0.5

mm thick, puberulous.

Plants of clearings, roadsides, and trails in de-

ciduous forest formations in Guanacaste and Ala-

juela from near sea level to 500 m elevation. While

only two collections (both by Brenes s.n.) have

been observed from Costa Rica, it is assumed that

this species is more common than this low number

suggests, since it is rather inconspicuous and can

easily be mistaken for another species which is

more frequently collected.

Chaetothylax leucanthus can be recognized most

easily by its inflorescences in sessile, headlike pan-

icles. It is easily mistaken for Chaetochlamys pan-

amensis, but can be distinguished from it by its

shorter corolla and capsule and its four-parted ca-

lyx.

Chamaeranthemum Nees

Herbs or small shrubs. Leaves petiolate to sessile; lam-

inae lanceolate to ovate-lanceolate, entire to undulate,

cystoliths dense and prominent on both surfaces. Inflo-

rescences in loose terminal spikes or spicate panicles.

Flowers sessile or pedicellate, 5-merous with inconspic-

uous bracts and bracteoles; corolla salverform, lobes sub-

equal; stamens 4, included, anterior stamens with 2-celled

anthers, posterior stamens with 1 -celled anthers or ster-

ile; capsules 2-4 seeded, with a long, narrow stipe.

Chamaeranthemum is native to tropical Amer-

ica with eight species reported from Costa Rica

and northern South America.

Key to Species of Chamaeranthemum

la. Leaves, stems, and rachises hirsute .

1 b. Leaves, stems, and rachises glabrous

C. durandii

.C. tonduzii

Chamaeranthemum durandii Leonard, Publ. Field

Mus. Nat. Hist., Hot. Ser. 18: 1208. 1938. Fig-

ure 4.

Herb to 56 cm tall; younger stems quadrangular, older

stems terete, internodes between leaf-bearing nodes 2.5-

7.5 cm long, 1-1.5 cm broad, hirsute. Leaves petiolate

to sessile, petioles to 6 mm long, pubescence like that of

the stem; laminae ovate-lanceolate, 4.5-10 cm long, 1 .6-

4.5 cm broad, apically acuminate, basally rounded to

obtuse, margins entire to undulate, sparingly pilose above,

pilose below, cystoliths dense and prominent on both

surfaces. Inflorescences of loose terminal spicate pani-

cles, to 1 5 cm long, pubescence of peduncle and rachis

that of the stem plus minute puberulence; bracts and

bracteoles subulate ca. 1.5-2.0 mm long, 0.5 mm broad,

puberulous. Flowers sessile, to 2 cm long; calyx segments

linear-subulate, 2.5-3 mm long, 0.5 mm broad, puber-

ulous-glandular; corolla pink or lavender, tube salver-

form, puberulous-glandular, 10-12 mm long, ca. 1 mm

broad, expanding to 1.5 mm broad at midpoint, lobes

5, subequal, oblong, ca. 8 mm long, 5 mm broad, apically

rounded; stamens 4, included, anthers of anterior sta-

mens 2-celled, ca. 1.5 mm long, 0.5 mm broad, apically

and basally acute, filaments short, 1 mm long, glabrous,

posterior stamens sterile, 1.1 mm long. Fruits clavate,

1 5 mm long, 3 mm broad, 2 mm thick, apically apiculate,

puberulous; seeds 2—4, flattened, suborbicular, ca. 2 mm

in diameter, slightly winged, glabrous.

Found in Costa Rica in the shade of both trop-

ical moist and wet forests of the Pacific watershed

ranging from Valle del General and Valle del Di-

quis to Quepos at elevations of 30 m to ca. 800

m. Flowering collections have been made from

January, February, and March.

Chamaeranthemum durandii can be recognized

by its lavender corollas with five equal lobes and

a narrow cylindrical tube with four short, included

stamens, the posterior pair being sterile. It can be

distinguished from C. tonduzii by its hirsute leaves,

stems, and rachises.

Chamaeranthemum tonduzii Lindau, Anales Inst.

Fis.-Geogr. Nac. Costa Rica 9: 188. 1898, and

in Pitt., Prim. fl. costaric. 2: 303. 1900.

Herb to 42 cm tall; younger stems quadrangular, older

stems terete, internodes between leaf-bearing nodes 1 .6-

4 cm long, 1 mm broad, puberulous along 2 lines. Leaves

petiolate to sessile, petioles to 5 mm long, pubescence

like that of the stem; laminae lanceolate to lanceolate-

ovate, 3-7.5 cm long, 1.5-3 cm broad, apically acumi-

nate, basally acute, margins entire to undulate, glabrous

above and below, cystoliths prominent and abundant on

36

FIELDIANA: BOTANY

both surfaces. Inflorescences of loose, terminal spikes or

spicate panicles to 10 cm long, peduncle and rachis gla-

brous; bracts and bracteoles subulate, to 2 mm long, 1

mm broad, puberulous. Flowers sessile or with pedicels

to 2 mm long; calyx segments linear-subulate, 1.5-2.5

mm long, 0.5 mm broad, glandular-puberulous; corolla

color not observed, salverform, tube ca. 6.5 mm long,

0.5 mm broad at base expanded to 1 .25 mm broad above

midpoint, puberulous, lobes 5, subequal, oblong, api-

cally obtuse, ca. 4.5 mm long, 2.5 mm broad, puberu-

lous; stamens 4, included, anthers of anterior stamens

2-celled, apically and basally acute, ca. 1.5 mm long,

0.75 mm broad, filament 0.5 mm long, glabrous, pos-

terior stamens sterile, 0.75 mm long. Fruits clavate, ca.

14 mm long, 3 mm broad, 1 mm thick, apically apiculate,

puberulous to subglabrous; seeds 4, suborbicular, flat-

tened, ca. 2 mm in diameter, including a wing ca. 0.5

mm broad surrounding the seed, glabrous.

resembles C. durandii and can be distinguished

from it by its glabrous leaves, stems, and rachises.

Dicliptera Juss.

Herbs or shrubs, stems more or less hexagonal in cross

section. Leaves petiolate; laminae mostly ovate with en-

tire or undulate margins. Inflorescences of cymes, spikes,

panicles, or thyrses with flowers or flower clusters sub-

tended by 2-4 pairs of conspicuous involucrate bracts.

Flowers sessile or pedicellate, often showy; calyx 5-mer-

ous, hyaline; corollas narrow, slightly funnelform, bila-

biate; stamens 2, anthers 2-celled, cells often unequal;

staminodes absent. Fruits of ovate, elliptic, or clavate

capsules; seeds 2 or 4.

Found in Costa Rica in tropical moist forests at

lower elevations of the Caribbean watershed, with

collections ranging from the Panamanian border

to Livingston on the Rio Reventazon. Flowering

collections have been made from March and July.

Chamaeranthemum tonduzii can be recognized

by its salverform corollas, four short, included sta-

mens, with the posterior pair being sterile. It closely

A total of 1 50 species of New and Old World

Tropics have been described for this genus, which

is most easily recognized by its hexagonal stems

and reduced cymes with bracts. It has been con-

fused with Blechum and Tetramerium, but can be

distinguished from these by the above character-

istics.

Key to Species of Dicliptera

la. Inflorescences of axillary cymes; corollas more than 2 cm long 2a

Ib. Inflorescences of spikelike racemes or thyrses or axillary heads with peduncles; corollas less than 2

cm long 5a

2a. Inflorescences with peduncles less than 2 cm long; capsules up to 8 mm long D. pallida

2b. Inflorescences with peduncles more than 2 cm long; capsules more than 10 mm long .... 3a

3a. Floral bracts lanceolate and glandular-puberulous; flowers subsessile, pedicels ca. 1 mm long

D. skutchii

3b. Floral bracts oblong, sparingly puberulous, or glabrous; flowers pedicellate, pedicels more than

2 mm long 4a

4a. Floral bracts apically rounded and apiculate; capsules puberulous; corolla puberulous

D. trifurca

4b. Floral bracts apically acute to obtusish; capsules glandular-puberulous; corollas glandular-pu-

berulous D. iopus

5a. Corollas white with lavender stripes; floral bracts to 7 mm long D. imbricata

5b. Corollas pinkish or lavender; floral bracts 9-14 mm long 6a

6a. Floral bracts spatulate, awned; inflorescence of spikelike racemes and/or axillary umbels

D. unguiculata

6b. Floral bracts obovate or elliptic, not awned; inflorescence of peduncled axillary heads

D. podocephala

Dicliptera imbricata Leonard, Publ. Field Mus.

Nat. Hist., Bot. Ser. 18: 1210. 1938. Figure 4.

Herb to 1 m tall; stems branched, sometimes rooting

at lower nodes, internodes between leaf-bearing nodes

3-6 cm long, 1.5-2.5 mm thick, puberulous mostly at

the angles in younger portions, basal portions terete and

glabrous. Leaves with petioles to 4 cm long, puberulous;

laminae ovate-elliptic, 3-10 cm long, 1-4 cm broad,

apically acuminate, basally attenuate, margins entire,

BURGER: FLORA COSTARICENSIS

37

glabrous, cystoliths dense and prominent on both sur-

faces. Inflorescences of terminal and axillary spikelike

thyrses, to 4 cm long, 1 .6 cm broad excluding corollas,

2 fanlike cymes of 3-5 flowers each at every node, each

cyme subtended by a pair of oblong-lanceolate bracts 3-

8 mm long, 0.5-1 mm wide, puberulous; bracts sub-

tending the flowers unequal, obovate, to 7 mm long, 3.5

mm broad, whitish hasally, greenish apically, strongly

3-nerved, puberulous with ciliate margins; outer pair of

bracteoles linear, slightly falcate, 5 mm long, 0.75 mm

broad, apically acuminate, ciliate, inner bracteoles lan-

ceolate, 4 mm long, 0.5 mm broad, ciliolate. Flowers to

16 mm long; calyx segments linear-subulate, 3 mm long,

0.25 mm broad, minutely puberulous; corolla white, lav-

ender-striped, puberulous, the tube cylindrical, 10 mm

long, 1 mm broad, the lips oblong, 6 mm long, 2 mm

broad, apically minutely dentate; stamens exserted to

near the tips of the lips, filaments to ca. 5 mm long,

puberulous; anther cells superposed, basally obtuse. Fruits

elliptic, ca. 4 mm long, 1 mm broad, puberulous toward

tip; seeds 2, oval, flattened, ca. 1 mm long, roughened.

Found in clearings and along trails in deciduous

forest of Pacific watershed in Alajuela and Guan-

acaste provinces at elevations of 600 to 2000 m.

Flowering collections have been made from late

December through March.

Dicliptera imbricata is recognized by its dense,

spikelike thyrses, its small (7 mm long), obovate

bracts subtending each flower, and its small (16

mm long), white corollas. It is most easily confused

with D. unguiculata, which has spine-tipped bracts

and rose-purple corollas.

Dicliptera iopus Lindau, Anales Inst. Fis.-Geogr.

Nac. Costa Rica 9: 1 89. 1 898, and in Pitt., Prim,

fl. costaric. 2: 305. 1900. Figure 4.

Shrub to 3 m tall; internodes between leaf-bearing

nodes 2-6 (9) cm long, 1.0-2.5 mm thick, strigose to

glabrous. Leaves with petioles to 7 cm long, glabrous to

sparingly puberulous; laminae ovate to elliptic, 3-15 cm

long, 1-5.6 cm broad, apically acuminate, basally acute,

margins entire, glabrous to sparingly strigose on both

surfaces, cystoliths visible on both surfaces. Inflores-

cences in axillary cymes, each cyme subtended by 2 el-

liptic to ovate-oblong bracts, to ca. 3 cm long, 1 cm

broad, apically acute, basally acute, glabrous to sparingly

pilosulous; 2 bracts of unequal length subtending each

flower, both linear-oblong, fused at base, the longer one

10-16 mm long, 4 mm broad at base, the shorter bract

to 1 2 mm long, 4 mm wide, both apically acute, keeled,

sparingly puberulous on both surfaces; 4 bracteoles, ob-

lanceolate, to 8 mm long, 2 mm broad, puberulous to-

ward tip, inner pair smaller, 3.5 mm long, 0.5 mm broad;

peduncles to ca. 6.5 cm long, subhexangular, puberulous

along ridges, pedicels of middle pair in a cyme longest,

to ca. 4 mm long, sparingly puberulous. Flowers usually

3 per cyme, although some have 2 or 4, 4-5 cm long;

calyx 5-merous, coalesced for ca. V* of its length, to 7

mm long, lobes linear, ca. 3.5 mm long, subtomentose;

corolla lavender, inflated at about the middle of the tube,

bilabiate, tube to 4 cm long, 2.5 mm broad at the base,

8 mm broad at the mouth, glandular-puberulous both

inside and out, upper lip entire, acute, ca. 9 mm long, 5

mm broad, lower lip ca. 1 1 mm long, 8 mm broad at

base, with 3 small, suborbiculate lobes at its apex, ca.

0.5 mm long; stamens exserted, filaments flattened, pi-

lose, anther cells equal; style exserted, glabrous. Fruits

clavate, ca. 1 5 mm long, apically apiculate, densely glan-

dular-puberulous, 2 seeds per capsule, flattened, subor-

bicular, ca. 3 mm in diameter.

Found rather commonly over a wide range in

Costa Rica in open meadows and cloud forests at

elevations of 1300 to 3200 m on both the Carib-

bean and Pacific slopes; also common in Panama

and probably Nicaragua. Flowering collections

have been made from September through March.

Dicliptera iopus is recognized by its peduncled

cymes usually in clusters of three, acute to obtusish

floral bracts, subtomentose pedicels, and glandu-

lar-puberulous corolla and capsule. It is easily con-

fused with D. trifurca, which differs in its rounded

and apiculate floral bracts, glabrous pedicels, and

puberulous corollas and capsules.

Dicliptera pallida Leonard, Publ. Field Mus. Nat.

Hist., Hot. Ser. 18: 1211. 1938. Figure 5.

Herb to 1.2 m tall; stems erect, branched, internodes

between leaf-bearing nodes 4-10 cm long, 1.5-3.5 mm

thick, subtomentose. Leaves with petioles to 1 .6 cm long,

subtomentose to glabrescent; laminae ovate, 3.5-8 cm

long, 2-3.5 cm broad, apically acuminate, basally acute

and slightly tapering, margins undulate, sparingly pilose

above and below, cystoliths visible, but not prominent.

Inflorescences in sessile or subsessile, axillary cymes,

peduncles to 7 mm long, subtomentose; flowers of the

upper cymes mostly sessile, those of the lower, short-

pedicelled, the pedicel of the middle flower longer than

that of the lateral flowers, all pedicels tomentose; bracts

subtending the cymes, leaflike, 2-3 cm long, 1-1.5 cm

broad, those subtending the flowers unequal, elliptic to

oblanceolate, the larger of the pair ca. 16 mm long, 5

mm broad, the shorter ca. 13 mm long, 2.5 mm broad,

both bracts apically acuminate, whitish toward the base,

hirtellous; bracteoles unequal, both narrowly lanceolate,

the longer to ca. 1 2 mm long, 1.25 mm broad, the shorter

slightly smaller, both ciliolate. Flowers 2.5-3.5 cm long;

calyx to ca. 5 mm long, fused ca. 'A of length, segments

lanceolate, ca. 3 mm long, 0.75 mm broad, minutely

ciliolate; corollas red, puberulous, the tube ca. 2 cm long,

2 mm broad at the base, narrowing to 0.5 mm broad

just above the ovary, expanding to 3 mm broad at the

mouth, the lips ca. 8 mm long, the upper emarginate,

the lower minutely 3-lobed; stamens exserted nearly to

the tip of the upper lip. Fruits clavate, to 8 mm long, 5

mm broad, 1 mm thick, puberulous; seeds discoid, ca.

3 mm in diameter, puberulous.

38

FIELDIANA: BOTANY

Found at middle to higher elevations in thickets

and along streams on the slopes of the Cordilleras

Central and Talamanca from 635 to 1700 m al-

titude. This species is apparently endemic to Costa

Rica. Flowering collections have been made from

December through February.

Dicliptera pallida is recognized by its axillary

cymes with sessile to subsessile flowers, floral bracts

and bracteoles with whitish bases, and compara-

tively long (more than 2 cm) red corollas. It can

be distinguished from the much more common

and weedy D. unguiculata, which it most closely

resembles, by these characteristics.

Dicliptera podocephala Donnell Smith, Hot. Gaz.

(Crawfordsville) 48: 299. 1909. Figure 5.

Herb to subshrub, to 1.5m tall; stems erect, branching,

internodes between leaf-bearing nodes 2-4 cm long, 1-

2 mm thick, glabrous. Leaves with petioles to 2.5 cm

long, puberulous; laminae ovate, 2.5-10 cm long, 1-4

cm broad, apically acuminate, basally acute, slightly ta-

pering, margins entire, cystoliths dense and prominent

on both surfaces, glabrous. Inflorescences in peduncled

axillary heads, 1 -several per axil, peduncles to 9.5 cm

long, minutely puberulous mostly along 2 lines; bracts

subtending the heads subulate, 3.5-5 mm long, 1 mm

broad, glabrous; fertile flowers usually 5-7 per head, bracts

subtending the flowers obovate to elliptic, 1 of each pair

slightly larger than the other, the middle pair the largest,

10-14 mm long, 8-9 mm broad, apically rounded or

obtuse, apiculate, narrowed at the base, subglabrous,

margins ciliolate; bracteoles 4, linear-lanceolate, 5-8 mm

long, puberulous. Flowers 14 mm long; calyx segments

resembling bracteoles, slightly shorter; corolla pinkish

lavender, 14 mm long, puberulous; stamens exserted.

Fruits ovoid, 6 mm long, 4 mm broad, 1 mm thick,

puberulous; seeds discoid, 2.5 mm in diameter, puber-

ulous.

Found in damp meadows, streamsides, and along

trails in Cartago and Puntarenas provinces near

1000 m elevation and at 90 m elevation at La

Selva in Heredia Province. Only two collections,

Allen 5902 (F) and Opler 597 (F), have been ob-

served other than the type collection. Flowering

collections have been made in February and March.

Dicliptera podocephala is recognized by its pe-

duncled axillary heads with five to seven flowers

per head and its obovate to elliptic bracts with

rounded, apiculate tips. With these characteristics

it can be distinguished from D. unguiculata, which

it most closely resembles.

Dicliptera skutchii Leonard, Publ. Field Mus. Nat.

Hist., Bot. Ser. 18: 1213. 1938. Figure 5.

Shrub to 3 m tall; erect, branching, internodes between

leaf-bearing nodes 5-7 cm long, 3-5 mm thick, puber-

ulous along 6 lines. Leaves with petioles 1-6.5 cm long,

sparingly puberulous; laminae ovate-elliptic, 9-17 cm

long, 3.5-7 cm broad, apically acuminate, basally acute,

slightly tapering, margins entire, cystoliths relatively ob-

scure above, more visible below, glabrous above and

below. Inflorescences in axillary cymes on peduncles to

8 cm long, peduncles with appressed puberulence; ped-

icels short, 1-2 mm long, hirtellous; bracts subtending

the cymes leaflike, to ca. 8 mm long; bracts subtending

the flowers lanceolate, glandular-puberulous, unequal,

the longer 10 mm long, 3 mm broad, the shorter 7 mm

long, 2.8 mm broad; 2 pairs of bracteoles subtending

each flower, the outer pair lanceolate, 6.5 mm long, 1.5

mm broad, glandular-puberulous toward apex, the inner

pair lanceolate-elliptic, 3.5 mm long, 1 mm broad, gla-

brous. Flowers 3 per cyme, to 6 cm long; calyx to ca. 7

mm long, coalesced for ca. [h its length, the lobes subu-

late, 3.5 mm long, 1 mm broad at the base, margins

glandular-ciliolate; corolla purple, glandular-puberulous,

the tube ca. 5 cm long, 3 mm broad at base, expanding

to 10 mm broad above the middle and narrowing to 7

mm broad at the mouth, the lips to 10 mm long, 1 acute

and finely notched at the tip, the other minutely 3-lobed,

the lobes suborbicular, ca. 0.75 mm in diameter; stamens

exserted to nearly the end of the lips. Fruits not seen.

Found in cloud forests at elevations around 1000

m in the vicinity of El General. Only one collec-

tion, Skutch 4140 (MO), in addition to the type

specimen, Skutch 3090 (us), has been observed.

Flower collection has been made in February.

Dicliptera skutchii can be recognized by its pe-

duncled cymes with short pedicels and its glan-

dular-puberulous bracts, bracteoles, and corollas.

By these characteristics it can be distinguished from

D. iopus, which it most closely resembles.

Dicliptera trifurca Oerst., Vidensk. Meddel. Dansk

Naturhist. Foren. Kj0benhavn 1854: 172. 1855.

Dicliptera trifurcata (Oerst.) Hemsl., Biol. Cen-

tralbl. 2: 525. 1881. Diapedium trifurcatum

(Oerst.) Kuntze, Revis. gen. pi. 2: 485. 1891.

Figure 5.

Herbs to subshrubs, to 2 m tall; stems branching, in-

ternodes between leaf-bearing nodes 1.5-8 cm long, 1-

3 mm thick, glabrous. Leaves with petioles to 5.3 cm

long, glabrous to sparingly puberulous along 2 lines; lam-

inae ovate-elliptic, 2.5-15 (18.5) cm long, 1-4 (5.6) cm

broad, apically acuminate, basally attenuate, margins

undulate, glabrous above, glabrous to sparingly puber-

ulous along costa and veins below, cystoliths more prom-

inent below. Inflorescences in axillary cymes of 3 flowers

each; peduncles to 8 cm long, mostly glabrous; bracts

subtending the cymes leaflike, frequently exceeding the

calyx; floral bracts oblong to oblong-spatulate, of un-

equal lengths, the larger to 1 1 mm long, 2 mm broad,

the smaller to 7 mm long, 1.5 mm broad, both apically

BURGER: FLORA COSTARICENSIS

39

rounded and apiculate, glabrous; bracteoles oblong-lan-

ceolate, to 5 mm long, 1 mm broad, sparingly puberu-

lous. Flowers with pedicels to 3 mm long, glabrous to

sparingly puberulous; calyx 6 mm long, segments subu-

late, 4 mm long, 0.6 mm broad, puberulous; corolla red-

violet or purplish, puberulous, 3.5-5.5 mm long, the tube

to 4 cm long, 2 mm broad basally, expanding to 7 mm

broad at the mouth, the lips oblong, to ca. 1.5 cm long,

the upper lip 7 mm broad, apically acute, lower lip 3-

lobed, the lobes subequal, ca. 0.75 mm long; stamens

exserted to the tip of the upper lip, filaments sparingly

puberulous; anther cells parallel, ca. 3 mm long, basally

apiculate. Fruits stipitate, to 1 8 mm long, 4 mm broad,

2 mm thick, puberulous; seeds 4, discoid, ca. 4 mm in

diameter, papillose.

Found in higher elevation rain forests or cloud

forests of the Cordilleras Central and Talamanca

at elevations from 1300 to 2300 m. Flowering

collections have been made from December

through June; also found in adjacent Panama.

Dicliptera trifurca is recognized by its pedun-

cled, three-flowered cymes with glabrous pedicels,

rounded and apiculate floral bracts, puberulous

corollas more than 3 cm long, and puberulous cap-

sules. These features distinguish it from D. iopus

which it closely resembles.

Dicliptera unguiculata Nees in Benth., Bot. voy.

Sulphur 149. 1844. Figure 5.

Herbs to 1 .5 m tall; steins erect or ascending, branched,

inter nodes between leaf- bearing nodes 5.5-10 cm long,

1-2.5 mm thick, increasingly pilose toward tips, terete

and glabrous below. Leaves soon deciduous, petioles to

9 cm long, pilosulous to subglabrous; laminae ovate to

elliptic, 5-9 cm long, 3-5.5 cm broad, apically acuminate

and often apiculate, basally acute to attenuate, margins

entire to undulate, both surfaces of younger leaves pilose,

older leaves subglabrous. Inflorescences composed of ax-

illary umbels or in crowded terminal and axillary spike-

like racemes to 9 cm long, 1.75 cm broad, bracts spat-

ulate, unequal, the larger 10-1 2 mm long, 5.5 mm broad,

the smaller 8-9 mm long, 3 mm broad, both apically

awned, the awn ca. 2 mm long, both basally clawed,

pilosulous, margins densely ciliate, 3-nerved; bracteoles

linear-lanceolate, 3.5 mm long, 0.5 mm broad, ciliolate.

Flowers sessile, to 15 mm long; calyx 2.5 mm long,

coalesced a little more than Vi its length, lobes 1 mm

long, minutely puberulous; corolla pink or lavender,

puberulous, the tube 7 mm long, 1 mm broad at the

base, 1.25 mm broad at the mouth, the lobes obovate,

8 mm long, 4 mm broad, 1 is apically obtuse, the other

apically minutely 3-lobed. Fruits elliptic, 4 mm long, 2

mm broad, 0.75 mm thick, puberulous; seeds discoid,

ca. 1.5 mm in diameter, puberulous.

This is the most common species of Dicliptera

in Costa Rica. It is found in moist thickets, open

fields, and along roadsides at elevations of 900 to

1500 m. Flowering collections have been made

between October and April. Its range extends north

to Mexico, but it has not yet been collected in

Panama.

Dicliptera unguiculata is recognized by its inflo-

rescences of axillary umbels and spikelike ra-

cemes, with conspicuous obovate, awned bracts,

short (to 1 5 mm long), pink or lavender corollas,

and often leafless condition. By these character-

istics it can be distinguished from D. pallida, a

much less commonly occurring species, which it

most closely resembles. It also resembles D. im-

bricata, which is much more common than D.

pallida, but more easily distinguished by the above

characteristics.

Dyschoriste Nees

REFERENCE— C. E. Kobuski, Ann. Missouri Bot.

Card. 15: 9. 1928.

Herbs to suffrutescent plants, stems prostrate, ascend-

ing or erect, glabrous or pubescent. Leaves sessile or

petioled; laminae mostly entire. Inflorescences cymose,

capitate, or spicate, terminal or axillary. Flowers sub-

tended by foliaceous bracts and bracteoles; calyx deeply

5-parted, usually subulate-setaceous, ciliate; corolla

5-lobed, obscurely or distinctly bilabiate, tube erect, limb

spreading; stamens 4, didynamous, anthers 2-celled, cells

oblong, minutely calcarate to mucronate at the base.

Fruit a capsule which is included in the persistent calyx,

oblong-linear, glabrous, 2-4 seeded, separating with dif-

ficulty at maturity into 2 valves; seeds flattened, subor-

bicular, with mucilaginous hairs when wet.

Dyschoriste is a widely distributed genus of about

100 species in both tropical and subtropical re-

gions of the world. A few species are found in

warm, temperate areas. Several species occur in

Central America.

Dyschoriste valeriana Leonard, Publ. Field Mus.

Nat. Hist., Bot. Ser. 18: 1215. 1938. Figure 5.

Herbs, prostrate, erect or ascending, to 60 cm tall,

frequently rooting at nodes of lower stem, stems qua-

drangular; internodes between leaf-bearing nodes 1.5-

4.5 cm long, 0.75-1 .5 mm thick, puberulous to glabrous.

Leaves petiolate to sessile, petioles to 1 cm long, glabrous

to puberulous along 2 lines; laminae ovate, 2.5-5 (6) cm

long, 1-2 (3) cm broad, apically acute to rounded or

obtuse, basally attenuate, margins entire to undulate,

cystoliths numerous, prominent above, obscure below,

40

FIELDIANA: BOTANY

glabrous to subglabrous on both surfaces. Inflorescences

of sessile axillary cymes, cymes of 2-5 flowers; 2 bracts

subtending each cyme, bracts oblanceolate, 10-18 mm

long, 2.5-6 mm broad, glabrous, margins minutely cil-

iolate; floral bracts linear to linear-elliptic, to 10 mm

long, 1 mm broad, bracteoles similar, but slightly small-

er, margins of both minutely ciliolate. Flowers sessile,

to 8-9 mm long; calyx 6-8 mm long at anthesis, 8-1 1

mm long at fructescence, coalesced for '/j-'/z of its length

by thin, easily torn tissue, lobes subulate, ca. 5.5 mm

long, 1 mm broad at base, single-nerved, sparingly pilose

along rib, margins ciliolate; corolla blue or purplish, pu-

berulous, the tube 6 mm long, 2 mm broad at base, 2.5

mm broad at the mouth, sub-bilabiate, lobes oblong,

apically rounded, upper lip with 3 lobes to 3 mm long,

2 mm broad, lower lip with 2 lobes to 2 mm long, 1.5

mm broad; stamens included, longer filaments to 2 mm

long, shorter filaments to 1 mm long, all glabrous, an-

thers 1 mm long, minutely calcarate at the base of each

cell, both cells joining to form an apiculate tip; ovary

glabrous. Fruit oblong-lanceolate, 9 mm long, 2 mm

broad, 1.75 mm thick, glabrous; seeds 4, ovate, flattened,

2.5 mm long, 2 mm broad, densely puberulous when

wet.

Found in disturbed habitats along trails and

roadsides in deciduous forest areas of Guanacaste

from near sea level to about 650 m elevation.

Flowering collections made from December to

April. One collection has been made from a similar

habitat in Panama. Since many collections of this

species have been made near Nicaragua, I expect

that D. valeriana occurs there also.

Dyschoriste valeriana is recognized by its square

stem, small blue or purplish flowers, four stamens

with two-celled anthers that have minutely cal-

carate bases, and two- to four-seeded capsules. It

closely resembles Hygrophila costata, which lacks

calcarate anthers and has 1 2 to 16 seeds per cap-

sule.

Elytraria Michx.

Herbs either caulescent or acaulescent. Leaves alter-

nate or subopposite, basal or restricted to the terminal

portion of the stem in a dense pattern, cystoliths lacking.

Inflorescences compact, peduncles with imbricate, co-

riaceous bracts. Flowers sessile with persistent perianth

except the soon deciduous corolla; bracts single; brac-

teoles 2; calyx 4-merous, segments narrow, entire or

toothed apically; corollas 5-merous, bilabiate, white or

blue, tube narrow, upper lip 1-lobed, lower lip 3-lobed;

stamens 2, barely exserted, anthers 2-celled which are

equal and parallel; ovary 2-celled; to 10 ovules in each

cavity, style 1 . Fruits a capsule, basally constricted and

apically acute; seeds borne on papilliform funicles.

About 1 7 species have been described for this

genus from southern North America plus New and

Old World Tropics.

Elytraria imbricate (Vahl) Pers., Syn. pi. 1: 23.

1805. Justicia imbricata Vahl, Eclogae ameri-

canae 1: 1. 1796. Tubiflora squamosa (Jacq.)

Ktze., Revis. gen. pi. 2: 500. 1891. is no longer

a valid synonym. The genus Tubiflora was re-

jected by the International Code of Botanical

Nomenclature 1972. Figure 5.

A weedy caulescent herb, occasionally acaulescent, to

50 cm tall; internodes between leaf- bearing nodes 0-15

mm long, 2-3 mm thick, glabrous to sparingly puberu-

lous or pilose. Leaves alternate, sessile to petiolate, pet-

ioles to 2 cm long, pilose; laminae ovate-lanceolate, 3.5-

13 cm long, 2—4.5 cm broad, apically acute, basally at-

tenuate, narrowing to a winged petiole, margins undu-

late, sparingly pilose above and below, more dense on

costa and lateral veins. Numerous scapes borne at leaf

axils, 5-28 cm long, branched or simple, covered with

lightly appressed, sheathing, ovate, or subulate bracts.

Inflorescences in terminal spikes, 1-several, 1.5-4 cm

long, 4 mm broad; bracts oblong to elliptic, 4-6 mm

long, 1.5-2 mm broad, awn-tipped and stiff with upper

bracts bearing 4 hyaline wings, 2 rounded extending from

base and 2 apical extending parallel to the apex; 2 brac-

teoles per flower, subulate, to 3 mm long, costa ciliate.

Flowers sessile, 5-8.5 mm long; calyx segments to 4.5

mm long, posterior segment elliptic, bidentate, 1 .25 mm

broad, lateral segments subulate, 0.75 mm broad, an-

terior segment elliptic, bidentate, 0.75 mm broad, all

segments transparent, puberulous; corolla pale lavender

or blue, glabrous, tube cylindrical, 4-7 mm long, 0.5 mm

broad, lower lip spreading, to 4 mm long, 3-lobed, the

middle lobe 2 mm broad, 2-lobed at apex, the lobes

rounded, ca. 1 mm long, the lateral lobes similar, but

slightly smaller, the upper lip erect, to ca. 2 mm long, 1

mm broad, apically 2-lobed. Fruits elliptic, ca. 3 mm

long, 1 mm broad, glabrous; mature seeds light brown,

rounded, triangular, ca. 0.5 mm long, glabrous.

Found in dry pastures and along roadsides and

trails of the Pacific watershed area at elevations

of 50 to 1 1 70 m. Flowering collections have been

made from November through March. This species

ranges from the southwestern United States to

western South America. Leonard (1951) reports

that in Central America this plant is used as a

remedy for dysentery, and the crushed leaves are

said to cure pimples.

Elytraria imbricata is recognized by its numer-

ous spikes borne on axillary, bracted culms, its

small bluish bilabiate corollas with awned, winged

bracts, and its capsules lacking hooklike funicles.

Superficially, this plant can be confused with Aphe-

landra tonduzii, which differs in all of the above

characteristics.

Graptophyllum Nees

Shrubs with branching, ascending stems. Leaves ses-

sile to petiolate, laminae often colored and variegated,

BURGER: FLORA COSTARICENSIS

41

margins entire or dentate. Flowers pedicellate, in short,

nearly sessile cymes, terminal panicles or solitary in the

axils of the leaves; calyx 5-merous; corollas red or purple,

bilabiate, the tube inflated above, with 2, short, recurved

lobes, the lower lip 3-lobed; stamens 2; staminodes 2.

Fruits of stalked, clavate capsules; seeds 2-4.

Ten species have been described for this genus,

occurring in West Africa, New Guinea, Australia,

and Polynesia. None is native to the New World.

Graptophyllum pictum (L.) Griff., Not. pi. asiat.,

part 4: 139. 1854. Based on Justicia picta L.,

Sp. pi. 1: 21. 1763. G. hortense (L.) Nees in

Wall., PI. asiat. rar. 3: 102. 1832. Figure 6.

Shrubs, stems to 3 m tall; ink-modes between leaf-

bearing nodes 4-7 cm long, 3-4 mm thick, glabrous.

Leaves sessile to short-petiolate, petioles to 4 mm long,

glabrous; laminae ovate to elliptic, 4.5-1 1 (20) cm long,

2-7 (9) cm broad, apically acuminate, basally acute to

obtuse, slightly tapering, purplish or green, variously

marked with yellow, margins entire, cystoliths dense and

generally prominent above, obscure below, glabrous on

both surfaces. Inflorescences in terminal racemes, the

rachis glabrous or sparingly puberulous. Flowers with

glabrous pedicels to 8 mm long, each subtended by 1

subulate bract, bract 3 mm long, 2 mm broad at base,

minutely ciliolate, bracteoles 2, similar to bracts al-

though slightly smaller; calyx segments subulate, ca. 3

mm long, 1 .25 mm broad at base, glabrous; corollas dark

purple or crimson, glabrous outside, glandular-puberu-

lous inside, 4.5-8 cm long, tube funnelform, ca. 3 cm

long, 4 mm broad basally, 1 1 mm broad at the mouth,

upper lip erect, oblong, ca. 1 6 mm long, 1 1 mm broad,

apically 2-lobed, lobes rounded, 5 mm long, 4 mm broad,

lower lip 3-lobed, lobes oblong, 1 5 mm long, narrowing

to a rounded tip, middle lobe 7 mm broad, lateral lobes

6 mm broad; stamens exserted to just below the tip of

the upper lip, filaments glabrous, anthers 4 mm long, 1

mm broad, cells slightly unequal at the base, acute api-

cally and basally, staminodes to 3 mm long. Fruits not

observed.

This is a species that is cultivated as an orna-

mental in the gardens of tropical America. Ac-

cording to Bailey ( 1 949) it is probably native to

New Guinea.

Graptophyllum pictum is recognized by its mot-

tled leaves, relatively large (to 8 cm long), bilabiate

corollas, two stamens with two-celled anthers and

two staminodes. It resembles various species of

Odontonema, but can be most easily distinguished

from them by its mottled leaves and larger corol-

las.

Habracanthus Nees

Herbs or shrubs. Leaves usually with ovate to elliptic

laminae. Flowers borne in loose to dense panicles; calyx

5-merous, segments narrowly lanceolate; corolla blue,

red, or yellow, salverform, bilabiate, upper lip narrow,

cuneate, lower lip usually oblong-cuneate, apically

3-lobed, lobes rounded; stamens 2, exserted, divergent,

anthers 1 -celled, filaments attached below middle of an-

ther; style exserted. Fruit a clavate capsule; seeds 4.

This is a genus of about 40 species found in

Mexico, Central America, and northern South

America.

Habracanthus silvaticus Nees in DC., Prodr. 11:

312. 1847. Figure 6.

Shrub to 2 m tall, younger stems quadrangular; inter-

nodes between leaf-bearing nodes 3-8 cm long, 1-2 mm

thick, glabrous to strigose along 2 lines. Leaves with

petioles 5-35 (75) mm long, glabrous to sparingly puber-

ulous; laminae elliptic to ovate-elliptic, 6-13 cm long,

2.5-5 cm broad, apically acuminate, basally acute, gla-

brous to sparingly puberulous along costa and lateral

veins, margins undulate, cystoliths prominent on both

surfaces. Inflorescences of loose, terminal thyrses to 10

cm long, 5 cm broad, lateral branches of thyrse dichot-

omously branched, branches of inflorescence minutely

strigose; bracts subulate, to 6 mm long, 1 mm broad at

base, glabrous. Flowers with pedicels ca. 2 mm long;

calyx whitish to purplish, segments subulate, 3-4 mm

long, 0.5 mm broad at base, glabrous; corolla blue, bi-

labiate, 12-18 mm long, glabrous, tube ca. 8 mm long,

near cylindrical, ca. 1 mm broad, the lips ca. 10 mm

long, the upper lip oblong, 2 mm broad, apically round-

ed, lower lip elliptic, ca. 4 mm broad, apically 3-lobed,

the lobes 0.5 mm long, middle lobe 1 mm broad, lateral

lobes 0.5 mm broad; stamens exserted to ca. the tip of

the upper lip. Fruit clavate, to 1 5 mm long, 3 mm broad,

2 mm thick, glabrous; seeds flattened, ovate, 2 mm long,

1.5 mm broad, puberulous.

A relatively common plant in wet forests and

adjacent clearings of higher elevations on Pacific

and Caribbean slopes of the Cordillera Central

from 975 to 2200 m elevation. Flowering collec-

tions have been made from July through March.

This species was first described in Chiapas, Mex-

ico, and extends from there to Panama.

Habracanthus silvaticus is recognized by its

bluish calyx and bilabiate corolla, with two sta-

mens that have one-celled anthers; also by its in-

florescence in a loose, terminal thyrse. It resembles

Razisea spicata, but can be distinguished from it

in flower color and inflorescence.

42

FIELDIANA: BOTANY

Hansteinia Oersted

Herbs or shrubs. Leaves opposite, usually petiolate;

laminae elliptic or ovate. Inflorescences in loose or com-

pact panicles which are sometimes thyrsoid; flowers with

5-merous calyx, calyx segments narrow, equal; corollas

red, orange, yellow, or white, cylindrical, narrow basally,

abruptly expanded just above the calyx, somewhat

obliquely enlarged and oblong, often constricted again

at the mouth; upper lip erect, the lower lip barely distinct

from the corolla tube, 3-lobed; stamens 2, attached at

base of corolla tube, exserted, anthers 1 -celled, basally

obtuse. Fruits a capsule, clavate, glabrous, 4-seeded.

Fourteen species have been described for this

genus, ranging from Mexico to Bolivia. In Costa

Rica the species are all found at elevations above

1000 m. Generally these species are not often col-

lected in Costa Rica, which may indicate infre-

quent occurrence. This genus closely resembles the

genera Habracanthus and Razisea. See the genus

description of Razisea for a discussion of these

differences.

Key to Species of Hansteinia

1 a. Leaf bases rounded or subcordate 2a

Ib. Leaf bases attenuate (except leaves subtending the inflorescence) 3a

2a. Corolla more than 20 mm long, glandular-puberulous; petioles mostly more than 2 cm long

H. ventricosa

2b. Corolla less than 20 mm long, glabrous; petioles less than 5 mm long H. sessilifolia

3a. Capsules less than 12 mm long; leafy internodes 7-8.5 cm long H. gracilis

3b. Capsules more than 1 2 mm long; leafy internodes 6 cm or less long 4a

4a. Inflorescences in fairly open racemose panicles, branches relatively few, erect or ascending; bracts

2-5 mm long H. stricta

4b. Inflorescences in compact cymose panicles, branches numerous, spreading; bracts 5-10 mm long

H. blepharorhachis

Hansteinia blepharorhachis (Lindau) Durkee,

comb. nov. Kolobochilus blepharorhachis Lin-

dau, Anales Inst. Fis.-Geogr. Nac. Costa Rica

9: 189. 1898, and in Pitt., Prim. fl. costaric. 2:

309. 1900. Glockeria blepharorhachis (Lindau)

Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser.

18: 1218. 1938. Razisea breviflora D. Gibson,

Fieldiana, Bot. 34: 77. 1972. Figure 6.

Herbs to suffrutescent plants to 2 m tall, stems

subquadrangular to quadrangular in younger portions;

internodes between leaf-bearing nodes 1.5-8 cm long,

1.5-2.5 mm thick, sparingly pilose mostly along 2 lines.

Leaves with petioles 2-6 cm long, sparingly pilose; lam-

inae elliptic-ovate, 7-18 cm long, 2-6.5 cm broad, api-

cally acuminate, basally attenuate, margins undulate to

crenulate, older leaves mostly glabrous, drying to very

dark green or nearly black above, younger leaves with

pilose hairs restricted to costa and veins, cystoliths abun-

dant though minute and obscure, more visible beneath.

Inflorescences in long, narrow, terminal thyrsoid pani-

cles to 30 cm long, 3-5 cm broad; peduncle to 7 cm long,

puberulous, the rachis pilose to subtomentose with brown

to reddish brown hairs plus longer, scattered, glandular

hairs, pedicels to 8 mm long, pubescence that of the

rachis; bracts narrowly subulate, 5-10 mm long, 1.5-3

mm broad, bracteoles similar, but smaller, both puber-

ulous plus a few longer glandular hairs. Flowers with 5-

merous calyx, segments linear, 4.5-10 mm long, 0.75-

1 mm broad, minutely puberulous, plus much longer

glandular hairs; corolla orange or orange-red, 1 5-22 mm

long, the tube to ca. 14 mm long, 4 mm broad at base,

expanding abruptly to 5-6 mm broad just above the

calyx, glabrous outside, pilosulous glandular hairs inside,

upper lip erect, oblong, 4-8 mm long, 3-4 mm broad,

apically rounded, lower lip short, truncate, 3-lobed, lobes

each ca. 1 mm long, suborbicular; stamens exserted to

well above the upper lip. Fruits clavate, 13-15 mm long,

3.5-4 mm broad, 2.5-3 mm thick, glabrous; seeds flat-

tened, ovate, 2.5-3.5 mm long, 2-2.5 mm broad, surface

muricate.

Plants of cloud forests at elevations of 1 500 to

2000 m. The locality of the type specimen ( War-

scewicz s.n.) is specified only as Costa Rica and

Veraguas (Panama). Flowering collections have

been made in April, August, and October from the

Cordilleras Tilaran, Central, and Talamanca. This

species occurs rather frequently in light gaps. It

occurs in habitats similar to those of Razisea spi-

cata and is also hummingbird-pollinated.

Hansteinia blepharorhachis is recognized by its

compact and comparatively narrow, but much-

branched, cymose panicles (thyrses) combined with

leaf blades which are elliptic-ovate with attenuate

bases. It is most similar to H. stricta, which differs

BURGER: FLORA COSTARICENSIS

43

in its generally smaller, less compact, less branched,

broader racemose panicles and smaller bracts (2.5

mm long vs. 5 to 10 mm).

Hansteinia gracilis Oerst., Vidensk. Meddel.

Dansk Naturhist. Foren. Kjobenhavn 1854: 143.

1855.

Herb to suffrutescent plant to ca. 60 cm tall; steins

subtetragonal, erect, simple or sparingly branched, in-

ter nodes between leaf-bearing nodes 7-8.5 cm long, pu-

berulous. Leaves with petioles to ca. 2.5 cm long; laminae

elliptic-ovate, to 1 8 cm long, 7 cm broad, apically acu-

minate, basally attenuate, margins subrepand-crenate,

glabrous on both surfaces except minutely strigose on

costa and veins. Inflorescences in terminal panicles to

1 5 cm long, rachis glandular-pubescent, hairs rust-col-

ored. Flowers with 5-merous calyx, calyx segments red,

7-9 mm long at anthesis, 13.5-16 mm long at fruiting

stage, glandular-villous; corolla red, 13.5-16 mm long,

tomentose. Fruit 9-1 1 mm long, puberulous.

Found in dense, damp woods on Mt. Jaris

northeast of Santiago (Santiago de Puriscal) in San

Jose Province, elevation 500 to 1000 m. I have

seen no collections of this species. The above de-

scription was adapted from the original, which was

based on a collection in both flower and fruit col-

lected in November.

Hansteinia gracilis differs from other Costa Ri-

can species in the genus by its shorter capsules (9

to 1 1 mm long vs. 1 2 mm or more) and its longer

internode lengths (7 to 8.5 cm long vs. 6 cm or

less).

Hansteinia sessilifolia (Oerst.) Durkee, comb. nov.

Glockeria sessilifolia Oerst., Vidensk. Meddel.

Dansk Naturhist. Foren. Kjobenhavn 1854: 141.

1855. Figure 6.

Herbs to suffrutescent plants to 2 m tall, older stems

terete, younger stems subquadrangular, glabrous to pu-

berulous along 2 lines; internodes between leaf-bearing

nodes 4.5-5.5 cm long, 1.5-2.5 mm thick. Leaves sessile

(in those subtending the inflorescence) to petiolate, pet-

ioles to 4.5 cm long, puberulous; laminae ovate, 5.5-1 1

cm long, 2-6 cm broad, apically acuminate, basally

rounded or subcordate in leaves at base of inflorescence

to acute in lower leaves, margins entire to undulate, gla-

brous or sparingly pilose on both surfaces, hairs mostly

restricted to costa and veins, cystoliths abundant on both

surfaces though minute and obscure. Inflorescences in

loose, terminal, dichotomously branching, thyrsoid pan-

icles to 38 cm long, 15 cm broad, sometimes with ad-

ditional axillary panicles, rachises and branches puber-

ulous to sparingly so, pedicels to 2.5 mm long, sparingly

puberulous; bracts narrowly triangular, to 3 mm long,

0.5 mm broad, glabrous to very sparingly puberulous.

Flowers with 5-merous calyx, calyx segments linear-su-

bulate, to 4-6 mm long, 0.3-0.5 mm broad, sparingly

glandular-puberulous; corolla red and yellow, 1 2-22 mm

long, glabrous, the narrow basal portion 2.5-3 mm long,

1-2 mm in diameter, the upper portion expanded to 3-

5 mm diameter, then gradually narrowing to 2.5-4 mm

diameter at the mouth, the upper lip spatulate, apically

acute, 3.5 mm long, 1.5 mm broad at the base, 0.5 mm

broad at the lip, lower lip short, recurved, ca. 1 mm long,

3 mm broad, lobes apically rounded; stamens exserted

to 5 mm beyond upper lip, anthers 2-2.8 mm long;

capsules clavate, 12-14 mm long, 2.3-3 mm broad, 1.8-

2.5 mm thick, glabrous; seeds flattened, elliptic 2-2.2

mm long, 1.2-1.4 mm broad, hispidulous.

A plant of the wooded, southern slopes of Vol-

can Barva (Barba), Volcan Poas, and the Monte-

verde cloud forest area in Puntarenas Province. I

have seen only two collections of this species, Leon

79 (F) and Dryer 1671 (F). Flowering collections

have been made in September and October.

Hansteinia sessilifolia is recognized by its loose,

dichotomously branching inflorescences and its

short corollas (12 to 22 mm long) with stamens

slightly exserted. It most closely resembles H. ven-

tricosa, but can be distinguished from it by the

above characteristics.

Hansteinia stricta (Leonard) D. Gibson, Field-

iana, Bot. 34: 63. 1972. Glockeria stricta Leon-

ard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18:

1219. 1938. Figure 6.

Herb to subshrub to 1.5 m tall, older stems terete,

glabrous, younger stems quadrangular, sparingly puber-

ulous or scattered brown puberulence along 2 lines; in-

ternodes between leaf-bearing nodes 3-6 cm long, 1.5-

2 mm thick. Leaves with petioles 1.5-3.3 cm long, gla-

brous to puberulous; laminae ovate, 6-12 cm long, 2.5-

5 cm broad, apically acuminate, basally attenuate, mar-

gins entire to undulate, sparingly and minutely strigose

on both surfaces, mostly restricted to costa and veins,

minute cystoliths abundant and apparent on both sur-

faces. Inflorescences in terminal and axillary racemose

panicles to 27 cm long, 8 cm broad; rachis and branches

minutely puberulous plus much longer (to 1 mm), scat-

tered glandular hairs, pedicels subsessile to 8 mm long,

pubescence that of the rachis; bracts and bracteoles subu-

late, 2-5 mm long, 1-2 mm broad, minutely puberulous

plus a few longer glandular hairs. Flowers with 5-merous

calyx, calyx segments narrowly linear or subulate, 4.5-

8 mm long, 0.75-1 mm broad, minutely puberulous plus

scattered glandular hairs; corolla orange-red and yellow,

15-19 mm long, 1 .5 mm broad at base expanding abruptly

to 6 mm broad above the calyx and narrowing to 5 mm

broad at the mouth, sparingly puberulous, upper lip erect,

4 mm long, 3 mm broad, apically acute, lower lip 3-lobed,

lobes rounded, 0.75 mm long, 1 mm broad; stamens

exserted well beyond the tip of the upper lip. Fruits

44

FIELDIANA: BOTANY

clavate, 13-15 mm long, 3.5 mm broad, 2.5 mm thick,

glabrous; seeds ovate, flattened, 2.5 mm long, 2 mm

broad, muricate.

Found in moist forests on the slopes of the Cor-

dilleras Central and Talamanca at elevations of

1500 to 1900 m. Flowering collections have been

made in November, December, and February.

Hansteinia stricta is recognized by its ovate

leaves with attenuate bases and its open paniculate

inflorescences with relatively few branches and

these strict, erect, or ascending. It most closely

resembles H. blepharorhachis, which has cymose

panicles (thyrses), with narrower, more compact

branching and larger bracts (5 to 10 mm long vs.

2 to 5 mm).

lections have been made in October, December,

January, and February.

There is variation in the degree of hairiness

among specimens from over the range of Han-

steinia ventricosa. Early flowering specimens in

particular may be confused with Razisea spicata,

but can be distinguished from it with their shorter

corollas (2.6 to 3 cm vs. 4 to 6.2 cm long), which

expand more abruptly, and by the more profuse

branching of the inflorescences. Hansteinia ven-

tricosa resembles H. stricta and H. blepharo-

rhachis somewhat, but can be distinguished from

them by its broader laminae, with rounded, sub-

cordate, or obtuse bases, as opposed to their nar-

rowed laminae with attenuate bases.

Hansteinia ventricosa (Donnell Smith) D. Gibson,

Fieldiana, Bot. 34: 63. 1972. Glockeria ventri-

cosa Donnell Smith, Bot. Gaz. (Crawfordsville)

27:439. 1899. Figure 6.

Herbs to subshrubs to 3 m tall, older stems terete,

younger stems subquadrangular ; internodes between leaf-

bearing nodes 2-7 (1 1) cm long, 1.5-3 mm thick, sub-

tomentose, hairs ca. 1 mm long. Leaves with petioles to

5.8 cm long, pilose to subtomentose; laminae ovate to

ovate-elliptic, 7-15 cm long, 3-12 cm broad, apically

acuminate, hasally subcordate to rounded or obtuse,

margins crenulate to undulate, both surfaces pilose, cys-

toliths minute. Inflorescences in terminal and axillary

thyrses to 29 cm long, 1 1 cm broad excluding corollas,

rachises and branches pilose plus glandular hairs to pu-

berulous, pedicels to 9 mm long, puberulous-glandular;

bracts and bracteoles oblong to narrowly lanceolate, to

9 mm long, 2 mm broad, apically acute to acuminate,

puberulous to pilose plus occasional glandular hairs.

Flowers with 5-merous calyx, segments linear to 7 mm

long at flowering and to 1 2 mm long at fruiting stage, 1

mm broad, puberulous plus occasional glandular hairs;

corolla orange, 2.6-3 cm long, tube curved outward,

sparingly glandular-pilose, 2 mm broad basally, con-

tracting just above the ovary and then expanding abrupt-

ly to 5 mm broad just above the calyx lobes, the upper

lip erect, elliptic, to 5 mm long, 2 mm broad, apically

bidentate, lower lip ca. 1.5 mm long, 3-lobed, the lobes

ca. 1 mm long, rounded; stamens exserted to ca. 1.5 cm

beyond the tip of the upper lip. Fruits clavate, 11-19

mm long, 3-4 mm broad, 3—4 mm thick, glabrous; seeds

4, oblong, flattened, basally oblique, 2 mm long, 1 .5 mm

broad, coarsely roughened.

This is a comparatively wide-ranging species

found in shade or partial shade of moist, evergreen

forests of the Cordilleras Tilaran, Central, and Ta-

lamanca and of adjacent areas in Panama at ele-

vations between 1000 and 2500 m. Flowering col-

Henrya Nees

REFERENCE— G. B. Happ, Monograph ofTetra-

merium and Henrya. Ann. Missouri Bot. Gard.

24: 501-582. 1937.

Herbaceous to suffrutescent perennials; stems erect,

spreading or decumbent, younger stems often pubescent

with glandular hairs, tetrangular to subterete, with pale,

often exfoliating, epidermis. Leaves petiolate, often de-

ciduous; laminae rotund-ovate to linear, entire, pubes-

cent to glabrous on both surfaces, cystoliths present. In-

florescences in terminal and axillary spikes with loosely

or densely imbricated bracts; cauline bracts solitary; in-

volucral bracts coalescent from base to near the apex on

the posterior side and closely enclosing the fruit, green,

mucronate at or near the apex. Flowers sessile or short-

pedicellate; calyx 5-merous; corolla narrowly funnel-

form, bilabiate, tube ca. '/3 the length of the corolla, an-

terior lip 3-lobed, the posterior lip shallowly bilobed or

entire; stamens 2, exserted; anthers 2-celled, nearly par-

allel, one cell longer than the other. Fruit a capsule, seeds

2, flattened, hispid to hispidulous on one side, glabrous

on the other.

Happ had recognized 20 species for this genus,

ranging from the northern states of Mexico and

adjacent United States to Costa Rica. He consid-

ered the position of the mucro on the involucral

bract to be a very important character in species

designations. This has been questioned by some,

and it appears that there are many fewer species

than he designated.

Henrya scorpioides Nees in DC., Prodr. 1 1: 491.

1847. Tetramerium scorpioides (Nees) Hems-

ley, Biol. Centralbl. 2: 526. 1882.

BURGER: FLORA COSTARICENSIS

45

Herbs to suffrutescent plants to 1 m tall; older steins

terete, internodes between leaf-bearing nodes 2-4 cm

long, 1-1.5 mm thick, puberulous. Leaves with petioles

1-1 5 mm long, puberulous; laminae ovate, 1-6 cm long,

0.5-3 cm broad, apically acute to acuminate, basally

obtuse to acute, margins entire, puberulous-glandular on

both surfaces, often glabrate with age, cystoliths prom-

inent on both surfaces. Inflorescences 2-10 cm long,

loosely imbricated cauline bracts linear-elliptic, elliptic

or linear-oblanceolate, 2-4 mm long, 1-2 mm broad;

involucral bracts oblanceolate, 7-9 mm long, glandular-

puberulous, the mucro just below the apex. Usually 2

flowers per bract, only 1 producing a fruit, calyx seg-

ments lanceolate, ca. 1.5 mm long; corolla pale yellow

to cream-colored, 10-15 mm long. Fruits clavate, 4.5-

6 mm long, 1.5-2.5 mm broad, ca. 1 mm thick, pu-

berulous; seeds 1 or 2, flattened, elliptic, ca. 1.5 mm

long, 1.25 mm broad, margins hispidulous.

This species ranges from Mexico to Costa Rica.

Habitats have been described only as "rocky soil"

or "rocky thorn forests." Only one collection from

Costa Rica, M. Quiros 671 (F), is known. It was

collected in flower at Desamparados de San Ma-

teo, Alajuela Province, in March at 100- to 500-m

elevation.

Henrya scorpioides is recognized by its spicate

inflorescences, with loosely imbricated bracts, its

coalesced involucral bracts that are conspicuously

glandular-puberulous and are 7 to 9 mm long, and

its cauline bracts that are conspicuously shorter

than the involucral bracts (2 to 4 mm long). This

may be confused with Tetrameriwn nervosum,

which differs in its more compact spikes with larg-

er bracts that are four-ranked and lack the co-

alesced involucral bract.

Herpetacanthus Nees

Herbs or shrubs. Inflorescences spicate; bracts loosely

imbricate. Flowers with a 5-merous calyx, calyx seg-

ments equal; corolla 5-merous, bilabiate, upper lip api-

cally bidentate, lower lip 3-lobed; stamens 4, didyna-

mous, inserted or barely exserted, the longer of each pair

2-celled with one of the cells superposed, the shorter of

each pair 1 -celled. Capsules clavate; seeds 4.

This is a genus of about 1 0 species, ranging from

Central America to Brazil. Most of the species

described are from Brazil. This genus is distin-

guished from other genera in the family with bi-

labiate corollas by its didynamous stamens, the

longer pair 2-celled, with one of the cells distinctly

superposed and the shorter pair of stamens one-

celled.

Herpetacanthus panamensis Leonard, J. Wash.

Acad. Sci. 32: 185. 1942. Standleyacanthus cos-

taricanus Leonard, Ceiba 3: 143. 1952. Figure 7.

Herbs or subshrubs to 1 m tall; stems ascending, spar-

ingly branched, rooting at lower nodes in some, younger

stems quadrangular, puberulous to pilose mostly along

2 lines, hairs with a purplish or reddish banding pattern,

older stems terete, glabrous; internodes between leaf-

bearing nodes 2.5-4 cm long, 0.75-1 .25 mm thick. Leaves

with short petioles to 1 cm long, pubescence that of the

stem; laminae ovate to ovate-elliptic, 4-14.5 cm long,

2.5-6 cm broad, apically acute to acuminate, basally

acute to attenuate, margins undulate, glabrous to strigose

on the costa and veins of both surfaces, cystoliths dense

and prominent on both surfaces. Inflorescences in lax

terminal and axillary spikes or spicate panicles, spikes

to 3.5 cm long, 2.5 cm broad; bracts loosely imbricate,

ovate to elliptic, 9-14 mm long, 4-7 mm broad, apically

acute to acuminate, glabrous or subglabrous, margins

ciliate; bracteoles linear to narrowly elliptic, 6-8 mm

long, 0.5-1 mm broad when fruits are mature, glabrous,

margins sparingly ciliate. Flowers with 5-merous calyx,

segments subulate, 3-4 mm long, 0.5 mm broad, mi-

nutely glandular-puberulous inside, glabrous outside,

tipped by 1 or 2 hairs; corolla white, to 1 7 mm long, the

tube 1 1 mm long, 2 mm broad at base, 3.5 mm broad

at the mouth, glabrous, the lips ca. 5.5-8 mm long, pu-

berulous toward the tip, upper lip ovate, 4 mm broad,

apically minutely bidentate, lower lip 3-lobed, lobes 2-

3 mm long, middle lobe rounded, 2.5 mm broad, lateral

lobes acute, ca. 1.5 mm broad, the stamens exserted or

included, filaments of the longer pair 3 mm long, of the

shorter pair 2 mm long, glabrous, anthers ca. 1 mm long,

0.3 mm broad. Fruits clavate, 9-10 mm long, stipe 3.5-

4 mm long, 0.5 mm thick, seed-bearing portion 3 mm

broad, 1.5 mm thick, glabrous; seeds 4, flattened, ovate,

2.5 mm long, 2.0 mm broad, roughened.

Collected in wet, shaded areas of forests, fre-

quently along streams from near sea level to ca.

800 m in Panama and Costa Rica. In Costa Rica

most collections have been made in Alajuela, Her-

edia, and Limon provinces of the Caribbean wa-

tershed area, but one, Allen 6297 (F, CR) is known

from Esquinas forest of Puntarenas Province.

Flowering collections in Costa Rica have been

made in March, July, September, and October.

The September collection, Allen 6297 (F), was from

an area which has a more extended dry season

than the others. One collection, Opler 119 (F), de-

scribes the bracts as purplish. This specimen has,

as have most others, dark purplish green bracts

when dried; some, however, such as Schnell 810

(F), collected inland from Puerto Viejo in Limon

Province, have yellowish green bracts. This vari-

ation may be regional, or it may be an age differ-

ence, since the Schnell collection was in an early

flowering stage with no fruits.

46

FIELDIANA: BOTANY

Herpetacanthus panamensis may be confused

with various of the larger-bracted species of Jus-

ticia, but can be easily distinguished from them,

with its second pair of stamens that are single-

celled (vs. two, two-celled stamens, with cells that

are frequently superposed). If stamens are missing,

the loosely imbricate bracts that are frequently

purplish are the best distinction.

Hygrophila R. Brown

Erect or diffuse herbs. Leaves opposite. Flowers in

axillary fascicles; bracts linear; calyx 5-merous, segments

subequal; corolla tubes subcylindric, small, the limb

2-lipped; stamens 4, didynamous, or 1 perfect stamen

and a staminode in each pair, the filaments in each pair,

united at the base; anthers 2-celled, some mucronulate;

posterior stigma lobe abortive. Fruit an oblong, non-

stipitate capsule; seeds 4-18 or more.

This genus is pantropical, with about 80 species.

Hygrophila has some resemblance to the genus

Dyschoriste, but can be distinguished from it in

Central America by its white flowers (vs. blue or

purple) and capsules with numerous seeds (14 to

48 vs. two to four).

Hygrophila costata Nees, PI. hort. bonn. icon. 2:

7. t. 3, 1824, and in DC, Prodr. 11: 88. 1847.

H. guianensis Nees, London J. Bot. 4: 634. 1 845.

H. conferta Nees in Mart., Fl. bras. 9: 21. 1847.

Figure 7.

Herbs to 1.5 m tall, sometimes declining and rooting

at lower nodes, steins obtusely quadrangular; internodes

between leaf-bearing nodes 2-7 (10) cm long, 1-3 mm

thick, glabrous. Leaves petiolate to subsessile, petioles

to 2.5 cm long, glabrous; laminae lanceolate to elliptic,

3.5-18 cm long, 0.5-5 cm broad, apically acuminate,

basally attenuate, margins entire to undulate, cystoliths

very numerous on the upper epidermis, less numerous

on the lower, glabrous above, sparingly puberulous be-

low, mostly restricted to costa and veins. Flowers sessile,

often in clusters of 3 or 4, bracts lanceolate-elliptic, 6-

9 mm long, 1.5-3 mm broad, sparingly puberulous, brac-

teoles similar, 2-3 mm long, ca. 0.5 mm broad, calyx

segments linear, equal, 5-9 mm long at anthesis, 6.5-12

mm long in fruit, sparingly puberulous; corolla white,

sometimes streaked or spotted with purple, 5.5-9 mm

long, tube ca. 1 .5 mm broad at base, expanding to 2 mm

broad at the throat, puberulous, 2-lipped, lips ca. 3 mm

long, upper lip 2-lobed, lower lip 3-lobed; stamens 4,

didynamous, filaments of each pair united at the base,

anthers 2-celled, ca. 1 mm long, cells basally mucron-

ulate; fruits narrowly oblong, sessile, glabrous, 8-1 3 mm

long, 6-8 seeds per locule.

This is a wide-ranging plant found from Mexico

to central Argentina in moist habitats. In Costa

Rica it has been collected along streams, rivers,

forested thickets, weedy roadsides, open pastures,

and rain forests from sea level of the Osa Peninsula

to mountain slopes of the Cordilleras Talamanca

and Central at 1 500 m.

Hygrophila guianensis and H. costata have been

distinguished by leaf shape and size, length of ca-

lyx, corolla, and capsule. I have brought H. gui-

anensis into synonymy because the numerous

measurements of all these characters in Costa Ri-

can collections intergrade and overlap. Further-

more, they show no habitat differences. This is a

common and wide-ranging complex, which on the

basis of these observations, appears to be one

species.

Hygrophila costata is recognized by its sessile

flowers often in clusters of three or four, its in-

conspicuous, white, bilabiate corollas with four

didynamous stamens with two-celled, basally mu-

cronulate anthers, and its oblong, sessile capsule

with 1 2 to 16 seeds.

Hypoestes Soland.

REFERENCE— W. T. Thiselton-Dyer, Flora of

Central Africa 5: 244. 1900.

Herbs or shrubs. Leaves petiolate. Inflorescences in

axillary and/or terminal spikes or panicles; floral leaves

resembling reduced vegetative leaves often present; bracts

2 per flower, free or coalesced for '/3-% of their length.

Flowers with 5-merous calyx, calyx lobes equal, subu-

late, free or coalesced 'A-% of their length; corolla bila-

biate, tube narrow, gradually expanding to the mouth,

upper lip 2-lobed, lower lip 3-lobed; stamens 2, anthers

1 -celled, basally rounded. Fruits clavate; seeds 2 or 4.

This is a genus of the Paleotropics, with 1 50

species extending from Africa to Australia. Most

species are found in Madagascar.

Hypoestes phyllostachya Baker, J. Linn. Soc., Bot.

22: 511. 1887. Figure 7.

Herbs to 70 cm tall, young basal shoots especially

velutinous; internodes between leaf-bearing nodes 2-6

cm long, 1.25-2.0 mm thick, subtetrangular, glandular

plus appressed hairs to appressed hairs at angles only.

Leaves petiolate, petioles to 2.5 cm long, appressed hairs

on upper surface only; laminae ovate, 2-4.5 cm long,

1.5-2.5 cm broad, apically acute, basally obtuse to

BURGER: FLORA COSTARICENSIS

47

rounded, margins entire, glabrous to sparingly puberu-

lous on the midrib of both surfaces, most surfaces dotted

with numerous pink spots to ca. 4 mm in size, cystoliths

present on both surfaces. Flowers borne singly and in

terminal and axillary spikes, often with a gradual tran-

sition from solitary flowers to spikes, rachises glandular-

puberulous; bracts 2 per flower, fused for up to half their

length, oblong, 12-15 mm long, 1.4-1.5 mm broad, api-

cally acute to obtuse, with one bract slightly longer than

the other, velutinous plus glandular hairs, bracteoles su-

bulate, apically tapered, 10-11 mm long, 1.2-1.3 mm

broad, velutinous; flowers sessile. Flowers with a 5-mer-

ous calyx, 6-7 mm long, segments equal, subulate, fused

for ca. half their length; corolla magenta, 24-25 mm long,

ca. 3 mm broad at the mouth, 0.7 mm broad basally,

puberulous, upper lip 1 1-12.5 mm long, 3-lobed, lobes

rounded, ca. 1 mm long, ca. 3.5 mm broad, lower lip

9.5-1 1 mm long, ca. 1.5 mm broad; stamens exserted,

anthers 1-celled, ca. 1.7 mm long, 0.6 mm broad. Fruits

clavate, 10-11 mm long, 1.4-1. 6 mm broad, 1.6-1. 7 mm

thick, puberulous toward apex; seeds 4, oval, 1.6-1.8

mm long, 1.1-1.2 mm broad, papillate.

This species is native to tropical Africa, but is

often cultivated for its attractive foliage. Escapes

from cultivation have been made along road banks

in two cloud forest areas in Costa Rica, the Mon-

teverde community at 1 500 to 2000 m elevation

in Puntarenas Province and near San Miguel at

about 1600 m elevation in Heredia Province.

Flowering collections were made in April and De-

cember.

Hypoestes phyllostachya is most easily recog-

nized by its pink-dotted leaves and magenta co-

rolla, with a three-lobed upper lip and one-lobed

lower lip. It is not easily confused with other Cen-

tral American species of Acanthaceae.

Justicia Linnaeus

Herbs, shrubs, or vines. Leaves petiolate, laminae usu-

ally ovate to elliptic and entire, cystoliths present and

usually prominent on both surfaces of dried specimens.

Inflorescences usually of spikes or panicles, occasionally

in cymes or solitary; bracts various, small and distant,

linear or subulate to large, conspicuous and imbricate.

Flowers with calyx segments usually narrow and equal,

some unequal, mostly 5(4)-merous; corollas bilabiate,

usually white, pink, or purple, the tube usually narrow,

funnelform, the upper lip 2-lobed, the lower lip 3-lobed;

stamens 2, usually slightly exserted, but not exceeding

the upper lip, anther cells 2, usually unequal, often su-

perposed, with 1 or both cells apiculate or calcarate, the

connective narrow to broad. Fruits a capsule, clavate, 4-

seeded.

This is the largest and most complex genus of

the Acanthaceae, with approximately 420 species

found mostly in the tropics, but extending well

into temperate habitats. It is worldwide in distri-

bution. The genus as treated here includes what,

until recently, was recognized as Beloperone and

Jacobinia.

The Costa Rican species of Justicia are wide

ranging, from near sea level to just above 2000 m

elevation. Most species are found between 500 and

2000 m in moist habitats around openings or along

trails in forests. Three species are found in dis-

turbed habitats. About five species are cultivated.

Justicia may be recognized by its two slightly

exserted stamens with two-celled anthers that are

frequently unequal, superposed, and basally apic-

ulate and calcarate. In addition, the bilabiate, nar-

row-tubed corollas and the lack of staminodes are

useful features.

Key to Species of Justicia

la. Inflorescences in spikes or cymes 2a

Ib. Inflorescences in panicles or thyrses (cymose panicles) 23a

2a. Inflorescences in axillary cymes 3a

2b. Inflorescences in spikes borne singly or in clusters 4a

3a. Leaves of a pair unequal; corolla violet or pale purple J. trichotoma

3b. Leaves of a pair equal; corolla orange-red to yellowish J. macrantha

4a. Leaves apically obtuse to rounded J. chamaephyton

4b. Leaves apically acuminate or acute 5a

5a. Bracts 4-19 mm long, imbricate 6a

5b. Bracts 1-3.5 mm long, lax (not imbricate) 20a

6a. Bracts spatulate; calyx segments 8-1 1 mm long 7a

6b. Bracts otherwise; calyx segments 4-7 mm long 8a

7a. Corolla 2.4-3.5 cm long; plant growing in open, disturbed habitats

J. carthaginensis

7b. Corolla 1.1-1.2 cm long; plant growing in shade of moist forests

J. costaricana

48

FIELDIANA: BOTANY

8a. Bracts reddish brown, 16-19 mm long; plants cultivated J. brandegeana

8b. Bracts green or brown, 4-1 1 mm long; plants not cultivated 9a

9a. Capsules 1 3-1 5 mm long; corollas 20-40 mm long 10a

9b. Capsules 5-10 mm long; corollas 7-19 mm long 1 la

lOa. Corollas white; bracts small, inconspicuous, 4-6 mm long, linear-subulate

J. urophylla

lOb. Corollas rose-purple; bract larger, 10-11 mm long, oblanceolate . . . J. brenesii

\ la. Leaf bases rounded to subcordate; leaves sessile to subsessile J. skutchii

1 Ib. Leaf bases acuminate, acute or obtuse; leaves with petioles mostly more than 5 mm

long (except J. refractifolid) 1 2a

12a. Calyx glandular-puberulous; lower cell of anther acute or rounded 13a

12b. Calyx puberulous or glabrous; lower cell of anther spurred 17a

1 3a. Bracts of inflorescence 4-6 mm broad; peduncles lacking glandular hairs

J. valerii

13b. Bracts of inflorescence 0.5-2.5 mm broad; peduncles with glandular hairs . 14a

14a. Corollas 15-19 mm long; calyx segments 5-7 mm long ..../. angustibracteata

14b. Corollas 7-1 1 mm long; calyx segments 2-4.5 mm long 1 5a

1 5a. Calyx segments (5) equal; spike lax J. tonduzii

15b. Calyx segments not equal (posterior calyx segment setaceous); spike compact

16a

1 6a. Bracts apically obtuse; leaves narrowly elliptical oblong J. parvibracteata

1 6b. Bracts apically acute; leaves lanceolate-elliptic J. pittieri

17a. Leaves no longer than 6.5 cm; posterior calyx segment much shorter than the rest . .

J. candelariae

17b. Leaves to as long as 18 cm; calyx segments of equal lengths 18a

18a. Bracts obovate to elliptic, strigose; corolla 16-17 mm long /. orosiensis

18b. Bracts ovate, linear-spatulate, spatulate or orbicular, pilose or glandular-puberulous;

corolla 7-1 2 mm long 1 9a

19a. Bracts linear-spatulate, spatulate or orbicular, glandular-puberulous; petioles glabrous

J. costaricana

1 9b. Bracts ovate, pilose; petioles pilose to tomentose /. refractifolia

20a. Leaves with auriculate bases, sessile to subsessile /. ephemera

20b. Leaves basally attenuate (lacking auricles), petiolate 2 la

2 la. Corolla 12-19 mm long, anther cells superposed; calyx segments 4 J. metallica

21b. Corolla 30-44 mm long, anther cells subequal; calyx segments 5 22a

22a. Leaves elliptic; corollas narrowly funnelform (not fusiform when unopened) . . . J. tinctoria

22b. Leaves ovate; corollas fusiform when unopened J. spicigera

23a. Capsules 4-6 mm long; corollas 4-6 mm long J. comata

23b. Capsules 7-25 mm long; corollas 8-70 mm long 24a

24a. Plant a climbing vine; corollas 65-70 mm long J. sarapiquensis

24b. Erect herbs or shrubs; corollas 8-60 mm long 25a

25a. Shrubs 2-6 m tall; anther cells equal or subequal, but not superposed 30a

25b. Herbs mostly less than 1 .5 m tall; anther cells superposed 26a

26a. Posterior calyx segment shorter, ca. half the length of the others; corollas 25-35 mm long

J. oerstedii

26b. Calyx segments of equal lengths; corollas 8-17 mm long 27a

27a. Inflorescence a compact spicate panicle resembling a single spike; bracts ovate to elliptic, 6-

8 mm long, 2.5-3.5 mm broad /. orosiensis

27b. Inflorescence a loose spicate panicle; bracts subulate to linear-subulate, 1 .5-2 mm long, 0.25-

1 mm broad 28a

28a. Leaves with auriculate bases, sessile to subsessile J. ephemera

28b. Leaves basally attenuate to rounded, lacking auricles, petiolate 29a

29a. Calyx segments 4, ca. 5 mm long; leaves ovate, 1.5-4 cm broad J. metallica

29b. Calyx segments 5, ca. 2 mm long; leaves lanceolate, 1-2 cm broad /. pectoralis

BURGER: FLORA COSTARICENSIS 49

30a.

30b.

33a.

Bracts subulate, 1-3 mm long; petioles short, 1 5 mm or less 3 la

Bracts linear-lanceolate, elliptic or obovate, 5-14 mm long; petioles long to more than 50 mm

long 33a

3 la. Flowers in axillary cymes; capsules 18-20 mm long /. macrantha

3 1 b. Flowers in loose spicate panicles; capsules 7-8 mm long 32a

32a. Leaves elliptic; corollas narrowly funnelform (not fusiform when unopened) . . . J. tinctoria

32b. Leaves ovate; corollas fusiform when unopened J. spicigera

Inflorescences in large, terminal spicate panicles, corollas red with yellow tip; capsules ca. 1.5 cm

long J. crenata

33b. Inflorescences in dense terminal or subterminal thyrses (cymose panicles), corollas yellow; capsules

2-2.5 cm long J. aurea

Justicia angustibracteata Leonard, Publ. Field

Mus. Nat. Hist., Bot. Ser. 18: 1227. 1938. Rhy-

tiglossa microphylla Oerst., Vidensk. Meddel.

Dansk Naturhist. Foren. Kjobenhavn 1854: 159.

1 855, non Justicia microphylla Lam., 1 79 1 . Fig-

ure 7.

Herb to 1 m tall, young stem quadrangular; internodes

between leaf-bearing nodes 1.5-4.5 cm long, 1.0 mm

thick, puberulous along 2 lines. Leaves with puberulous

to subglabrous petioles to 1 cm long; laminae ovate-

lanceolate, 1.5 (2)-5 (6) cm long, 1-1.5 (2) cm broad,

apically acuminate, basally acute, margins undulate, gla-

brous to sparingly and minutely scabrous above, more

so on costa and veins, puberulous below. Inflorescences

in lax terminal spikes to 6 cm long, 1 cm broad excluding

corollas; peduncle quadrangular, to 2 cm long, glandular-

puberulous, rachis quadrangular, glandular-puberulous;

bracts imbricate, oblanceolate, 10-11 mm long, 2-2.5

mm broad, ciliate, sparingly puberulous, bracteoles lin-

ear, 6.5-8.5 mm long, 0.5-0.75 mm broad, glandular-

puberulous. Flowers with 5-merous calyx, calyx seg-

ments unequal, linear subulate, the posterior segment

much smaller, ca. 3.5 mm long, 0.25 mm broad, the

other segments 5-6.5 mm long, 0.5-0.75 mm broad,

sparingly puberulous-glandular; corolla white with pur-

plish markings, 15-19 mm long, 2-2.5 mm broad at

base, ca. 3.5 mm broad at the throat, upper lip erect, 6-

7 mm long, 5-6 mm broad at base, narrowing to a keeled,

rounded apex, lower lip 6.5-7 mm long, 6-7 mm broad

at base, 3-lobed, the lobes ca. 2.5 mm long, semiorbic-

ular, stamens exserted, but not beyond the upper lip,

anthers strongly superposed, anthers with relatively wide

separation, connective 0.5 mm long, anthers 1.0 mm

long, 0.5 mm broad, apically and basally acute. Fruits

clavate, 5-7.5 mm long, 2.25 mm broad, 1-1.5 mm

thick, puberulous; seeds 2, flat, ovoid, dark brown, 1.5

mm long, 1.25 mm broad.

Found along streams and on steep slopes through

cloud forest areas of Cartago and San Jose prov-

inces at elevations of 1800 to 2000 m. Flowering

collections have been made in January, February,

March, and June.

Justicia angustibracteata is recognized by its ob-

lanceolate bracts, unequally segmented, five-mer-

ous calyx, and white corollas with purplish mark-

ings. This species is not easily confused with other

Justicia species when all of the above features are

visible. It bears a superficial resemblance to J. par-

vibracteata, but differs from it in its unequal calyx

segments, its longer (10 to 1 1 mm vs. 4 to 6 mm),

oblanceolate bracts (vs. elliptic) and its longer co-

rollas (15 to 19 mm vs. 6.5 to 7.5 mm).

Justicia aurea Schlecht., Linnaea 7: 393. 1832,

non Justicia aurea (Rose) Lindau, Bull. Herb.

Boissier. 5: 675. 1897. Justicia umbrosa Benth.,

PI. hartw. 79. 1 84 1 . Adhatoda umbrosa Nees in

DC., Prodr. 11: 406. 1847. Jacobinia aurea

Hemsl., Diagn. pi. nov. mexic. 35. 1879, non

Hiern, 1877-1878. Ecbolium umbrosum Kuntze,

Revis. gen. pi. 2: 981. 1891. Jacobinia umbrosa

Blake, Contr. Gray Herb. 52: 103. 1917. Fig-

ure 7.

A shrub to 6 m tall, stem subquadrangular to qua-

drangular in younger portions; internodes between leaf-

bearing nodes 2-5 cm long, 4-7 mm thick, puberulous

to tomentose. Leaves with petioles to 5.7 cm long, pu-

bescence that of the stem; laminae ovate to ovate-elliptic,

11.5-30 (40) cm long, 5.5-12 (17) cm broad, apically

acuminate, basally attenuate, margins entire, undulate

or crenulate, glabrous to sparingly strigose above, hairs

more dense on costa, glabrous to strigose below, cysto-

liths dense and prominent on both surfaces. Inflores-

cences densely thyrsoid at or near the ends of branches,

to 18 cm long, 6 cm thick excluding corollas; bracts

linear-lanceolate, 1 1-14 mm long, 1-2 mm broad; brac-

teoles similar, but slightly smaller, both puberulous.

Flowers with 5-merous calyx, segments lanceolate, 5.5-

7 mm long, 1.5 mm broad at base, puberulous; corolla

yellow, bilabiate, cylindrical, to 5.5 cm long, 3 mm broad,

puberulous-glandular outside, glabrous inside except for

bearded throat, the lips ca. 2.5 cm long, upper lip erect,

7 mm broad, apically acute, lower lip 6 mm broad, api-

cally 3-lobed, lobes ca. 1 mm long, apically rounded and

slightly hooded, especially the middle lobe; stamens

50

FIELDIANA: BOTANY

exserted to ca. the tip of the upper lip, anthers 3 mm

long, glabrous, the cells unequal, basally acute, filaments

stout, ca. 0.75 mm broad, glabrous; ovary 2 mm long,

1 mm thick, glabrous. Fruit clavate, puberulous, 2-2.5

cm long, ca. 5 mm broad, 4 mm thick; seeds 4, flattened,

suborbicular, ca. 2 mm in diameter, surface roughened.

This is a very attractive shrub, and it is fre-

quently cultivated. In the wild it is a relatively

wide-ranging species and is found from southern

Mexico to Panama. In Costa Rica it has been col-

lected in all provinces except Limon and Cartago,

generally in and adjacent to rain and cloud forest

from elevations of 100 to 2000 m.

Flowering collections have been made in all

months except May and August.

Justicia aurea is recognized by its large, attrac-

tive thyrsoid inflorescences with large, yellow co-

rollas (more than 5 cm long), its shrubby habit,

and its large leaves with long laminae (to 30 cm

or more long). It is not easily confused with other

species of Justicia.

Justicia brandegeana Wassh. & Smith, Fl. Ilus.

Catarinense ACAN:102. 1969; Beloperone

guttata Brandegee, Univ. Calif. Publ. Bot. 4:

278. 1912; non Justicia guttata Wall., PI. asiat.

rar. 1: 24. pi. 28. 1830. Figure 9.

Herb to subshrub, 1-2 m tall, steins subquadrangular;

in U- modes between leaf- bearing nodes 6-9.5 cm long,

1.5-2.5 mm thick, strigose mostly along 2 lines in older

portions. Leaves with densely strigose petioles 0.5-1 cm

long; laminae ovate-lanceolate, 3-3.5 cm long, 1-1.5 cm

broad, margins entire, apically acute to slightly attenuate,

apex blunt, basally acute, densely pilose on both surfaces,

cystoliths visible on both surfaces although obscured by

hairs. Inflorescences in terminal and axillary spikes to 6

cm long, 2 cm broad, excluding corollas; bracts reddish

brown, imbricate, ovate, 16-19 mm long, 11-13 mm

broad, apically acute, basally rounded to subcordate,

puberulous, ciliolate, bracteoles ovate-elliptic, 12-14 mm

long, 5-6 mm broad, apically acute, basally obtuse, pu-

berulous with ciliolate margins, peduncles 5-10 mm long,

densely strigose. Flowers with 5-merous calyx, segments

subulate, ca. 5 mm long, posterior segment 1.25 mm

broad basally, others ca. 0.75 mm broad, all densely

puberulous; corolla white, 3 longitudinal rows of pur-

plish dots in throat, 3.2 cm long, the tube narrow, 2 mm

broad at base expanding to only 3.5 mm at the mouth,

bilabiate, upper lip erect, 1 2 mm* long, 5 mm broad at

base, apically acute, lower lip ca. 1 1 mm long, 2-3 mm

broad, apex of 3 shallow, close, obtuse lobes; stamens

exserted to the tip of the upper lip, anther cells unequal,

the cells superposed ca. 3 mm long, basally spurred,

filaments glabrous. Fruits clavate, 1 2 mm long, 4 mm

broad, 3.5 mm thick, puberulous; seeds 4, dark brown,

subglobose, 3 mm long, 2.5 mm broad, 0.5 mm thick,

glabrous.

This plant is native to Mexico and is commonly

cultivated in much of Latin America.

Justicia brandegeana is recognized by its at-

tractive red bracts. These rather compact inflo-

rescences of reddish bracts give the plant its com-

mon name, the shrimp plant. In Costa Rica it is

called camaroncillo.

Justicia brenesii (Leonard) D. Gibson, Fieldiana,

Bot. 34: 66. 1972. Beloperone brenesii Leonard,

Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1201-

1202. 1938. Figure 8.

Herbs to 1 m tall, erect or decumbent with branches

frequent, occasionally rooting at lower nodes, younger

stems quadrangular with dense brownish pilose hairs to

1 mm long at the angles, older stems terete and glabrous;

intern (»d os between leaf-bearing nodes, 2-9 cm long, 1 .5-

2.5 mm thick. Leaves with petioles to 2.2 cm long, pilose;

laminae ovate-elliptic, 3.5-6.5 cm long, 2-3.5 cm broad,

apically short-acuminate, basally acute, margins entire

with pilose hairs, both surfaces glabrous to sparingly

pilose, cystoliths prominent and dense on both surfaces.

Inflorescences in dense terminal or lateral spikes or

spikelike thyrses to 5 cm long, 2.5 cm broad excluding

corollas; bracts imbricate, oblanceolate, 10-11 mm long,

3.5-4 mm broad, pilose and ciliate; bracteoles of similar

shape and length, but 1-2 mm broad, pilose and ciliate.

Flowers with 5-merous calyx, segments subulate, all ca.

6 mm long, posterior segment 1.5 mm broad at base,

lateral segments 1.25 mm broad, anterior segments 1

mm broad, all ciliolate; corolla rose-purple, to 2.5 cm

long, 2.5 mm broad at base expanding to 5.5 mm broad

at the mouth, glandular-puberulous, the lips ca. 10 mm

long, the upper lip erect, apically acute and notched,

lower lip 3-lobed, lobes ca. 3 mm long, apically obtuse;

the stamens exserted to just below the tip of the upper

lip, anther cells superposed, upper cells ca. 2 mm long,

lower cells slightly longer, all cells basally calcarate. Fruits

clavate, capsules ca. 13 mm long, 4 mm broad, 2.5 mm

thick, puberulous, hairs retrorse, seeds not seen.

Found in or adjacent to wet forest areas from

800 to 2200 m elevation. This is apparently an

endemic species collected in Alajuela and Heredia

provinces on both Caribbean and Pacific wa-

tershed zones. Flowering collections have been

made in September through April, except March.

Justicia brenesii is recognized by its stems with

densely pilose angles, its relatively broad, short,

and crowded spikes or thyrsi with pilose and ciliate

bracts, and its rose-purple corollas. It has some

resemblance to J. carthaginensis, but differs from

it in its generally smaller laminae, long-pilose

stems, and pilose and ciliate bracts.

Justicia candelariae (Oerst.) Leonard, Carnegie

Inst. Wash. Publ. 461: 231. 1936. Rhytiglossa

BURGER: FLORA COSTARICENSIS

51

candelariae Oerst., Vidensk. Meddel. Dansk

Naturhist. Foren. Kjebenhavn 1854: 158. 1855.

Dianthera candelariae Hemsl., Biol. Centralbl.

2:517. 1882. Figures.

Herbs to 45 cm tall, steins frequently decumbent, often

rooting at the lower nodes, quadrangular, pilose, densely

so in younger portions; internodes between leaf-bearing

nodes, 2.5 cm long, 1-1.5 mm thick. Leaves with short

petioles 3-10 mm long, pilose; laminae ovate-elliptic to

elliptic, 2.5-6.5 cm long, 1-2 cm broad, apically and

basal ly acute, margins entire, glabrous to sparingly pilose

above, sparingly pilose beneath, cystoliths visible on both

surfaces, more prominent beneath. Inflorescences in ter-

minal and axillary spikes to 7 cm long, 1.3 cm broad

excluding corollas, peduncles to 2.2 cm long, pilose; bracts

imbricate, spreading outward at an thesis, elliptic, 6.5-9

mm long, 3.5-5 mm broad, apically obtuse and mu-

cronulate, basally acute, vein pattern prominent, pilose

and ciliate; bracteoles narrowly elliptic, ca. 6 mm long,

1.5 mm broad, pilose and ciliate. Flowers with 5-merous

calyx, calyx segments unequal, segments narrowly subu-

late, posterior segment 2.5-3 mm long, 0.25 mm broad,

all others 4-6 mm long, 0.5-0.75 mm broad, ciliolate;

corollas white to pale purple, ca. 10 mm long, 2 mm

broad at base expanding to 2.5 mm at the throat, gla-

brous, upper lip acute, 3.5 mm long, 2.5 mm broad at

base, lower lip 4.5 mm long, 3-lobed, lobes 2 mm long,

middle lobe 2.75 mm broad, lateral lobes ca. 2 mm

broad, all apically rounded; stamens barely exserted, an-

ther cells unequal, ca. 0.5 mm long, upper cell slightly

larger, lower cell calcarate. Fruits clavate, capsules 7 mm

long, 2 mm broad, 1.5 mm thick, puberulous; seeds sub-

orbicular, 1.2 mm in diameter, papillate.

Herbs or suffrutescent plants to 1.5 m tall, stems te-

trangular and grooved in younger portions, sparingly

strigose mostly along grooves (sometimes along 2 lines)

in younger stems, glabrous in older stems; internodes

between leaf-bearing nodes, 2-7 cm long, 1-3 mm thick.

Leaves with petioles 0.5-2.5 cm long, puberulous; lam-

inae ovate to elliptic-ovate, 3-7 (14) cm long, 1.5-4(6.5)

cm broad, apically acuminate, basally attenuate, margins

entire, glabrous to strigose on the veins on both surfaces,

cystoliths abundant and prominent on both surfaces.

Inflorescences in terminal and axillary spikes, frequently

in clusters of 2 or 3, spikes 2-5.5 cm long, 2-3 cm broad

excluding corollas, peduncles short, frequently less than

1 cm long, puberulous; bracts imbricate, oblong-spatu-

late to spatulate, sizes varying from 10 mm long, 4 mm

broad near apex to 15(18) mm long, 7 (8) mm broad at

the base of the inflorescence, apiculate, puberulous with

glandular hairs toward the apex, margins ciliate; brac-

teoles similar, but slightly smaller. Flowers sessile, calyx

with 5 equal segments, segments linear-lanceolate, 8-15

mm long, 1-1.5 mm broad, puberulous, margins white,

ciliate; corolla rose-purple, 24-35 mm long, sparsely and

minutely puberulous outside, ca. 3 mm broad basally,

expanding to ca. 5 mm broad at the mouth, tube bearded

at 4 points inside just above the ovary, upper lip erect,

1 5-20 mm long, notched, lower lip 3-lobed, 1 5-20 mm

long, middle lobe ca. 7.5 mm broad, lateral lobes ca. 5.5

mm broad, middle lobe personate; stamens exserted ca.

10 mm beyond the mouth of the corolla tube, filaments

glabrous, anther cells 2.5-3 mm long, superposed, upper

cell 1.5 mm above the lower, lower cell calcarate at base,

upper cell blunt. Fruit clavate, capsules 13-17 mm long,

5-6 mm broad, 4 mm thick, apically apiculate, puber-

ulous; seeds very dark brown, shiny, suborbicular, 3-3.5

mm in diameter.

Collected in moist forests including oak forests

at elevations of 1200 to 2000 m in provinces of

Cartago and adjacent parts of San Jose. Flowering

collections have been made in November, Decem-

ber, January, and April.

Justicia candelariae is recognized by its pilose

stems, elliptic, subreticulate bracts with ciliate

margins, relatively small, inconspicuous corollas

( 1 cm long or less), and its unequal calyx segments.

It bears the same common Spanish name Zornia

as Blechum brownei, which it resembles superfi-

cally, but B. brownei is found at lower elevations,

has a near regular corolla, and has four stamens

instead of two. It also has some resemblance in

its inflorescence to Justicia parvibracteata, which

has much longer and narrower leaves, smaller

bracts, and a setaceous posterior calyx segment.

Justicia carthaginensis Jacq., Enum. syst. pi. 11.

1760, and in L., Syst. veg. ed. 16: 2: 153. 1817.

Adhatoda carthaginensis (Jacq.) Nees in DC.,

Prodr. 11: 403. 1847. Figure 8.

Found at lower elevations (0 to 100 m) of Gua-

nacaste and Puntarenas provinces in and adjacent

to tropical dry forests, frequently in secondary

growth and along roadsides. This plant is found

over a wide range from Mexico to South America.

It is abundant where found and occurs most often

in disturbed habitats. It is most frequently found

at lower elevations, but has been collected from

as high as 1300 m in some areas, although not

above 100 m in Costa Rica. This species exhibits

considerable variation in such features as corolla

length as well as size, shape, and hairiness of bracts,

leaves, and calyx. None of these variations is suf-

ficiently consistent to warrant the designation of

new species or varieties.

Justicia carthaginensis is recognized by its dense

spikes, conspicuous, large (13 to 17 mm long),

broad, clavate capsules. Because it is weedy also,

it has been confused with Blechum brownei, but

can be easily distinguished from it in number of

stamens (two vs. four), larger, bilabiate corolla,

bract shape (oblong-spatulate vs. ovate), and much

larger capsule (13 to 17 mm vs. 6 mm).

52

FIELDIANA: BOTANY

Justicia chamaephyton D. Gibson, Fieldiana, Hot.

34: 67. 1972. Beloperone variegata Lindau, An-

ales Inst. Fis.-Geogr. Nac. Costa Rica, 9: 189.

1898, and in Pitt., Prim. fl. costaric. 2: 315.

1900. Figure 8.

Herbs to 30 cm tall, stems ascending, rooting at lower

nodes, younger portion quadrangular, densely puberu-

lous, the hairs retrorse; internodes between leaf-bearing

nodes, 2.5-4 cm long, 1.5-2.5 mm thick, puberulous.

Leaves subsessile to petioles 1 cm long, petioles puber-

ulous; laminae elliptic, 5-12 (15) cm long, 2.5-5 (6.5)

cm broad, apically obtuse to rounded, basally attenuate,

margins undulate, glabrous above, puberulous on costa

and veins below, drying to a lighter green along the veins

above, cystoliths abundant and conspicuous on both sur-

faces. Inflorescences in loose terminal spikes to 8.5 cm

long, 4 mm broad (excluding corollas), peduncle to 3 cm

long, puberulous, rachis puberulous; bracts subulate, 2.5-

6 (7) mm long, 1 mm broad, puberulous, bracteoles sim-

ilar, but ca. 2 mm long, 0.5 mm broad. Flowers 1 per

bract; calyx 4-merous, segments subequal, 5-6 mm long,

0.5 mm broad, puberulous; corolla white or cream col-

ored suffused with yellow-green markings, ca. 10 mm

long, the tube narrow, 1 mm broad at base, 1.5 mm

broad at the throat, puberulous, the upper lip 5-6 mm

long, 3 mm broad, minutely bidentate at the tip, the

lower lip 5 mm long, middle lobe ca. 2.5 mm broad,

lateral lobes 2.5 mm long, 2 mm broad; stamens exserted

to just beyond the tip of the upper lip, anther cells su-

perposed, the lower calcarate. Fruits clavate, 10-1 1 mm

long, puberulous; seeds flat, glabrous, 2.5 mm in di-

ameter.

Collected in and along the margins of moist pri-

mary forests from the lowlands of Osa Peninsula

and the Golfo Dulce area at near sea level to the

Boruca area at ca. 300 m elevation, all in Puntar-

enas Province. This species is also found in low-

land areas of both the Pacific and Caribbean sides

of Panama, ranging from the Costa Rican border

to near Colombia.

Justicia chamaephyton is recognized by its small

size, slender, lax spikes, subulate bracts, and lam-

inae with obtuse or rounded bases and apices. Su-

perficially, J. chamaephyton resembles Aphelan-

dra tonduzii, but in addition to its different flowers,

it can be distinguished from it by its obtuse or

rounded leaf bases.

3.2-7 cm long, 2-2.5 mm thick. Leaves sessile; laminae

somewhat variable, ovate, elliptic or elliptic-oblong, 3.5-

10 cm long, 2.2-3.5 cm broad, apically acuminate, ba-

sally acute, rounded or obtuse, margins entire to slightly

undulate, glabrous to sparingly strigose along the costa,

cystoliths dense on both surfaces, especially so above.

Inflorescences in terminal and axillary spicate panicles,

the spikes whorled at the rachis nodes, the flowers secund

on the peduncle, the rachis and the branches grooved

with the branches generally somewhat flattened; rachis

and branches puberulous with a few longer glandular

hairs; bracts subulate, ca. 2 mm long, 0.5 mm broad,

keeled; bracteoles similar, slightly shorter. Flowers ses-

sile with 5-merous calyx, segments equal, lanceolate,

1.5-2 mm long, 0.5 mm broad at the base, glabrous;

corolla bilabiate, white, purplish, or white with purplish

dots or streaks, 4-6 mm long, 0.8 mm broad at base

expanding to 1.5 mm broad just above the base, gla-

brous, tube 3 mm long, the upper lip 3 mm long, 1.8

mm broad at base, apically acute, the lower lip 4 mm

long, 1.5 mm broad at base, expanding to 3 mm broad

above, 3-lobed at the tip, the lobes about equal, 0.5 mm

long, 1 mm broad, apically obtuse; stamens exserted ca.

2 mm beyond the corolla mouth, anther cells superposed,

ca. 0.3 mm long, filaments pilose toward base, glabrous

above. Fruits clavate, 4-5 mm long, 1.5 mm broad, 1

mm thick, puberulous; seeds 4, reddish brown, subor-

bicular, flattened, ca. 1 mm in diameter, papillose.

This is a species found in various disturbed hab-

itats, stream banks, cutover hills, roadsides and

along trails. It occurs throughout tropical America

and is generally found in lower elevations below

500 m, although one collection (Standley & Val-

erio 51873 F), has been made at La Hondura, San

Jose Province at 1200 to 1500 m. It has been

collected from all provinces in Costa Rica. Flow-

ering collections have been made in all months

except June, August, and September.

Justicia comata is recognized by its spicate pan-

icles that are both terminal and axillary, its small

corollas (4 to 6 mm long) and capsules (4 to 5 mm

long). It is most easily confused with J. pectoralis,

which has only terminal spicate panicles, longer

corollas (8 to 12 mm long), and capsules (7 to 8

mm long).

Justicia costaricana Leonard, Publ. Field Mus. Nat.

Hist., Bot. Ser. 18: 1229. 1938. Figure 8.

Justicia comata (L.) Lam., Encycl. 1: 632. 1783.

Dianthera comata L., Syst. nat. 10: 850. 1759.

Figure 8.

Herbs to suffrutescent plants to 1 m tall, stems some-

what grooved and hexagonal in younger portions, gla-

brous to pilose; internodes between leaf-bearing nodes,

Herbs to suffrutescent plants to 2 m tall, younger stems

quadrangular, older stems terete; internodes between leaf-

bearing nodes 2 (l)-7 cm long, 2 mm thick, puberulous

along 2 lines toward the apex. Leaves with glabrous pet-

ioles to 2 cm long; laminae ovate to elliptic, 5-15 cm

long, 2-5.5 cm broad, apically acuminate, basally acute

to obtuse, margins undulate, glabrous, cystoliths visible

on both surfaces, but more prominent above. Inflores-

BURGER: FLORA COSTARICENSIS

53

cence in loose, axillary spikes to 3 cm long, 1 cm broad,

rachis glandular-pubcrulous. peduncle to ca. 1 cm long,

puberulous along 2 lines; bracts linear-spatulate, spat-

ulate, or orbicular, 8-10 mm long, 0.5 mm to 8 mm

broad, glandular-puberulous, bracteoles linear, 8-10 mm

long, 0.25-0.5 mm broad, glandular-puberulous; bracts

and bracteoles usually spreading. Flowers with 5-merous

calyx, segments lanceolate, equal, 5.5-6.5 mm long, 1-

1.25 mm broad, puberulous; corolla saccate, glandular-

puberulous, pale green or greenish yellow with broad

dark red or purplish streaks at the upper part of the tube,

1 1.5-12.5 mm long, 1.5 mm broad basally, expanding

immediately to 4 mm broad just above base, bilabiate,

upper lip ca. 6 mm long, 6.5 mm broad, apically bilobed,

lobes acute, 0.5 mm long, 1 mm broad, lower lip 7-7.5

mm long, lateral lobes 4 mm long, 3.5 mm broad, middle

lobe 4.5 mm long, 3.5 mm broad, all lobes apically

rounded; stamens exserted to just below the upper lip,

anther cells superposed, upper cells basally apiculate, 1.5

mm long, 0.75 mm broad, filaments glabrous. Fruit not

seen.

This is an endemic species of mid-altitudes, 1 000

to 1500 m. It grows in the shade of moist forests

in both the Pacific and the Caribbean watersheds,

including the slopes of the Cordilleras Central,

Talamanca, and Tilaran. The collections of Palm-

er 144 and 161 (NY) from Monte verde, Puntar-

enas, have broader bracts, orbicular in shape, which

are different from those of all other collections,

but are otherwise identical. Flowering collections

have been made from January through April.

Justicia costaricana is recognized by its spread-

ing, spatulate to orbicular bracts, spreading linear

bracteoles, and greenish white, red-throated, sac-

cate corollas. There is variation in the shape of the

bracts which requires that other features be ex-

amined before identification is confirmed. It may

be confused with J. carthaginensis, but this species

has longer corollas (25 to 30 mm) and grows in

disturbed open habitats.

Justicia crenata (Leonard) Durkee, Ann. Missouri

Bot. Card., 65: 217. 1978. Jacobinia crenata

Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser.

18: 1223. 1938. Figure 9.

Shrub to 5 m tall, stem terete; internodes between leaf-

bearing nodes 2.5—4 cm long, 2-3 mm thick, glabrous.

Leaves with petioles to 5 cm long, glabrous; laminae

elliptic, 9-24 cm long, 2.5-6 (9) cm broad, apically acu-

minate, basally acute to attenuate, margins crenate, gla-

brous, cystoliths visible on both surfaces. Inflorescences

in large, terminal, spicate panicles to 27 cm long and 1 5

cm broad, peduncles and rachises puberulous; bracts el-

liptic to obovate, apically rounded to acute, soon decid-

uous, 5-10 mm long, 2-5 mm broad, sparingly glan-

dular-puberulous; bracteoles similar, but smaller. Flowers

with 5-merous calyx, the segments linear lanceolate,

equal, to 7 mm long, 1.25 mm broad basally, sparingly

glandular-puberulous, the margins minutely ciliolate; co-

rolla bilabiate, red with yellow tip, to 5 cm long, pu-

berulous glandular, the tube narrow, 3 mm broad at base

narrowing to 2 mm broad at the mouth, the lips erect,

each to 2.5 cm long, 6 mm broad, gradually narrowing

from base to tip, the upper lip apically notched, the lower

lip with 3 small lobes at the apex, the lobes each ca. 2

mm long, rounded and hooded apically; stamens exsert-

ed to the tip of the lobes, the anther cells unequal, each

to 2.5 mm long, each slightly spurred basally, the lower

cell more so. Fruit clavate, 1.5 cm long, glabrous or with

a few glandular hairs at the apex.

Found in rain forests of Limon Province, rang-

ing to cloud forests of higher elevations (to 1 500

m) of the Cordillera Central region. Also found in

similar habitats of adjacent Panama. Flowering

collections made in February and May through

November, except July.

Justicia crenata is recognized by its large, ter-

minal panicles of short spikes, its rounded bracts,

and its comparatively large, crenate leaves. It has

been confused with J. urophylla, which differs in

its dense spikes, linear, subulate bracts, entire to

undulate leaf margins, and white corolla.

Justicia ephemera Leonard, Contr. U.S. Natl.

Herb. 31: 606. 1958.

SufTrutescent plants to 2 m tall, stems upright, much-

branched; internodes between leaf- bearing nodes 5-1 1

cm long, 2-3.5 mm thick, upper internodes tetrangular,

glabrous to puberulous along 2 lines. Leaves sessile or

nearly so; laminae elliptic, 5-17 cm long, 1.3-5.5 cm

broad, apically long-acuminate, gradually narrowing to

an auriculate base, margins crenulate, glabrous on both

surfaces, cystoliths numerous, minute, visible on both

surfaces. Inflorescences in terminal, lax, spicate panicles

or single spikes in early flowering stages, to 10 cm long,

peduncles 2-3 cm long, puberulous; floral bracts subu-

late, 2-6 mm long, ca. 1 .0 mm broad. Flowers with 5-

merous calyx, calyx segments subulate, 2.5 cm long, ca.

0.75 mm broad, sparingly ciliolate; corolla white, bila-

biate, 10-13 mm long, 2 mm broad basally, glabrous,

the lips equal, 8-9 mm long, the upper lip ovate, apically

bidentate, 3.5-4 mm broad, lobes of lower lip ovate,

apically obtuse, 2.5-3 mm broad, middle lobe grooved

and laterally sculpted to form a nectar guide, slightly

broader than lateral lobes; stamens exserted to ca. 'A-'/s

the length of the corolla lobes, 3.5 mm long, glabrous,

anthers moderately superposed, 1 mm long, apically and

basally obtuse. Fruits not observed.

This species is found in wet areas of rain forests

in Colombia, Panama, and Costa Rica. One Costa

Rican collection, (Allen 5223, us), is known from

54

FIELDIANA: BOTANY

Puntarenas Province along the banks of the Rio

Grande de Terraba, on the trail from Palmar Norte

to Cana Blanca, at an elevation of 0 to 100 m.

This flowering collection was made in late March.

Flowering collections from Panama and Colombia

were also made in late March or April.

Justicia ephemera is recognized by its elliptic

leaves with auriculate bases and small (10 to 13

mm long), white corollas with barely exserted sta-

mens on broad, lax, spicate panicles. Because of

its unusually shaped leaves, it is not easily con-

fused with other Costa Rican species of Justicia.

Justicia macrantha Benth., PI. hartw. 78: 1839.

Figure 9.

Shrub to 3 m tall, older stems terete, quadrangular

toward apex; internodes between leaf-bearing nodes 2-

6 cm long, 1.5-3.5 mm thick, pilose to glabrate. Leaves

with petioles to 1.5 cm long; laminae elliptic, 6-16 (19)

cm long, 2.5-5.5 (6) cm broad, apically acuminate, ba-

sally attenuate, margins crenulate, glabrous on both sur-

faces except for pilose hairs in some, both surfaces fre-

quently gland-dotted, glands more abundant below,

cystoliths abundant on both surfaces. Inflorescences in

axillary cymes; bracts and bracteoles subulate, 2-3 mm

long, ca. 1 mm broad basally, glabrous, peduncles to 4.5

cm long, glabrous to sparingly pilose; pedicels 3-5 mm

long, glabrous. Flowers with 5-merous calyx, segments

subulate, 3-4 mm long, ca. 1 mm broad basally, keeled,

glabrous, usually ciliolate; corolla orange red to yellow-

ish, 4.5-5.5 cm long, deeply bilabiate, glabrous, the tube

funnelform, basally 4 mm broad, 8 mm broad at the

mouth, both lips ca. 2.8 cm long, the upper lip erect, 8

mm broad, apically dentate in 2 short lobes and hooded

to enclose part of the style, the lower lip spreading, 1 cm

broad, 3-lobed at the tip, the lobes about equal, 4 mm

long, apically rounded; stamens exserted to just below

the tip of the upper lip, the anthers pendulous, converg-

ing apically and diverging basally, the cells 2.5 mm long,

slightly unequal, basally rounded, the filaments stout, 1

mm broad, glabrous. Fruits clavate, 1 8-20 (22) mm long,

4 mm broad, 2.5 mm thick, puberulous; seeds oval,

flattened, 3 mm long, 2 mm wide, roughened.

This species is found through most of Central

America in forests at altitudes above 500 m. In

Costa Rica it has been collected in cloud forests

as high as 1 750 m, including the Cordilleras Gua-

nacaste, Tilaran (Monteverde), Central, and Tala-

manca. It has been collected in all provinces except

Limon. Flowering collections have been made from

November through April, excluding March.

Justicia macrantha is recognized by its axillary

cymes with large colorful corollas, which are

strongly bilabiate, and its pendulous anthers with

cells lacking spurs. It has been confused with /.

secunda, which has mostly terminal inflorescences

which are thyrsoid (cymose panicles) or panicu-

late, shorter corollas (2.5 to 4.3 cm long), upright

anthers, and shorter capsules (ca. 1 cm long).

Justicia metallica Lindau, Anales Inst. Fis.-Geogr.

Nac. Costa Rica, 9: 189. 1898, and in Pitt., Prim,

fl. costaric. 2: 311. 1900. Figure 9.

Herb to 1 m tall, stem ascending, rooting at the lower

nodes, younger stems quadrangular, the angles rounded,

pilosulus in 2 lines, mixed with glandular hairs toward

the apex, purplish; internodes between leaf-bearing nodes

1.5-6.5 cm long, 1-2 mm thick. Leaves with petioles 3-

10(15) mm long, pubescence of petioles that of the stem;

laminae ovate, 3-9 (12) cm long, 1.5-4 (6) cm broad,

apically acuminate to short acuminate, basally sparingly

attenuate to rounded, margins entire, undulate or cre-

nate, glabrous to sparingly strigose along the midrib and

the costa, the cystoliths prominent on both surfaces, dried

leaves greenish above, shiny metallic blue beneath. In-

florescences of lax, terminal or axillary, spicate panicles,

the spikes ca. 8 cm long, the first internode ca. 1 cm

long, the peduncles and rachises glandular-puberulous,

purplish; bracts and bracteoles linear subulate, to 1.5

mm long, 0.25 mm broad basally, glandular-puberulous.

Flowers with a 4-merous calyx, calyx segments subulate,

to 5 mm long, ca. 0.5 mm broad basally, glandular-

puberulous; corollas pink to purplish, 12-14 mm long,

sparingly puberulous, the tube cylindrical, to 9 mm long,

2 mm broad, the upper lip erect, to 4.5 mm long, 2 mm

broad, apically minutely bidentate, the lower lip spread-

ing, to 5.5 mm long, the lobes to 2 mm long, the middle

lobe 2 mm broad, the lateral lobes 1.5 mm broad, all

apically rounded; stamens exserted to just below the tip

of the upper lip, the anther cells superposed, ca. 1 mm

long, the lower cell spurred. Fruits clavate, capsules 7-

8 mm long, ca. 2 mm broad, 2 mm thick, sparingly

puberulous; seeds reddish brown, suborbicular, 1.5 mm

in diameter, roughened.

Found along stream banks and in wet places of

rain forests at elevations generally under 600 m.

This species has been collected in much of Panama

and in adjacent Costa Rica on the Pacific slope,

mostly in the vicinity of the basin of the Rio Gen-

eral and the Osa Peninsula. Flowering collections

have been made in Costa Rica in January and

February.

Justicia metallica is recognized by its leaves with

shiny blue metallic undersides, its purplish stems

and rachises, its four-parted calyx, and its lax spi-

cate panicles. It is not easily confused with other

species of the genus.

Justicia oerstedii Leonard, Publ. Field Mus. Nat.

Hist., Bot. Ser. 18: 1230. 1938. Figure 9.

BURGER: FLORA COSTARICENSIS

55

Herb to suffrutescent plant, 1.5 m tall, steins erect,

terete to subquadrangular when young; internodes be-

tween leaf-bearing nodes, 1.5-4.5 cm long, 0.75-1.25

mm thick, glabrous to puberulous. Leaves with petioles

to 5 mm long, petioles glabrous; laminae elliptic to ovate,

3-10 (12) cm long, 1-2.5 (3) cm broad, apically acu-

minate to acute, basally attenuate, margins entire to un-

dulate, glabrous on both surfaces, cystoliths prominent

on both surfaces. Inflorescences in terminal spicate pan-

icles to 20 cm long, the spikes lax, the peduncles and

rachises glandular-puberulous; bracts and bracteoles

narrowly subulate, 1-1.5 mm long, ca. 0.5 mm broad

basally, puberulous. Flowers with 5-merous calyx, the

calyx segments subulate, 3—4 mm long, 0.5 mm broad,

the posterior segment smaller, 1.5-2 mm long, 0.25 mm

broad, all segments keeled, puberulous; corolla bright

red, 2.5-3.5 cm long, puberulous, the tube narrow, 1

mm broad basally, 5-6 mm broad at the mouth, the

upper lip oblong, to 9 mm long, 5 mm broad, apically

minutely bidcntatc. the lower lip to 10.5 mm long,

3-lobed, the middle lobe 2.5 mm broad, the lateral lobes

1.5 mm broad, all lobes to 3 mm long, apically rounded;

stamens exserted to about the tip of the upper lip, the

anther cells superposed by a connective, 0.5 mm broad,

the lower cell rounded, acute at the base. Fruits clavate,

to 1 2 mm long, 2 mm broad, puberulous; seeds subor-

bicular, flattened, ca. 1.5 mm in diameter, roughened.

This species is abundant where found in rain

forest and cloud forest areas in moist, shaded lo-

cations along streams and trails at elevations of

500 to 1 800 m in Costa Rica and Panama. In many

locations this is the most abundant herb in flower.

In Costa Rica it is found mainly on the Pacific

watershed areas of all the Cordilleras from Pan-

ama to the Cordillera Guanacaste. It has not been

collected in Limon Province. Flowering collec-

tions have been made from December through

April.

Justicia oerstedii is recognized by its bright red

flowers with corollas more than 2 cm long, its lax,

spicate panicles with glandular-puberulous ra-

chises, and its five-parted calyx with one segment

much shorter than the others. It may be confused

with J. macrantha, which differs in its longer co-

rolla (more than 4 cm long), its axillary, cymose

inflorescences, and its four-parted calyx.

Justicia orosiensis Durkee, sp. nov. Figure 9.

Herbae vel suffrutices, usque ad 1.2 m altae, caulibus

erectis et difrusis. Folia petiolata, laminis ellipticis, 8-

23 cm longis, 2-7 cm latis, apice acuminatis, basi atten-

uatis, supra glabris, inferne plerumque glabris, cystolithis

nullis. Inflorescentiae paniculatae spiciformes, compac-

tae, terminales, bracteis laxe imbricatis, obovatis vel el-

lipticis, 6-8 mm longis, bracteolis similibus bractearum

sed minoribus. Flores virello-albi ad lilacini, corollis 1 2-

14 mm longis, staminibus exsertis. Fructus clavati, 12-

14 mm longi, seminibus 4.

Herbs to suffrutescent plants to 1.2m tall, stems erect,

but diffuse, quadrangular when young and dark maroon,

internodes between leaf-bearing nodes, 1.5-9.5 cm long,

1.5-3.5 mm thick, glabrous to sparingly puberulous.

Leaves with glabrous to sparingly puberulous petioles 1-

2 cm long; laminae elliptic, 8-23 cm long, 2-7 cm broad,

apically acuminate, basally attenuate, margins entire to

slightly undulate, glabrous above, glabrous to sparingly

puberulous along costa below, cystoliths obscure on both

surfaces. Inflorescences in compact terminal, spicate

panicles superficially resembling single spikes, 7-1 1 cm

long, 1.5-2 cm broad excluding corollas, peduncles and

rachises strigose; bracts and bracteoles obovate to ellip-

tic, bracts loosely imbricate, 6-8 mm long, 2.5-3.5 mm

broad, apically obtuse to rounded, basally attenuate, stri-

gose mostly at base, bracteoles similar although slightly

smaller. Flowers with 5-merous calyx, the calyx seg-

ments subulate, 6-7 mm long, 1.25-1.5 mm broad, gla-

brous; corolla greenish white to lilac, 16-17 mm long,

secund, glabrous, the tube cylindrical, 6 mm long, 2 mm

broad, the upper lip curved, oblong, partially enclosing

the stamens, to 11 mm long, 6 mm broad, apically

notched, the lower lip to 9.5 mm long, 3-lobed, the mid-

dle lobe 4 mm broad, the lateral lobes 3 mm broad, all

lobes to 7 mm long, apically rounded; stamens exserted

to about the tip of the upper lip, the anther cells super-

posed, 2.5 mm long, the lower cell calcarate, the upper

cell with apiculate base, pollen is 4-porate. Fruits clavate,

12-14 mm long, 4—5 mm broad, 3.5—4 mm thick, gla-

brous; seeds 4, suborbicular, ca. 3 mm in diameter.

TYPE— Costa Rica; Cartago Prov., 10 km south

of Tapanti, elevation 1 600 m, in primary forest

high over Rio Grande de Orosi, Lent 954 (holo-

type, F). Additional collections: Cartago Province,

forested slopes above Platanillo, elevation 1000

m, Wilbur and Stone 10611 (DUKE); along stream

near Tausito, elevation 1400 m, Lent 3818 (CR,

F). Flowering collections were made in mid-May,

late August, and February.

Justicia orosiensis is recognized by its compact

spicate panicles superficially resembling spikes and

its secund, greenish white corollas. It may possibly

be confused with the much more common and

wide-ranging J. carthaginensis of disturbed habi-

tats, which differs further in its dense spikes, larger

rose-purple corollas (24 to 35 mm long vs. 16 to

17 mm long), and larger (25 to 35 mm long), ob-

long-spatulate to spatulate bracts versus the 6- to

8-mm long, obovate to elliptic bracts of J. oro-

siensis.

Justicia parvibracteata Leonard, Publ. Field Mus.

Nat. Hist., Hot. Ser. 18: 1231. 1938. Figure 10.

56

FIELDIANA: BOTANY

Erect herb to 35 cm tall, stems quadrangular, terete in

older portions; inter nodes between leaf-bearing nodes 1-

4 cm long, 1 mm thick, puberulous mostly along 2 lines.

Leaves with puberulous petioles to 8 mm long; laminae

narrowly elliptic-oblong, 3.5-7 cm long, 0.7-1.2 cm

broad, margins entire to undulate, apically acute, basal 1 \

attenuate, cystoliths prominent on both surfaces, gla-

brous. Inflorescences in terminal and axillary spikes or

subpaniculate spikes to 6 cm long, 6 mm broad excluding

corollas; bracts barely imbricate, elliptic to obovate, 4-

6 mm long, 1-2 mm broad, apically obtuse, apiculate,

veins prominent, bracteoles narrowly elliptic, 3-4 mm

long, 0.5-0.75 mm broad, sparingly ciliolate, peduncles

0.5-2.5 cm long, puberulous mostly along 2 lines, rachis

glandular-puberulous. Flowers in fascicles of 2—4 per

bract; calyx 5-merous, posterior segment setaceous, 1.5-

2.5 mm long, the remainder lanceolate, 3.5-4.5 mm long,

ca. 0.5 mm broad, all glandular-puberulous, corolla white,

7.5-8 mm long, 1.5 mm broad basally, expanding to 2

mm broad at the mouth, sparingly puberulous, upper lip

3.5-4 mm long, 2 mm broad basally, apically acute,

lower lip 4-4.5 mm long, 2 mm broad, 3-lobed, lobes 1

mm long, ca. 1.5 mm broad, apically rounded; stamens

exserted to ca. Vi the length of the upper lip, anther cells

superposed, ca. 0.5 mm long, lower one slightly smaller,

rounded basally. Fruits clavate, 4.5-5.5 mm long, 1-1 .25

mm broad, 1-1.25 mm thick, sparingly puberulous, seeds

reddish brown, ovate, 1 mm long, 1 mm wide, papillate.

Found in wet areas along streams and river banks

on the Caribbean slope of the Cordillera Central

and on steep forested slopes of the Osa Peninsula,

at altitudes of 450 to 1400 m. Flowering collec-

tions have been made from January through

March.

Justicia parvibracteata is recognized by its nar-

row spikes with small (4 to 6 mm long), elliptic to

obovate bracts with narrow apices, short, white,

inconspicuous corollas (less than 1 mm long), and

its narrowly elliptic-oblong laminae. It has some

resemblance to J. candelariae and J. valerii, but

can be distinguished from these by its smaller bracts

(4 to 6 mm long vs. 6 to 10 mm) and narrower

leaves.

Justicia pectoralis Jacq., Enum. syst. pi. 11. 1760.

Figure 10.

Herbs to 2 m tall, stems weak, erect or ascending,

subquadrangular, grooved; internodes between leaf-

bearing nodes 2-4.5 mm long, 1-2 mm thick, retrorse

hairs mostly along 2 lines. Leaves with short strigose

petioles to ca. 5 mm long; laminae narrowly to broadly

lanceolate, 4-8 ( 1 0) cm long, 1 -2 (2.5) cm broad, margins

entire to undulate, apically acuminate, basally acute to

obtuse, cystoliths dense and prominent above, not vis-

ible beneath, glabrous to strigose at the costa above,

glabrous beneath. Inflorescences in terminal, lax, spicate

panicles to 25 cm long, 1 5 cm broad, the lower inter-

nodes of spicate branches ca. 2 cm long, rachises and

peduncles glandular-puberulous; bracts and bracteoles

subulate, 2 mm long, 1 mm broad basally, puberulous

with a few glandular hairs. Flowers with a 5-merous

calyx, segments narrowly subulate to 2 mm long, 0.5

mm wide, basally glandular puberulous; corollas white,

lilac or purple, 8-12 mm long, puberulous outside, the

throat transversely plicate, often spotted with dark pur-

ple, the tube narrowly funnelform, basally 1 .25 mm broad,

2.5 mm broad at the mouth, the upper lip erect, trian-

gular 3.5 mm long, 3 mm broad, apically acute, the lower

lip spreading, to 5 mm long, 3-lobed, the lobes 1 mm

long, the middle lobe 2 mm broad, the lateral lobes 1.5

mm broad, all apically obtuse; stamens exserted to about

the tip of the upper lip, the anther cells slightly super-

posed, basally apiculate, the filaments glabrous. Fruits

clavate, capsules 7-8 mm long, 1.5-2 mm broad, ca. 1

mm thick, puberulous; seeds orbicular, flattened, 1.5 mm

in diameter.

This is a wide-ranging species of roadsides and

waste places in tropical America. In Costa Rica it

occurs from near sea level to 900 m altitude and

has been collected in all provinces except Alajuela.

Flowering collections have been made from Jan-

uary through May.

Justicia pectoralis is recognized by its terminal

spicate panicles, with corollas 8 to 12 mm long,

puberulous capsules 7 to 8 mm long, and lanceo-

late laminae 4 to 8 mm long. It is most easily

confused with J. comata, but can be distinguished

from it by its terminal spicate panicles (vs. ter-

minal and axillary spicate panicles), longer corol-

las (8 to 12 mm vs. 4 to 6 mm long), and longer,

puberulous capsules (7 to 8 mm long vs. papillose

capsules, 4 to 5 mm long).

Justicia pittieri Lindau, Anales Inst. Fis.-Geogr.

Nac. Costa Rica 9: 1 89. 1 898, and in Pitt., Prim,

fl. costaric. 2: 312. 1900. Figure 10.

Herbs to 50 cm tall, stems ascending, younger stems

quadrangular, grooved, retrorse hairs mostly along 2 lines;

internodes between leaf-bearing nodes 5.5-7 cm long,

1.5-2 mm thick. Leaves petiolate, petioles short, 1-10

mm long, strigose; laminae lanceolate-elliptic, 5-12 cm

long, 2-3 cm broad, margins entire, apically acuminate,

basally acute, glabrous to sparingly strigose, cystoliths

dense and prominent on both surfaces. Inflorescences in

terminal and axillary spikes, to 1 5 cm long, 1 cm broad,

excluding corollas, rachises and peduncles with retrorse

pubescence mostly along 2 lines, plus some glandular

hairs, peduncle to 1 1 cm long, bracts imbricate, spread-

ing at anthesis, elliptic, 4.5-5.5 mm long, 1.5-2.2 mm

broad, apically and basally acute, subglabrous, glandu-

lar-puberulous, ciliate, bracteoles elliptic, 3.5-4 mm long,

0.5-0.75 mm broad, glandular-puberulous, ciliate. Flow-

BURGER: FLORA COSTARICENSIS

57

ers in fascicles of 2-3 per bract, calyx 5-merous, posterior

segment setaceous, 2 mm long, remaining segments su-

bulate, 3.5-4.5 mm long, 0.5-0.75 mm broad, glandular-

puberulous; corollas white with purple dots, 7-9 mm

long, 1 mm broad at the base expanding to 2.25 mm

broad at the mouth, puberulous with a few glandular

hairs, the throat transversely plicate, the plaits purplish,

upper lip 5 mm long, 2 mm broad basally, apically bi-

dentate, lower lip 3.5 mm long, the lobes orbicular, ca.

1 mm in diameter; stamens exserted ca. '/z the length of

the upper lip, anther cells superposed, ca. 0.5 mm long,

basally obtuse, lower cell slightly smaller. Fruits not seen.

Only two collections of this species have been

observed: the type, Pittier 8642 (CR) from lowland

rain forests of the upper Rio Yorkin between Costa

Rica and Panama and Skutch 4652 (F, MO) at 600

to 850 m altitude in the vicinity of Pejibaye (Pe-

jivalle), Cartago Province. These flowering collec-

tions were made in January and March.

Justicia pittieri is recognized by its inflores-

cences with elliptic bracts (4.5 to 5.5 mm long),

which spread at anthesis, flowers in fascicles of

two to three per bract, unequal calyx segments,

and stems with retrorse hairs mostly along two

lines. Justicia pittieri may be confused with J. ton-

duzii, but differs from it in its unequal calyx seg-

ments, elliptic bracts (vs. oblanceolate), and gla-

brous to sparingly strigose leaves (vs. pilose).

lobes 2-2.5 mm long, apically obtuse, the middle lobe

2-2.5 mm broad, the lateral lobes 1.5-2 mm broad, the

stamens exserted to ca. the tip of the upper lip, the fil-

aments attached to the tube just below the mouth, pu-

bescence of minute glandular hairs, the anther cells su-

perposed, the upper cell mucronate, the lower cell spurred.

Fruits clavate, capsules 7.5-10 mm long, 2 mm broad,

2 mm thick, puberulous; seeds ovate, ca. 1.5 mm long,

1.5 mm broad, flattened, roughened.

Found at lower elevations frequently in moist,

disturbed areas that are shaded in both Costa Rica

and Panama. It has been collected only in the Ca-

ribbean lowlands of Costa Rica. Flowering collec-

tions have been made in November, December,

and July.

Justicia refractifolia is recognized by its axillary

spikes with small, imbricate, ovate bracts which

are brown when dried and have a prominent re-

ticulate venation and ciliate margins; its small,

relatively inconspicuous flowers (ca. 1 5 mm long)

with superposed anther cells that are spurred only

on the lower cell; and its rather large (11 to 18 cm

long), elliptic-obovate leaves. Justicia refractifolia

has some resemblance to J. refulgens, but can be

distinguished from it by its unbranched spikes (vs.

trifurcate spikes), ovate ciliate bracts (vs. lanceo-

late without cilia), and longer bracts (8 to 10 mm

vs. 6 mm long).

Justicia refractifolia (Kuntze) Leonard, Publ. Field

Mus. Nat. Hist., Bot. Ser. 18: 1232. 1938. Ec-

bolium refractifolium Kuntze, Revis. gen. pi. 2:

487. 1 89 1 . Justicia pseudopolystachia Cuf., Arch.

Bot. Sist. 10: 49. 1934. Figure 10.

Herbs to suffrutescent plants, 1 m tall, stems simple,

erect, subquadrangular toward the apex, subterete below;

internodes between leaf- bearing nodes 4-10 cm long,

1.75-2.5 mm thick, pilose to tomentose. Leaves with

pilose to tomentose petioles to 3 mm long; laminae el-

liptic to elliptic-obovate, 1 1-18 (21)cm long, 3-6.5 (8.5)

cm broad, margins entire to undulate, apically acumi-

nate, basally acute to attenuate, glabrous to strigose on

the costa and veins above, pilose to sparingly strigose

beneath, cystoliths prominent to obscure above, obscure

beneath. Inflorescences in terminal and axillary spikes,

borne singly and in clusters, to 1 5 cm long, 1 cm broad,

the rachis pilose to strigose; bracts ovate, 8-10 mm long,

3.5-5 mm broad, imbricate, apically acute, basally ob-

tuse, drying brown, the veins prominent, sparingly pilose

on both surfaces, ciliate; bracteoles lanceolate, slightly

falcate, 6.5-9 mm long, 1.5-2 mm broad, puberulous,

ciliolate. Flowers with 5-merous calyx, segments nar-

rowly subulate, 5.5-6 mm long, 0.5-0.75 mm broad,

puberulous, ciliolate; corolla white to purple, the tube

cylindrical, 7-8 mm long, 1.5 mm broad, puberulous,

the upper lip erect, ca. 4—4.5 mm long, 2 mm broad,

apically bidentate, lower lip 4-5 mm long, 3-lobed, the

Justicia sarapiquensis McDade, Syst. Bot. 7: 489-

493. 1982. Figure 10.

Suffrutescent vine climbing to 15 m; internodes be-

tween leaf-bearing nodes 1.5-2.5 cm long, 2-2.5 mm

thick, subquadrangular, sparingly puberulous, older stems

terete. Leaves with sparingly puberulous petioles 5-10

mm long; laminae ovate, 4.5-7.7 cm long, 2.2-3.5 cm

broad apically acuminate, basally obtuse to rounded or

slightly cordate, margins entire, glabrous on both sur-

faces, cystoliths prominent on both surfaces. Inflores-

cences in lax, terminal panicles of up to 15 flowers, pe-

duncles and branches sparingly puberulous; bracts

subtending the flowers borne singly or in pairs, ovate,

magenta, 15-21 mm long, 14-16 mm broad, apically

obtuse, glabrous, bracts on lower parts of peduncle small-

er; bracteoles 2 per flower, lanceolate, magenta, 16-19

mm long, 6-10 mm broad, apically acute, glabrous.

Flowers with 5-merous calyx, calyx lobes equal, 1 5-22

mm long, 2-4 mm broad, white, apically acute and apic-

ulate, glabrous; corolla white, 6.5-7 cm long, outside

with fairly dense white pilose hairs 1-1.5 mm long, the

tube 38—40 mm long, 5-6 mm broad at base, narrowed

to 2.5-3 mm broad above the ovary, expanding to 5-6

mm broad at the throat, the upper lip erect, 23-26 mm

long, apically acute, the lower lip slightly reflexed at an-

thesis, 27-29 mm long, apically 3-lobed, each lobe 3-4

mm long; stamens extending to just below the upper lip,

anther cells slightly superposed, ca. 4 mm long. Fruits

58

FIELDIANA: BOTANY

clavate, dark brown, 15-20 mm long, 5-6 mm broad,

3-4 mm thick, glabrous, floral bracts, bracteoles, and

calyx are strongly reflexed when fruits are mature; seeds

4, orbicular, flattened, 3—4 mm in diameter, glabrous.

This species grows in shaded areas of the rain

forest lowlands of northern Costa Rica at eleva-

tions of 100 to 575 m. It has been collected in

only two sites, near Quesada (Villa Quesada), San

Carlos, and the La Selva Research Station, both

in Heredia Province. Flowering collections have

been made in March.

Justicia sarapiquensis is recognized by its viny

habit, its large magenta bracts and bracteoles, its

white calyx, and its white, pilose corolla. It is not

easily confused with other species of Justicia from

Costa Rica; however, the Panamanian species, /.

graciliflora (Standley) D. Gibson, has a viny habit,

but lacks the large, showy bracts and bracteoles.

Justicia skutchii Leonard, Publ. Field Mus. Nat.

Hist., Bot. Ser. 18: 1234. 1938. Figure 10.

Shrub to 1 m tall, stems quadrangular, older portions

terete; internodes between leaf-bearing nodes 1-3.5 cm

long, 1.5-2 mm thick, densely pilose. Leaves subsessile;

laminae ovate-elliptic, 3.5-13 cm long, 3.5-5 cm broad,

margins entire to slightly undulate, apically acuminate,

basally rounded or subcordate, cystoliths prominent on

both surfaces, sparingly pilose above, pilose beneath.

Inflorescences in terminal spicate panicles, spikes lax, to

9.5 cm long, 8 mm broad excluding corollas, peduncle

and rachis pilose-glandular, bracts elliptic, 3.5-4 mm

long, 1 mm broad, sparingly puberulous, ciliolate, brac-

teoles linear, 3-3.5 mm long, ca. 0.25 mm broad, spar-

ingly glandular-puberulous. Flowers in clusters of 2-3

per bract, calyx 5-merous, posterior segment setaceous,

3.5-4 mm long, puberulous, other segments subulate-

linear, 3.4-5 mm long, 0.5 mm broad, puberulous; co-

rolla white and purple, glabrous, 7.5-9 mm long, tube

cylindrical 4.5-5 mm long, 2 mm broad, upper lip acute,

ca. 3 mm long, 2 mm broad, lower lip ca. 3 mm long,

lobes 0.5 mm long, rounded, lateral lobes 1 .25 mm broad,

middle lobe 2 mm broad; stamens exserted to ca. '/2 the

length of the upper lip, cells unequal, 0.5 mm long, ba-

sally acute. Fruits clavate, capsules 8.5-9 mm long, 2

mm broad, 2 mm thick, puberulous, seeds reddish brown,

ovate, 1.75 mm long, 1.5 mm broad.

Found in moist habitats on rocks along a stream

in a forest in the vicinity of General Viejo (El

General), San Jose Province, at 9 1 5 m elevation.

Known only from the type collection, Skutch 3012

(MO).

Justicia skutchii is recognized by its lax, spicate

panicles with small (3.5 to 4 mm long), elliptic

bracts, dense yellowish pilose hairs on the stem,

and basally rounded to subcordate leaves. It re-

sembles J. tonduzii, but can be distinguished from

it by its leaves with rounded to subcordate bases

(vs. obtuse or narrowed bases) and its longer cap-

sules (8.5-9 mm long vs. 5 mm).

Justicia spicigera Schlecht., Linnea 7: 395. 1832.

Jacobinia spicigera (Schlecht.) L. H. Bailey,

Stand. Cycl. Hort. 1715. 1915. Figure 11.

Erect or scandent shrub, 1-1.5 m tall; internodes be-

tween leaf-bearing nodes 1-5 cm long, 1-2 mm thick,

quadrangular, glabrous to puberulous along 2 lines. Leaves

with petioles 5-10 mm long, petioles mostly puberulous

along 2 lines; laminae ovate, 6.5-17 cm long, 3.5-9 cm

broad, apically acuminate, basally attenuate, margins en-

tire to undulate, cystoliths abundant and prominent on

both surfaces, dried leaves often purplish black. Inflo-

rescences in terminal and axillary spicate panicles, flow-

ers secund, peduncles and rachises puberulous; bracts

and bracteoles subulate, 1-1.5 mm long, glabrous. Flow-

ers with a 5-merous calyx, calyx segments subulate, 2.5-

3 mm long, 0.75-1 mm broad basally, glabrous; corollas

orange, 3.5-4.4 cm long, appearing fusiform before

opening, glabrous, the upper lip erect, ca. 1 5 mm long,

6.5 mm broad, apically acute, the lower lip ca. 15 mm

long, 5 mm broad, frequently rolled up, lobes small,

suborbicular, 1.5 mm long, 1.5 mm broad; stamens

exserted approximately to the tip of the upper lip, the

anther cells subequal, ca. 2.5 mm long, basally apiculate.

Fruits not seen.

Found along stream banks in thickets and for-

ests of Mexico and Central America, this plant is

often cultivated for use as laundry bluing and in

some areas as a dye. Only one collection of this

plant, Skutch 4809 (F, MO, NY), has been made in

Costa Rica, in the Rio General basin in March. It

had abundant flowers, but no fruits.

Justicia spicigera is recognized by its frequently

darkened, ovate leaves, axillary spicate panicles

with secund flowers, and orange corollas (3.5 to

4.4 cm long) which are fusiform when unopened.

It is easily confused with J. tinctoria, which is also

cultivated for the same purposes, but can be dis-

tinguished from it by its fusiform corollas (vs. nar-

rowly funnelform) and its larger, ovate leaves (6.5

to 1 7 cm long vs. elliptic leaves 4.5 to 14 cm long).

Justicia tinctoria (Oerst.) D. Gibson, Fieldiana,

Bot. 34(6): 74. 1972. Sericographis tinctoria

Oerst., Vidensk. Meddel. Dansk Naturhist. For-

en. Kjobenhavn 1854: 150. 1855. Jacobinia

tinctoria (Oerst.) Hemsley, Biol. Centralbl. 2:

552. 1882. Figure 11.

BURGER: FLORA COSTARICENSIS

59

Shrubs to 2 m tall, younger steins subquadrangular,

the angles rounded; internodes between leaf-bearing nodes

1.5-5 cm long, 1.5-2.5 mm thick, glabrous. Leaves with

glabrous petioles to 1.2 cm long; laminae elliptic, 4.5-

14 cm long, 1.5-4.5 cm broad, margins entire to un-

dulate, apically acute to acuminate, basally attenuate,

cystoliths visible on both surfaces, but more prominent

below. Inflorescences in lax axillary spikes or spicate

panicles with secund flowers, occasionally branched,

spikes to 4 cm long, peduncles to 2 cm long, glabrous to

puberulous; rachis subquadrangular, internodes 4-5 mm

long; bracts and bractoles subulate, 1-2 mm long, 0.5-

1 mm broad at base. Flowers with 5-merous calyx, calyx

segments subulate, 2.25-3 mm long, 1 mm broad at base,

glabrous; corolla orange or red, 3-4 cm long, the tube

slightly funnelform, 3 mm broad basally, 5 mm broad

at the mouth, the lips subequal, ca. 16 mm long, the

upper lip erect, 7 mm broad basally, apically acute and

keeled, the lower lip oblong, erect or slightly spreading,

4.5 mm broad, apically 3-lobed, the lobes equal, 2 mm

long, 1 mm broad, apically obtuse and keeled; stamens

exserted to about the tips of the lips, the anther cells

subequal, 2.5 mm long, obliquely attached to the con-

nective, mucronulate, filaments glabrous. Fruits not seen.

First described from Costa Rica, this species is

found in dry thickets of Central America and is

often cultivated. When steeped in hot wtaer, its

leaves produce a bluing solution, azul de mata,

used in laundering white clothes. Flowering col-

lections have been made in January, March, April,

and June.

Justicia tinctoria is recognized by its short ax-

illary spikes with secund flowers, red or orange,

narrowly funnelform, corollas (3 to 4 cm long),

and short calyx segments (2.25 to 3 mm long). It

is easily confused with J. spicigera, but can be

distinguished from it by its smaller elliptic leaves

(4.5 to 14 cm long vs. larger ovate leaves 7-17 cm

long), and narrowly funnelform corollas (vs. fu-

siform corollas).

Justicia tonduzii Lindau, Anales Inst. Fis.-Geogr.

Nac. Costa Rica, 9: 189. 1898, and in Pitt., Prim,

fl. costaric. 2: 313. 1900. Figure 1 1.

Herbs to 25 cm tall, stem ascending, branching, youn-

ger stems terete to subquadrangular, pilose hairs to 3

mm long; internodes between leaf-bearing nodes, 2-4

cm long, 1-1.5 mm thick. Leaves with pilose petioles to

5 mm long; laminae ovate to ovate-elliptic, 5-8 cm long,

2-3.2 cm broad, margins undulate, apically acuminate,

basally obtuse to rounded, pilose on both surfaces, cys-

toliths dense and prominent on both surfaces. Inflores-

censes in lax terminal and axillary spikes to 7 cm long,

rachiscs and peduncles pilose plus shorter glandular hairs,

bracts not imbricate at anthesis. oblanceolate, 2.5-3 mm

broad, apically acute, pilose, bracteoles linear ca. 2.5 mm

long, 0.5 mm broad, pilose. Flowers with 5-merous ca-

lyx, calyx segments equal, subulate, 2-2.5 mm long, glan-

dular-puberulous; corollas purplish, ca. 5-7 mm long,

tube cylindrical, ca. 1.5 mm broad, glabrous, the throat

longitudinally furrowed, upper lip 2.5 mm long, 1.5 mm

broad, apically acute, upper lip 2.5 mm long, 1.5 mm

broad, apically 3-lobed, lobes ca. 0.5 mm long, apically

rounded; stamens exserted ca. '/2 the length of the upper

lip, anther cells superposed, upper cell larger, basally

acute, lower cell basally rounded. Fruits clavate, 6 mm

long, 2 mm broad, puberulous, seeds 4, orbicular, 1 mm

in diameter.

Only one collection of this species has been ob-

served, Pittier 16073 (us), from the forests of Vic-

toria Farm, plains of Zent, 200 m elevation, Li-

mon Province. This flowering collection was made

in January.

Justicia tonduzii is recognized by its lax inflo-

rescences, with minute, oblanceolate bracts (2.5 to

3 mm long), equal calyx segments, and long pilose

hairs of stem, inflorescence, rachis, and leaves.

Justicia tonduzii may be confused with J. pittieri,

but differs from it in its equal calyx segments,

oblanceolate bracts (vs. elliptic), and long pilose

hairs.

Justicia trichotoma (Kuntze) Leonard, Publ. Field

Mus. Nat. Hist., Bot. Ser. 18: 1236. 1938. Ec-

bolium trichotomum Kuntze, Revis. gen. pi. 2:

488. 1891. Justicia asymmetrica Lindau, Anales

Inst. Fis.-Geogr. Nac. Costa Rica, 9: 189. 1898,

and in Pitt., Prim. fl. costaric. 2: 310. 1900.

Figure 1 1 .

Herbs or small shrubs to 2 m tall, stem erect, subqua-

drangular, lower portion terete; internodes between leaf-

bearing nodes 1-5 cm long, 1-2 mm thick, strigose. Leaves

with strigose petioles to 2 cm long; laminae ovate to

ovate-lanceolate, 1 of each pair conspicuously larger than

the other, the larger 5-9 cm long, 2-4.5 cm broad, the

smaller 2-5.5 cm long, 1-2 cm broad, all apically acu-

minate, basally attenuate, margins undulate, sparingly

strigose to glabrous above, similar below but more dense,

cystoliths often inconspicuous, more visible on lower

surface. Inflorescences in small, axillary cymes, pedun-

cles to 2.7 cm long, strigose; bracts and bracteoles subu-

late, to 1.5 mm long, 0.5 mm broad basally, strigose.

Flowers sessile, calyx 5-merous, segments narrowly su-

bulate, to 6 mm long, 1 mm broad, puberulous; corolla

white, pink or violet, 16-20 mm long, puberulous, the

tube funnelform, 1 .5 mm in diameter basally, expanding

to 4 mm in diameter at the mouth, upper lip to 6 mm

long, 4 mm broad, apically acute with a rounded apex,

lower lip to 7 mm long, 3-lobed, lobes 3 mm long, api-

cally obtuse, middle lobe 2.5 mm broad, lateral lobes 2

mm broad; stamens exserted to about the tip of the upper

lip, anther cells superposed, lower cells calcarate. Fruits

clavate, to 1 1 mm long, 2.5 mm broad, 1 mm thick,

puberulous; seeds 4, dark brown, suborbicular, ca. 2 mm

in diameter, roughened.

60

FIELDIANA: BOTANY

Found in wet habitats adjacent to streams, in

clearings of rain forests and wet thickets in Pan-

ama and Costa Rica, mostly at elevations below

1000 m. In Costa Rica collections have been made

only in the Caribbean watershed. Flowering col-

lections have been made in February, March, April,

and June.

Justicia trichotoma is not easily confused with

other species in the genus, because of the unequal

size of the leaf pairs.

Justicia urophylla (Lindau) D. Gibson, Fieldiana,

Bot. 34: 74. 1972. Beloperone urophylla Lindau,

Anales Inst. Fis.-Geogr. Nac. Costa Rica, 9: 1 89.

1898, and in Pitt., Prim. fl. costaric. 2: 316.

1900. Figure 11.

Shrub to 2 m tall, stems subterete; internodes between

leaf-bearing nodes 2-7 cm long, 1.5-2.5 mm thick, gla-

brous to sparingly puberulous in 2 lines. Leaves with

petioles to 3 cm long, pubescence that of the stem; lam-

inae ovate-elliptic, 4.5-13 cm long, 2-5 cm broad, mar-

gins entire to undulate, apically acuminate, basally at-

tenuate, glabrous, cystoliths dense, plainly visible on both

surfaces. Inflorescences in 1 -several dense terminal spikes

to 6 cm long, ca. 6 mm broad excluding corollas, the

peduncles to 2 cm long, puberulous mostly along 2 lines,

the rachis glabrous to puberulous; bracts imbricate, lin-

ear-subulate 4-6 mm long, ca. 1 mm broad basally, pu-

berulous, ciliolate, bracteoles triangular, ca. 1 mm long,

0.5 mm broad basally, puberulous. Flowers imbedded

in the rachis, calyx segments 5, equal, subulate, 2-3 mm

long, ca. 0.5 mm broad, puberulous; corolla white, 3-4

cm long, puberulous, the tube slightly funnelform, 2 mm

broad basally, narrower just above the ovary, ca. 3 mm

broad at the mouth, the lips 1 1-16 mm long, the upper

lip erect, apically bidentate, the lower lip 3-lobed, the

lobes 7-10 mm long, the middle lobe 4 mm broad, the

lateral lobes 3 mm broad, all apically rounded; stamens

exserted ca. 2 mm below tips of the lips, anther cells

superposed, basally acute, ca. 2 mm long, connective ca.

1 mm long, the filaments glabrous. Fruits clavate, cap-

sules 13-15 mm long, 4 mm broad, 3 mm thick, gla-

brous, seeds light brown, suborbicular, ca. 3 mm in di-

ameter, glabrous.

Found in rain forests of Costa Rica and Panama

from sea level to about 1 100 m elevation. In Costa

Rica numerous collections have been made, but

all are from the vicinity of San Ramon in Alajuela

Province. Flowering collections have been made

in Costa Rica in February and March.

Justicia urophylla is recognized by its dense, ter-

minal spikes witih short, inconspicuous bracts (4

to 6 mm long), its white corollas (more than 3 cm

long), and its equally five-parted calyx with short

segments (2 to 3 mm long). It may be confused

with J. secunda, which is found in adjacent regions

of Panama, but has not been collected in Costa

Rica. It can be distinguished from J. secunda by

its dense spikes (vs. a looser, paniculate to thyrsoid

inflorescence), white corollas (vs. red or purplish

red), and its leaves with basally attenuate laminae

(vs. acute or obtuse).

Justicia valerii Leonard, Publ. Field Mus. Nat.

Hist, Bot. Ser. 18: 1236. 1938. Figure 11.

Decumbent herb to 30 cm tall, stems subquadrangular

to near terete in older portions; internodes between leaf-

bearing nodes 1.5-7 cm long, ca. 1 mm thick, puberu-

lence retrorse. Leaves petiolate, petioles short, to 5 mm

long, puberulence retrorse to pilose; laminae elliptic-

ovate, 3.5-10 (13) cm long, 1.5-4 (5) cm broad, margins

entire to crenulate, apically acuminate, basally attenuate,

cystoliths dense and prominent on both surfaces, gla-

brous. Inflorescences in terminal and axillary spikes to

7 cm long, ca. 1 cm broad excluding corollas; bracts

imbricate, ovate-elliptic, 6-10 mm long, 4-6 mm broad,

apically obtuse to rounded apiculate, basally obtuse to

rounded, strigose hairs mostly restricted to midrib, veins

very prominent, forming a coarse reticulation, ciliolate;

bracteoles lanceolate, 7-8 mm long, 1 .25-1.5 mm broad,

apically acute to acuminate, apiculate, basally acute, veins

very prominent, pubescence that of the bracts, ciliolate,

peduncles 1-3 cm long, puberulence retrorse. Flowers

with a 5-merous calyx, posterior segment setaceous, ca.

2 mm long, other segments subulate, 5.5-6.5 mm long,

0.5-0.75 mm broad at base, glandular-puberulous, co-

rollas white or bluish white, 9-1 1 mm long, 2 mm broad

at base expanding to 3.5 mm broad at the mouth, spar-

ingly puberulous, upper lip ca. 4.5 mm long, 2.5 mm

broad at base, apically acute, lower lip 5 mm long, 3-

lobed, lobes 2.5 mm long, middle lobe 2.5 mm broad,

lateral lobes 2 mm broad, all apically rounded; stamens

exserted ca. 1 mm beyond the mouth, anther cells su-

perposed, upper cell ca. 0.75 mm long, lower cell ca. 0.5

mm long, both cells basally rounded. Fruits clavate, 7-

8 mm long, 2 mm broad, 1.5 mm thick, puberulous;

seeds reddish brown, flattened, suborbicular, ca. 1.25

mm in diameter, papillate.

Found in pastures, along trails, and in clearings

of premontane moist to premontane rain forest

from 750 to 2000 m elevation in both the Carib-

bean and Pacific watersheds of the Cordilleras Ti-

laran, Central, and Talamanca. Flowering collec-

tions have been made from December through

March.

Justicia valerii is recognized by its stems with

short retrorse hairs, spikes with imbricate bracts

which are ovate-elliptic with a coarse reticulation

pattern, and unequal calyx segments. It is easily

confused with J. candelariae and differs from it in

its stems with short retrorse hairs (vs. pilose stems),

BURGER: FLORA COSTARICENSIS

61

longer laminae (3.5 to 10 cm vs. 2.5 to 6.5 cm),

and glandular-puberulous calyx (vs. ciliolate).

I outeridium S. Watson

REFERENCE— A. Richardson, Revision of Lou-

teridium (Acanthaceae). Tulane Studies in Zool-

ogy and Botany 17: 63-76. 1972.

Shrubs or soft- wooded trees, at least some species

deciduous. Leaves petiolate, laminae ovate, elliptic, ob-

lanceolate or obovate with entire or crenate margins,

cystoliths present and usually prominent in dried spec-

imens. Inflorescences in terminal, cymose, bracteate

panicles, with a pair of cymes at each node, bracts and

bracteoles small. Flowers witih a 3-merous calyx, co-

rollas large, red, light green or yellow, the tube expanding

abruptly just above the ovary, bilabiate, 2 lobes above,

3 lobes below, throat prominently gibbous; stamens 2 or

4, exserted from an arched position in the bud, anthers

2-celled, cells equal. Fruits a sessile capsule with 2 car-

pels; retinacula present, seeds numerous.

This is a genus of the New World tropics, with

nine woody species found in Mexico through Pan-

ama. Collections of L. costaricense have been made

in Darien Province of Panama. This province is

adjacent to Colombia, and it is likely to be found

in that country as well.

Louteridium is not easily confused with other

genera of the Acanthaceae since it is the only genus

with a large, three-merous calyx. This, plus the

large flowers and sessile, many-seeded capsules,

makes it relatively easy to identify.

In Costa Rica this species occurs in moist forests

from 500 to 1000 m elevation. It is found in hab-

itats such as on rotted logs or humus-covered rocks

in streams and along weedy, shaded roadsides. It

is known from Vueltas (Las Vueltas), Alajuela

Province, near Siquirres in Limon Province and

San Vito of Puntarenas Province. Numerous col-

lections have been made in lowland rain forests

of most provinces in Panama. Flowering collec-

tions were made in March and August.

Louteridium costaricense is recognized by its

large three-merous calyx with narrowly elliptic

segments and its large, showy corolla with two

stamens. It is most similar to L. tamaulipense of

Mexico, but differs from it in its persistent bracts

(vs. caducous) and glabrous inflorescence (vs. glan-

dular-puberulous).

Megaskepasma Lindau

Shrub. Leaves petiolate; laminae large, elliptic, cys-

toliths lacking. Inflorescences in loose terminal spikes,

the bracts and bractlets large and conspicuously colored.

Flowers with 5-merous calyx of equal segments; corolla

bilabiate, the upper lip bifid, the lower lip 3-lobed; sta-

mens 2, the anthers 2-celled; stigma subcapitate. Fruit a

clavate capsule, 4-seeded, the seeds borne on hooklike

funicles.

This is a monotypic genus and is endemic to the

Neotropics. It is recognized by its large, showy

bracts and its large leaves, which when dried, are

shown to lack cystoliths.

Louteridium costaricense Radlk. & Donnell Smith,

Hot. Gaz. (Crawfordsville) 37: 422. 1904. Figure

12.

Shrubs to 1.5 m tall; stems tetragonal, inter nodes be-

tween leaf-bearing nodes 1.5-10 cm long, 3-4.5 mm

thick, glabrous. Leaves with petioles to 5 cm long, gla-

brous; laminae elliptic, 2 1-32 cm long, 9.5-15 cm broad,

apically acuminate, basally attenuate, margins entire, both

surfaces glabrous, cystoliths dense and prominent on

both surfaces. Inflorescences in narrow terminal pani-

cles, peduncles to ca. 15 cm long, pedicels 1.5-3 cm long,

glabrous, bracts and bracteoles ovate-lanceolate. 8-12

mm long, 3-5 mm broad, glabrous. Flowers with 3-mer-

ous calyx, segments loosely enclosing the bud, charta-

ceous, narrowly elliptic, apically acute, subequal. 2-4.5

cm long, 1-2 cm broad, cystoliths numerous, venation

conspicuous; corolla pale green or yellow, 3.5 cm long,

glabrous, lobes apically rounded, 8-9 mm long, 6-7 mm

broad, throat prominently gibbous; stamens 2, exserted.

anthers ca. 10 mm long. Fruit clavate, 3-3.5 cm long, 7

mm broad, 7 mm thick, glabrous.

Megaskepasma erythrochlamys Lindau, Bull.

Herb. Boissier 5: 666. 1897. Figure 13.

Shrub to 4 m tall; stems subquadrangular. internodes

between leaf-bearing nodes 2.5-4 cm long, 6-8 mm thick,

puberulous. Leaves with petioles to 4 cm long, petioles

puberulous; laminae elliptic, 13-28 cm long, 3.5-13 cm

broad, apically acuminate with a rounded, apiculate tip,

basally acute, margins entire to crenulate, sparingly stri-

gose on the costa and lateral veins of both surfaces, cys-

toliths none. Inflorescences in lax, clustered spikes to 20

cm long, 4 cm broad excluding corollas, peduncle to 3.5

cm long, brown puberulous, rachis terete, brown puber-

ulous, bracts deep reddish purple drying to reddish brown,

solitary, ovate to elliptic, 3.5-4.7 cm long, 1.5-2 cm

broad, apically acute, basally acute, sparingly strigulose

on both surfaces, mostly restricted to the costa, margins

entire; bracteoles elliptic to 20 mm long, 5 mm broad,

apically acute, basally attenuate, sparingly strigulose.

Flowers with 5-merous calyx, calyx segments equal, lan-

ceolate, ca. 10 mm long, 2 mm broad basally, densely

strigose; corolla white, tube cylindrical, 20-25 mm long,

2-3 mm broad basally, pilosulous, the upper lip lanceo-

62

FIELDIANA: BOTANY

late to 35 mm long, 5 mm broad basally, apically 2-lobed,

the lobes subulate, 1.5 mm long, 0.5 mm broad basally,

the lower lip elliptic, 35 mm long, 8 mm broad, pilo-

sulous outside, apically 3-lobed, the lobes lanceolate, 7

mm long, 2 mm broad basally; stamens exserted ca. 3

cm beyond throat of the corolla, the filaments glabrous,

the anthers 4.5 mm long, 1.5 mm broad, basally mu-

cronate; ovary 2 mm long, glabrous. Fruits clavate, 30-

35 mm long, 8-10 mm broad, 6-7 mm thick, apically

attenuate and mucronate, glabrous; seeds 4, sub-

orbiculate, flattened, ca. 7 mm in diameter, glabrous.

A native of Venezuela, this plant is cultivated

throughout much of the Neotropics.

Megaskepasma erythrochlamys is recognized by

its shrubby habit, its large leaves which lack cys-

toliths, and its flowers with large (3.5-4.7 cm long)

reddish purple bracts, and its large (50-60 mm

long) white, bilabiate corollas with two exserted

stamens having two-celled anthers with equal cells.

It most closely resembles the genus Justicia, but

differs in its leaves, which lack cystoliths, and its

much larger bracts.

Mendoncia Veil.

Tall, suffrutescent vines. Leaves opposite, petiolate,

laminae pinnately veined, margins mostly entire, cys-

toliths lacking. Flowers axillary, 1 to several or occa-

sionally many in each axil, each flower pedicellate and

subtended by 2 flat or keeled bracts, connate prior to

anthesis and variably so at anthesis; calyx reduced to an

annular, often membranous structure, usually glabrous;

corolla 5-merous, the tube funnelform or tubular, usually

expanded at the base, the lobes equal or nearly so, re-

flexed or spreading, the throat often oblique; stamens 4,

didynamous, the filaments short, the anthers linear-lan-

ceolate, 2-celled, usually flattened, apically acute, glan-

dular-puberulous, basally lobed, the lobes more or less

unequal; ovary oblique. Fruit a compressed drupe, often

apically oblique, the endocarp pulpy; seeds 1 or 2.

This is a genus of approximately 60 species from

Central and tropical South America, tropical Af-

rica, and Madagascar. In Mendoncia gracilis, M.

lindavii, and M. littoralis (of Colombia and Pan-

ama), the bracts are completely joined before an-

thesis, enclosing the flower in a clear, colorless

liquid until it opens. While this has not been re-

ported for other species of Mendoncia, the connate

habit of the bracts before anthesis suggests that

other species may have fluid-enclosed flowers as

well.

Mendoncia is recognized by its viny habit, its

paired, connate bracts partially enclosing five-

merous flowers with four didynamous stamens,

and its drupaceous fruits.

Key to Species of Mendoncia

la. Corolla 1 .5 cm long; flowers in axillary umbels M. brenesii

Ib. Corolla 2.5 cm long or more; flowers axillary, borne singly or in 2's or 3's 2a

2a. Young stems narrowly winged; hairs on upper leaf surface not arising from stellate bases

M. retusa

2b. Young stems not winged; hairs on upper leaf surface arising from stellate bases 3a

3a. Corollas less than 3.5 cm long; bracts less than 2 cm long 4a

3b. Corollas more than 3.5 cm long; bracts more than 2.5 cm long 5a

4a. Petioles 2-4 cm long; bracts obovate to elliptic and basally subcordate M. costaricana

4b. Petioles 1-2 cm long; bracts elliptic to ovate and basally rounded M. gracilis

5a. Bracts oblong-lanceolate; widest below the middle; hairs on stem evenly distributed, somewhat

appressed M. landavii

5b. Bracts oblong to oblong-elliptic; widest at the middle; hairs on stem mostly at the angles and

spreading . M. tonduzii

Mendoncia brenesii Standley & Leonard, Publ.

Field Mus. Nat. Hist., Bot. Ser. 18(4): 1239.

1938. Figure 12.

Suffrutescent vine; stems scandent, subquadrangular,

grooved; internodes between leaf-bearing nodes 4.5-12

cm long, 1.5-3 mm thick, glabrous to sparingly and mi-

nutely strigose. Leaves with petioles to 2.6 cm long, pet-

ioles sparingly strigose; laminae ovate-elliptic, 6-11.5

cm long, 3-7.5 cm broad, apically acuminate with a

mucro 1-2 mm long, basally rounded, margins entire,

glabrous above and below except for sparingly minutely

strigose on the costa. Inflorescences in axillary umbels

of 5-10 flowers radiating from short, flat, rounded, ax-

illary spurs, pedicels ca. 1 cm long, sparingly strigose;

bracts ovate, 8-12 mm long, 5-7 mm broad, apically

BURGER: FLORA COSTARICENSIS

63

acuminate, hasally rounded, sparingly strigose, bracts

frequently fused to ca. V* their length. Flowers with a

membranous calyx; corolla white, ca. 1 5 mm long, gla-

brous, the tube ca. 8 mm long, 3 mm broad, slightly

expanded basally, the lobes obcordate, ca. 7 mm broad;

stamens included, the anthers 3 mm long, apically cal-

carate, basally obtuse.

Few collections of this species have been made,

but these have been in shady areas of cloud forests

of the Cordillera Central, Alajuela Province, at

elevations of 1000 to 1500 m. It has also been

collected in adjacent Panama at 1 500 to 2000 m.

Flowering collections have been made from June

through August.

Mendoncia brenesii is most easily recognized by

its axillary umbels of small, white flowers. It is not

easily confused with other Mendoncia species if

its axillary umbels are present.

Mendoncia costaricana Oerst., Vidensk. Meddel.

Dansk Naturhist. Foren. Kjobenhavn 1854: 113.

1855. Figure 12.

Herbaceous vine; stems subquadrangular, grooved; in-

ter nodes between leaf-bearing nodes 5.5-14 cm long,

1.5-2.5 mm thick, strigose. Leaves with petioles to 3.8

cm long, petioles strigose; laminae elliptic to elliptic-

ovate, 7.7-13.5 cm long, 3-6.7 cm broad, apically

abruptly acuminate and sometimes falcate, the apiculate

tip to ca. 2 mm long, basally acute to near rounded,

margins entire, sparingly scabrous above with hairs aris-

ing from stellate bases, sparingly strigose beneath. Flow-

ers axillary, borne singly or in 2's or 3's, pedicels ca. 2

mm long, strigose; bracts obovate to elliptic, 15-18 mm

long, 8-10 mm broad, apically obtuse and apiculate,

basally subcordate, strigose outside, glabrous within; co-

rollas white with dark spots at the throat, to 3 cm long,

8 mm broad at the throat, 4-5 mm broad, ovary gla-

brous. Fruit flattened, ovoid with double margins, ca.

20 mm long, 1 3 mm broad, 4 mm thick, puberulous.

The seven collections of this species in Costa

Rica indicate that it is wide ranging in rain forests

at elevations of 500 to 1000 m in the Pacific wa-

tershed area of San Jose Province and the Cor-

dillera Central portions of Alajuela, Cartage, and

Puntarenas provinces. It is also found in adjacent

Chiriqui Province of Panama. Flowering collec-

tions have been made in July, August, and Sep-

tember.

Mendoncia costaricana is recognized by its

comparatively short (3 cm long), white corollas

borne singly or in pairs of axillary flowers, its stri-

gose stems, and its relatively long petioles (2 to 4

cm long). It may be confused with M. gracilis, but

can be distinguished from it by its longer petioles

(2 to 4 cm long vs. 1 to 2 cm) and its obovate to

elliptic bracts, which are basally subcordate.

Mendoncia gracilis Turrill, Bull. Misc. Inform. 418.

1919.

Suffrutescent vine; young stems quadrangular, strigose

to sericeous, hairs appressed upward; internodes between

leaf-bearing nodes, 4.5-10.5 cm long, 1.2-2.5 mm thick.

Leaves with strigose to sericeous petioles to 2 cm long,

mostly ca. 1 .5 cm long; laminae elliptic to elliptic ovate,

8.5-12.5 cm long, 4-7 cm broad, apically acuminate,

often with a mucro to 2 mm long, basally rounded to

obtuse, margins entire, the veins prominent, usually 4

pairs, sparingly strigose above, hairs emerging from stel-

late bases, strigose below. Inflorescences axillary, the

pedicels borne singly to 3 or 4, 1.2-1.5 cm long, strigose

to sericeous; bracts elliptic to ovate, entire, 15-19 mm

long, 6-9 mm broad, apiculate, basally rounded, strigose

to sericeous outside. Flowers with the calyx reduced to

an entire membranous cup ca. 1 mm deep, glabrous;

corolla white, funnelform, ca. 3 cm long, basally 3 mm

broad expanding to 6 mm broad at the throat, the throat

streaked with reddish brown, oblique, the lobes 5-6 mm

long, 4-5 mm broad, shallowly emarginate, the tube

glandular puberulous inside below the stamens, remain-

der of corolla glabrous; stamens included. Fruits ovoid,

ca. 1.7 cm long, purple-black and fleshy at maturity.

This species, originally described from along

trails and openings in the rain forests of the Pacific

slope and intermountain areas of Colombia, has

also been collected in similar habitats in central

Panama. One collection, Durkee 83-4 (F, GRI), has

been made in the Golfo Dulce area, Puntarenas

Province of Costa Rica, in mature rain forest along

the new road to Jimenez from the Interamerican

Highway near the Rio Olla. The flowering collec-

tion was made in January.

Mendoncia gracilis can be recognized by its white

corollas about 3 cm long borne singly to three or

four in leaf axils, elliptic bracts ( 1 5 to 1 9 mm long),

basally rounded with upwardly appressed seri-

ceous hairs; young stems and leaves also have up-

wardly appressed hairs, and petioles are 1 to 2 cm

long. It may be confused with M. costaricana,

which differs in its longer petioles (2 to 4 cm long)

and its obovate to elliptic bracts, which are basally

subcordate.

Mendoncia lindavii Rusby, Mem. Torrey Bot. Club

4: 241. 1895. Figure 12.

Suffrutescent vine; younger stems quadrangular, in-

ternodes between leaf-bearing nodes, 3.5-17 cm long,

1.5-3 mm thick, wooly hairs rust-colored. Leaves peti-

olate, petioles to 3 cm long, wooly with rust-colored

hairs; laminae elliptic, 5.8-11.5 cm long, 3.3-6.4 cm

64

FIELDIANA: BOTANY

broad, apically acuminate, hasally obtuse to rounded,

margins entire, sericeous above with hairs rising from

stellate bases, sericeous beneath, but hairs more dense

and prostrate. Flowers axillary, pedicellate, borne singly

or in pairs, pedicels to 3 cm long, pubescence that of the

petiole; mature bracts oblong-lanceolate, subfalcate, 3.2-

4 cm long, 0.8-1.2 cm broad, apically attenuate, basally

rounded, wooly with rust-colored hairs; corolla red,

pink, or white, ftmnelform, 4-5 cm long, 6 mm broad

basally expanding to 8 mm broad at the throat, glabrous,

the 5 equal lobes to 6 mm long, 4.5 mm broad, apically

retuse to rounded; stamens attached at about the mid-

point of the corolla tube, anthers 10-13 mm long, 1.5

mm broad, filaments 2 mm long; ovary velutinous, ca.

4 mm long, 2.5 mm broad, style to 5 cm long, stigma

forked. Fruits flattened, obovate, to 20 mm long, 1 1 mm

broad, 8 mm thick, apically oblique and terminated by

a portion of the persistent style, dark brown (dull purple

when fresh) and densely puberulous.

Ranging from Belize and Guatemala to Bolivia,

this species has been collected in rain forest areas

of Costa Rica at elevations of 500 to 1000 m on

the Pacific Watershed only in Puntarenas Province

near Panama and San Jose Province in the vicinity

of General Viejo (El General). Flowering collec-

tions have been made in June and January.

Mendoncia lindavii is recognized by its large red,

pink, or white corollas (4 to 5 cm long), its large

(3.2 to 4 cm long), oblong-lanceolate, subfalcate

bracts with conspicuous, reddish brown wooly

hairs, and its large (to 20 mm long) dark brown

or purplish, obovate fruits. It may be confused

with M. retusa, which also has large flowers, but

can be distinguished from it by its wooly stems

(vs. strigose), its nonwinged stems (vs. narrowly

winged), and its wooly, oblong-lanceolate bracts

(vs. mostly glabrous, oblong).

Mendoncia retusa Turrill, Bull. Misc. Inform. 423.

1919. Figure 12.

Suffrutescent vines with narrowly winged, quadran-

gular stems; internodes between leaf-bearing nodes 4-

14 cm long, 1-2.5 mm thick, sparsely strigose to gla-

brous. Leaves petiolate, petioles to 5.2 cm long, sparsely

strigose to puberulous; laminae elliptic, 7-15 cm long,

2.5-7.5 cm broad, apically acuminate and mucronulate,

basally acute, obtuse or rounded, margins undulate,

sparsely strigose on costa beneath. Flowers axillary, ped-

icellate, borne singly, pedicels 1.75-3 cm long, sparingly

strigose to glabrous; bracts green, elliptic to elliptic-ob-

long, 1.5-3.0 cm long, 1-2 cm broad, apically retuse or

rounded, mucronate, basally rounded, mostly glabrous;

corolla white with a purplish throat, 4-5 cm long, gla-

brous, lobes equal, 5-6 mm long, apically obtuse. Fruit

irregularly obovate to 19 mm long, 9 mm broad, 8 mm

thick, puberulous.

Found in lowland rain forests ranging from

Mexico (Chiapas) through Panama, this species is

one of the most commonly found in Central Amer-

ica, although, again, few collections have been made

in Costa Rica. However, these collections are from

both the Caribbean and Pacific areas at elevations

of 15 to 100 m. Flowering collections have been

made in June and October.

Mendoncia retusa is recognized by its narrowly

winged, quadrangular stems, its large (4-5 cm long),

white corollas with purplish throats, and its elliptic

to elliptic-oblong bracts, which are apically retuse

or rounded and mucronate. The bracts are some-

what variable, and in those specimens with more

oblong bracts, there may be confusion with M.

lindavii, which differs in its rust-colored, wooly

hairs, its lack of winged stems, and its red, pink,

or white corollas.

Mendoncia tonduzii Turrill, Bull. Misc. Inform.

413. 1919. Figure 12.

Suftrutescent vines with quadrangular stems; inter-

nodes between leaf-bearing nodes 9-12 cm long, 2-3 mm

thick, densely pilose, hairs tawny and mostly restricted

to the angles. Leaves petiolate, petioles to 2 cm long,

densely pilose tawny hairs; laminae elliptic to elliptic-

ovate, 7-13 cm long, 5-8.3 cm broad, apically acumi-

nate, often with an apicule to 2 mm long, basally acute

to rounded, margins entire, pilose hairs with stellate bas-

es above, more densely pilose below. Flowers solitary or

in pairs in the axils of the upper leaves, pedicellate, ped-

icels 2-4 cm long, pilose; bracts oblong to oblong-elliptic;

ca. 4 cm long, 1.5 cm broad, apically rounded and mu-

cronate, basally rounded, densely pilose, hairs tawny;

corolla white, ca. 5 cm long, 8 mm broad at the oblique

throat, the lobes equal, emarginate, spreading, ca. 8 mm

long, 5-6 mm broad, glabrous. Fruit purplish black at

maturity, ca. 2 cm long, 1 cm broad, 6 mm thick, densely

puberulous.

This uncommon species has been collected in

forests at 500 to 1000 m elevation of the Tilaran

area, Guanacaste Province, Brenes 12675 (F), and

Tucurrique, Cartage Province, Tonduz 12946 (us,

holotype). Three collections have also been made

in similar habitats in Code and Colon provinces

of Panama.

Mendoncia tonduzii is recognized by its dense,

tawny pilose hairs, its oblong to oblong-ovate

bracts, 4 cm long, apically rounded and mucro-

nate, and laminae with pilose hairs on stellate bas-

es. This distinguishes it from the more common

M. lindavii, which it most closely resembles.

Nelsonia R. Br.

Herbs, stems softly villous. Leaves sessile; laminae

with entire margins. Inflorescences of dense, terminal

BURGER: FLORA COSTARICENSIS

65

and axillary spikes; bracts imbricate, 1 per flower. Flow-

ers with 4-merous calyx; corolla bilabiate; stamens 2,

anthers 2-celled, the cells parallel, each cell globose; stig-

ma 2-lobed. Fruits a 2-chambered capsule, capsules ob-

long; seeds borne on papilliform funicles.

There is only one species described for this ge-

nus, which was first collected in Java. The genus

was named for David Nelson, a gardner who ac-

companied Cook on his last voyage.

Nelsonia canescens (Lam.) Spreng., Syst. veg. ed.

16. 1:42. 1825. Figure 13.

Herbs, decumbent to ascending with narrow, branch-

ing stems to 40 cm long; internodes between leaf-bearing

nodes 1-5 cm long, 1-2 mm thick, villous to wooly.

Leaves sessile; laminae ovate, 1-8 cm long, 0.5-3 cm

broad, apically acute to obtuse, basally acute to obtuse,

margins entire, cystoliths lacking, villous above and be-

neath. Inflorescences in dense spikes to 6.5 cm long, 5.5

mm broad excluding corollas, pedunculate or sessile,

mostly axillary, flowers sessile to pedicels ca. 1 mm long;

bracts imbricate, 1 per flower, ovate, 4-6 mm long, 4-

5 mm broad, apically acuminate, velutinous. Flowers

with 4-merous calyx, segments ca. 4 mm long, unequal,

posterior segment ovate, 2.5 mm broad, anterior seg-

ment lanceolate, 1.5 mm broad, lateral segments lan-

ceolate, 1 mm broad; corolla deep purple to blue, bila-

biate, the tube ca. 3 mm long, upper lip 2-lobed, ca. 1

mm long, lower lip 3-lobed ca. 1 mm long. Fruit sessile,

ca. 5 mm long, ca. 10 seeds per locule.

Found in disturbed habitats, especially in moist,

sandy areas along roadsides, trails, and streambeds,

this species occurs in Mexico, Central America,

northern South America, tropical Africa, India to

Indo-China, Malay Peninsula, and Australia. Col-

lected in Costa Rica at elevations of 150 to 1500

m in Alejuela, Guanacaste, and San Jose prov-

inces. Flowering collections have been made in

February through May.

Nelsonia canescens is recognized by its decum-

bent stems with numerous, compact spikes, ve-

lutinous, imbricate bracts subtending minute flow-

ers with bilabiate bluish corollas, and two stamens

with two equal-celled anthers. Its capsules are ses-

sile, and its seeds lack the hooklike funicles which

serve to eject the seeds of most Acanthaceae. Cer-

tain specimens may resemble Elytraria imbricata,

which has larger leaves (6 to 20 cm vs. 1 to 8 cm

long) and stiff, awned bracts (vs. soft, awnless

bracts).

Odontonema Nees

REFERENCE— V. M. Baum and J. L. Reveal, A

new proposal to conserve 8037 Odontonema

(Acanthaceae). Taxon 31: 757-759. 1982.

Herbs or shrubs. Leaves sessile to petiolate; laminae

often large with cystoliths visible on both surfaces of

dried specimens. Inflorescences in terminal panicles,

thyrses, or verticillate inflorescences, bracts inconspic-

uous. Flowers with 5-merous, equally segmented calyx;

corolla bilabiate, the upper lip 2-lobed, the lower lip

3-lobed; stamens 2, exserted or extending to the mouth,

the anthers 2-celled with the cells parallel and basally

blunt, staminodes 2. Fruits in clavate capsules, seeds 4

on hooklike funicles.

This is a genus of 29 species, all endemic to the

Neotropics.

Odontonema tub! for me (Bertol.) Kuntze, Re vis.

gen. pi. 2: 493. 1891. Justicia tubiformis Bertol.,

Novi Comment. Acad. Sci. Inst. Bononiensis 4:

405. 1840. ThyrsacanthusflagellumOerst.,Vi-

densk. Meddel. Dansk Naturhist. Foren. Kj0-

benhavn 1854: 146. 1855. Odontonema flagel-

lum (Oerst.) Kuntze, Revis. gen. pi. 2: 494. 1891.

Thyrsacanthus strictus Nees in DC., Prodr. 11:

324. 1947. Odontonema strictum (Nees) Kuntze,

Revis. gen. pi. 2: 494. 1891. Figure 13.

Suffrutescent herbs to shrubs 2.5 m tall; internodes

between leaf-bearing nodes 1.5-9 cm long, 1 .5-3.25 mm

thick, puberulous. Leaves sessile to petiolate, petioles to

3 cm long, puberulous to subglabrous; laminae elliptic

to broadly elliptic-ovate, 1 1-36 cm long, 3.5-1 1 cm broad,

apically acuminate, basally attenuate, margins crenate

to entire (often on the same plant), cystoliths more vis-

ible on upper surface, obscure beneath, glabrous. Inflo-

rescences verticellate to 35 cm long, 1.5 cm broad ex-

cluding corollas or fruits, peduncles 1.5-7 cm long,

puberulous, rachis puberulous to tomentose, flowers

pedicellate with 1 to 10 pedicels per cluster, pedicels

puberulous; bracts and bracteoles inconspicuous, subu-

late, 3-10 mm long, 1-2 mm broad, puberulous. Flowers

with 5-merous calyx, calyx lobes equal, subulate, 2-4

mm long, 0.6-1 mm broad, minutely puberulous; corolla

pink or red, funnelform, 15-30 mm long, 4-5.5 mm

broad at the throat, 1-3 mm broad at the base, glabrous,

upper lip 7-8 mm long, 4.5-5 mm broad, bilobed, lobes

apically acute, 3.5-4 mm long, lower lip 7-8 mm broad,

lobes elliptic, apically rounded, 3-4 mm broad, all lobes

with glandular hairs inside; stamens exserted, anthers ca.

2.5 mm long, filaments glabrous, staminodes 1-3.5 mm

long. Fruits clavate, 2-3 cm long, 4 mm broad, 3-4 mm

thick, glabrous, seeds 4, ovate with an oblique base,

flattened, 5-6 mm long, 3.5-4.5 mm broad, 1 mm thick,

surface densely rugose.

This species is found in light gaps, clearings, and

along streams and trails in rain forests of lower

elevations to rain forest transition at higher ele-

vations of Mexico, Central America, and Panama.

In Costa Rica it has been collected in both the

Atlantic and Pacific watersheds from all provinces

except Guanacaste at elevations of 50 to 1200 m.

66

FIELDIANA: BOTANY

Flowering collections have been made in July and

October through April. This is a wide-ranging

species which is subject to much variation in leaf,

flower, and fruit size, as well as pubescence of the

rachis and number of flowers per cymule. Many

of the early species were erected on these bases,

but additional collections show intergradation in

all of these characters. The only character that

appears unique to Odontonema Jlagellum (Oerst.)

Kuntze is the narrowed, flagellate tip of the inflo-

rescence which I believe is an early flowering stage

of O. tubiforme.

Odontonema tubiforme is recognized by its ver-

ticillate inflorescences with near-equal, funnel-

form corollas that are pink or red with two exserted

stamens with two-celled anthers and two short

staminodes. It is most frequently confused with

Razisea species, but can be distinguished from

them by its staminodes, two-celled anthers, and

more broadly funnelform corollas.

Poikilacanthus Lindau

Herbs or shrubs. Leaves of a pair unequal, petiolate.

Flowers borne in small terminal or axillary, bracted spikes

or heads. Flowers with 5-merous calyx; corolla narrow,

bilabiate; stamens 2, anthers 2-cclled. the cells unequally

attached, the cells rounded or ending in a short spur;

pollen elliptic, polyporate. Fruits a clavate, 2-chambered

capsule with 4 seeds, each borne on hooklike retinacula.

subulate, 7-9 mm long, 1.3-1.5 mm broad, ciliolate to-

ward apex; corolla lavender or purplish, very narrowly

funnelform, 6.5-7.8 cm long, 4-5 mm broad at the throat,

1.5 mm broad basally, glandular-puberulous, upper lip

30-35 mm long, 7-9 mm broad, apical K minutely

2-lobed, lower lip 22-27 mm long, 9-12 mm broad at

ca. midpoint, apically 3-lobed, lobes ca. 2 mm long, 3

mm broad; stamens exserted to ca. the tip of the upper

lip or just beyond, anther cells ca. 2.25 mm long, lobes

obtuse. Fruits clavate, 12-15 mm long, 3.5-5 mm broad,

3.1-3.5 mm thick, sparingly puberulous.

Found in gaps or clearings, along trails and

streams in cloud forest areas of Mexico (Chiapas),

Honduras, Guatemala, Nicaragua, and Costa Rica.

Collected in Costa Rica in cloud forest areas of

the Cordilleras Tilaran, Central, and Talamanca

at elevations of 1000 to 1750 m, mostly on the

Pacific watershed, but occasionally on the Carib-

bean as well. Flowering collections have been made

in February, March, and September through No-

vember.

Poikilacanthus macranthus is recognized by its

unequal leaf pairs, its lavender, narrowly funnel-

form bilabiate corolla (6.5 to 7.8 cm long), two

exserted stamens with widely superposed anther

cells, and elliptic, polyporate pollen. It is most

easily confused with species of Justicia, but can

usually be distinguished from these in its unequal

leaf pairs (excluding Justicia trichotoma), its large,

narrowly funnelform corollas, and its polyporate

pollen (vs. two-, rarely three-, or four-, porate).

This is a Neotropical genus of about 12 species

found in Central America and tropical South

America. It is similar to the genus Justicia, but

differs from it in its unequal leaf pairs and its

pollen morphology.

Poikilacanthus macranthus Lindau, Bull. Herb.

Boissier 3: 481. 1895. Figure 13.

Shrubs to 3 m tall; internodes between leaf-bearing

nodes 2-6.5 cm long, 1-2.25 mm thick, quadrangular,

puberulence of appressed hairs varying from a few hairs

restricted to the angles to dense. Leaves of a pair unequal,

petioles 0.5-2.6 cm long, pubescence that of the stem;

laminae ovate to ovate-elliptic, 3-7 (8.5) cm long 1.5-

3.5 (4) cm broad, apically acuminate, basally obtuse to

rounded and often oblique, margins entire, appressed

hairs restricted to costa and veins on both surfaces to

pilose, cystoliths dense and prominent on both surfaces.

Inflorescences in terminal, sessile, headlike clusters of

2-4 flowers; bracts oblong to spatulate, 6-9 mm long,

1.5-3 mm broad, ciliate, bracteoles similar, but smaller.

Flowers with 5-merous calyx, calyx segments equal,

Pseuderanthemum Radlk.

Herbs or small shrubs. Leaves commonly petiolate;

laminae elliptic, ovate, or lanceolate, cystoliths generally

visible on both surfaces of dried specimens. Inflores-

cences in terminal or axillary racemes, spikes or panicles;

1 bract and 2 bracteoles subtending each flower or flower

cluster, bracts and bracteoles inconspicuous. Flowers with

4 or 5 subulate calyx segments; corolla white, blue, pur-

plish or pink, salverform, the tube narrow, the limb

spreading, the 5 lobes subequal; stamens 2, included or

exserted, inserted at or above the middle of the corolla

tube; anther 2-celled, the cells basally obtuse or acute;

staminodes 2, minute. Fruits in clavate capsules, seeds

2 or 4 on hooklike retinacula.

This is a genus of about 1 20 species from the

New and Old World tropics. The flowers of this

genus resemble those of the genus Chamaeran-

themum which differs in its four stamens that are

included, with the posterior pair either one-celled

or sterile.

BURGER: FLORA COSTARICENSIS

67

Key to Species of Pseuderanthemum

la. Inflorescence a cymose panicle (thyrse); rachis of inflorescence glabrous P. atropurpureum

Ib. Inflorescence a raceme, spike, or spicate panicle; rachis glandular-puberulous or puberulous . . 2a

2a. Petioles conspicuously winged, laminae bases long-attenuate; stamens exserted . . . P. cuspidatum

2b. Petioles not winged, laminae bases acute or attenuate, but not long-attenuate; stamens included 3a

3a. Flowers in opposite fascicles of 2-3 or more; laminae mostly more than 9 cm long . . . . P. pittieri

3b. Flowers borne singly and opposite, or if in pairs, alternate; laminae mostly less than 9 cm long . . . 4a

4a. Flowers borne singly and opposite; capsules 16-19 mm long, puberulous P. praecox

4b. Flowers borne singly or in pairs, mostly alternate; capsules 1 1-15 mm long, glabrous . . . . P. standleyi

Pseuderanthemum atropurpureum (Bull) Bailey,

Gentes Herb. 1: 130. 1923. Eranthemum atro-

purpureum Bull. Card. Chron. 1 : 6 19. 1 875. Fig-

ure 13.

Shrubs or small trees to 4 m tall; internodes between

leaf-bearing nodes 2.5-5 cm long, 2-4 mm thick, te-

trangular. glabrous. Leaves petiolate, petioles 0.5-1 .8 cm

long, glabrous except for a tuft of hair at the stipular

ring; laminae elliptic to ovate, 4.5-14.5 cm long, 4.5-

6.5 cm broad, apically acute, some apiculate, basally

attenuate, margins entire, greenish beneath and dark pur-

ple above, glabrous on both surfaces, cystoliths dense

and near punctiform on both surfaces. Inflorescences in

terminal and axillary cymose panicles with the flowers

in subsessile fascicles of mostly 3 flowers; pedicels, pe-

duncles and rachises glabrous, pedicels to 5 mm long,

peduncles to 7 cm long, rachises to 1 3 cm long, bracts

elongate, lower bracts ca. 5-7 mm long, 2-3 mm broad,

upper bracts progressively smaller, all ciliolate. Flowers

with 5-merous calyx, the lobes equal, subulate, 2-3 mm

long, 0.75-1 mm broad, puberulous; corolla white with

magenta spots in the throat, the tube cylindrical, 10-12

mm long, 2-2.5 mm broad basally, the limb 20-22 mm

broad, the lobes elliptic, 8-10 mm long, 5-8 mm broad,

apically rounded, ciliolate; stamens exserted, filaments

attached just below the mouth, glabrous anther cells

equally attached, ca. 1.5 mm long, basally acute, stam-

inodes 1.5-2 mm long. Fruits not observed.

This species, frequently cultivated in tropical

America, is believed to be a native of Polynesia.

It sometimes escapes from cultivation. Flowering

collections have been made in January and No-

vember. Pseuderanthemum atropurpureum is rec-

ognized by its cymose panicles and purplish leaves,

which are frequently variegated with green, white,

and pink. These features make it easily distin-

guishable from other species of the genus.

Pseuderanthemum cuspidatum (Nees) Radlk., Sit-

zungsber. Math.-Phys. Cl. Konigl. Bayer. Akad.

Wiss. Munchen 13: 286. 1883. Eranthemum

cuspidatum Nees in Bentham, PI. hartw. 148.

1839. Figure 13.

Herbs to 60 cm tall; stems erect; internodes between

leaf-bearing nodes 1.3-7 cm long, 0.5-3.5 mm thick,

quadrangular, puberulous along 2 lines. Leaves petiolate,

petioles subsessile to ca. 3.5 cm long, glabrous to spar-

ingly puberulous along 2 lines; laminae ovate, 8-19 cm

long, 3.2-8.2 cm broad, apically acuminate, basally long-

attenuate, margins entire, glabrous to very sparingly sca-

brous on both surfaces, cystoliths dense and prominent

on both surfaces. Inflorescences in lax terminal and ax-

illary spikes frequently branched from the base, to 16

cm long, 4 mm broad excluding corollas, the rachis glan-

dular-puberulous; lowermost bracts subtending the in-

florescence, suborbicular, to 2 cm long, 1.5 cm broad,

floral bracts subulate, 2-3 mm long, 0.75-1 mm broad,

ciliolate; bracteoles similar, though smaller; flowers ses-

sile, paired or in pseudowhorls of up to 5 flowers. Flowers

with a 5-merous calyx, calyx lobes equal, subulate, 2.5-

3.5 mm long, 0.5-0.75 mm broad, glabrous to sparingly

puberulous plus occasional glandular hairs; corolla pur-

plish, glabrous, tube cylindrical 1.5-2.8 cm long, 0.8-1

mm broad, lobes of the limb elliptic, 8.5-9 mm long,

4.5-5 mm broad, apically obtuse; stamens barely exsert-

ed, filament ca. 2.5 mm long, staminodes 0.5 mm long,

anther cells 1.25 mm long. Fruits clavate, 11-15 mm

long, 2-2.5 mm broad, 1.5-2 mm thick, glabrous, seeds

4, ovate, 2.5-3 mm long, 2-2.5 mm broad, rugose.

Found in shady, moist, forested areas from

southern Mexico to northwestern South America,

this species is wide ranging and not uncommon as

suggested by the relatively numerous collections.

In Costa Rica it is found in both major watersheds

from elevations of about 700 to 1575 m. It has

not been collected in Guanacaste, Heredia, or Li-

mon provinces. Flowering collections have been

made in May and August through December.

Pseuderanthemum cuspidatum is recognized by

its often-branched, terminal and axillary spikes

with lax flowers in pairs or pseudowhorls, glabrous

to subglabrous five-merous calyx, and its long,

conspicuously winged petioles (or long-attenuate

laminae). It is most easily confused with P. pittieri,

which differs in its more pronounced glandular-

puberulous calyx, its elliptic laminae with more

abruptly attenuate bases, and its shorter corolla

tubes (7.5 to 1 1 mm long vs. 15 to 28 mm long)

with stamens included.

68

FIELDIANA: BOTANY

Pseuderanthemum pittieri Leonard, Publ. Field

Mus. Nat. Hist., Hot. Ser. 18: 1245. 1938. Figure

14.

Herbs or small shrubs to 80 cm tall, stems erect; in-

ter nodes between leaf-bearing nodes 1-7 cm long, 2-4

mm thick, subquadrangular, puberulous. Leaves with

short petioles to 6 mm long, glabrous to sparingly puber-

ulous; laminae elliptic, 8.5-19 cm long, 3-8.5 cm broad,

apically acuminate, basally attenuate or acute, margins

undulate, cystoliths moderately dense and visible on both

surfaces, glabrous above, purplish puberulence on costa

and main veins beneath. Inflorescences in terminal and

axillary spikelike racemes, frequently branched; flowers

opposite, frequently in fascicles of 2-3, pedicels short,

ca. 0.5 mm long, puberulous-glandular; bracts subulate,

2.5-3 mm long, 1-1.5 broad basally, puberulous plus

occasional glandular hairs. Flowers with a 5-merous ca-

lyx, calyx lobes equal, long-subulate, 3-4 mm long, 0.4-

0.5 mm broad basally, glandular- puberulous: corolla

white, pink, or lavender, the tube subcylindrical, 7.5-1 1

mm long, 1-1.5 mm broad basally and up to ca. mid-

point where the stamens are attached and then expanding

to 1.5-2 mm broad at the limb, tube glandular-puber-

ulous, lobes of the limb 10-11 mm long, 4-5 mm broad,

apically obtuse; stamens included, attached at midpoint

of corolla tube, filament ca. 2 mm long, staminodes 0.5

mm long, anther cells 1 .25 mm long. Fruits clavate, ca.

1 8 mm long, 2.8 mm broad, 2.2 mm thick, apically acute,

puberulous; seeds 4, oval, flattened, ca. 3 mm long, 2.5

mm broad, tubercled.

Found growing in shade of moist lowland forest

areas of primary and secondary forests in the Golfo

Dulce area and the Osa Peninsula at elevations of

50 to 200 m. Flowering collections have been made

in December and January.

Pseuderanthemum pittieri is recognized by its

comparatively large, elliptic leaves with short-at-

tenuate lamina bases, spikelike racemes, glandu-

lar-puberulous calyx lobes and corollas, and short,

subcylindric corolla tubes (7.5-1 1 mm) with in-

cluded stamens. It is most easily confused with the

more widespread P. cuspidatum, which differs in

its conspicuously winged petioles (long-attenuate

laminae), true spikes, glabrous to subglabrous ca-

lyx lobes and corollas, and longer cylindric corolla

tubes (15-28 mm) with exserted stamens.

Pseuderanthemum praecox (Bentham) Leonard, J.

Wash. Acad. Sci. 31: 99. 1941. Eranthemum

praecox Benth., PI. hartw. 291. 1848. Figure 14.

Herbs to 30 cm tall; Intel-nodes between leaf-bearing

nodes 1.8-5 cm long, 1-1.5 mm thick, quadrangular,

puberulous mostly along 2 lines. Leaves with petioles 4-

10 mm long, glabrous to sparingly puberulous; laminae

elliptic to elliptic-ovate, 1 .5-8 cm long, 0.8-2.5 cm broad,

apically acuminate, basally attenuate, margins undulate,

cystoliths dense and visible on both surfaces, sparingly

strigose above, strigose on costa and main veins below.

Inflorescences in short terminal and axillary spikes 1-

6.5 cm long, flowers opposite, borne singly, bracts subu-

late, 1.5-3 mm long, 0.5-0.75 mm broad, sparingly pu-

berulous, bracteoles similar though smaller; rachises

glandular-puberulous. Flowers with a 5-merous calyx,

calyx lobes long-subulate, 2.5-3.5 mm long at early an-

thesis, 5-6 mm long in fruit, 0.5-0.75 mm broad, glan-

dular-puberulous to mostly puberulous; corolla laven-

der, pink or bright rose, glabrous outside, the tube

cylindrical, 16-19 mm long, ca. 1.5 mm broad, lobes

elliptic, 8-12 mm long, 3.5-5 mm broad, apically acute;

stamens included, filaments 1.75-2 mm long, stami-

nodes ca. 0.5 mm long, anthers 1.5-2 mm long. Fruits

clavate, 15-18 mm long, ca. 3 mm broad, 4 mm thick,

puberulous plus occasional glandular hairs on some; seeds

suborbicular, ca. 2 mm in diameter, rugose.

In Costa Rica this species is found in or adjacent

to premontane or montane rain forests of the Cor-

dilleras Tilaran and Central at elevations of 1450

to 2000 m, where flowering collections have been

made in October, November, February, and April.

This species has a range extending to northern and

central Mexico.

Pseuderanthemum praecox is recognized by its

comparatively small, elliptic laminae that are

mostly 2 to 6 cm long, its lax spicate inflorescences

with flowers opposite and borne singly, and its

lavender to bright rose corollas with cylindrical

tubes (16 to 19 mm long) and stamens included.

It most clearly resembles P. standleyi, which dif-

fers in its inflorescences with flowers solitary or in

pairs mostly alternating on the rachis, slightly

shorter corolla tube (ca. 1 5 mm long vs. 1 6 to 19

mm long), its staminodes that are slightly longer

than the filaments (vs. much shorter), and its

shorter, glabrous capsules (10 to 15 mm vs. 15 to

1 8 mm long and puberulous).

Pseuderanthemum standleyi Leonard, Publ. Field

Mus. Nat. Hist., Bot. Ser. 18:1 246. 1 938. Figure

14.

Herbs to 1 m tall; Intel-nodes between leaf-bearing

nodes 8-20 mm long, 0.9-1.1 mm thick, puberulous

mostly along 2 lines, terete. Leaves with petioles 3-6

mm long, puberulous; laminae ovate to lanceolate, 3-8

(9.5) cm long, 1.2-5 cm broad, apically acuminate, ba-

sally attenuate, margins entire to undulate, glabrous to

pilose above, puberulous beneath, mostly at the costa

and veins, the cystoliths dense, moderately prominent

on both surfaces. Inflorescences in terminal spicate pan-

icles and in axillary spikes, the spikes lax, to 9 cm long,

the rachises glandular puberulous; flowers solitary or in

pairs, mostly alternate; bracts and bracteoles subulate,

1.5-3 mm long, ca. 0.5 mm broad basally, puberulous-

glandular. Flowers with 5-merous calyx, calyx segments

linear-subulate, 5-6 mm long, puberulous-glandular; co-

rolla purplish, the tube cylindrical ca. 15 mm long, 1

mm broad, sparingly and minutely puberulous, lobes of

BURGER: FLORA COSTARICENSIS

69

the limb elliptic, ca. 7 mm long, 3.5 mm broad, apically

obtuse; stamens included, the staminodes slightly longer

than the filaments. Fruits clavate, 1 1-15 mm long, 2.5-

3 mm broad, 1.5-2 mm thick, glabrous; seeds 4, oblique,

ca. 2.5 mm long, 2 mm broad, tuberculate.

Found in moist forests and thickets of the Pacific

slopes of the Cordilleras Central and Talamanca,

adjacent Panama, and up to west-central Mexico.

Flowering collections have been made in Decem-

ber and May.

Pseuderanthemum standleyi is recognized by its

lax spicate inflorescences with flowers solitary or

in pairs, mostly alternate on the rachis, its purplish

corolla with a tube about 15 mm long, and its

staminodes, which equal or exceed the length of

the filaments. It most closely resembles P. praecox,

which has the flowers borne singly and opposite,

longer corollas ( 1 6 to 1 9 mm), much shorter stam-

inodes, and longer, puberulous capsules (15 to 18

mm long vs. glabrous, 10 to 15 mm long).

Razisea Oerst.

Shrubs or herbs to 2.5 m tall. Leaves petiolate except

those subtending the inflorescence which are frequently

sessile or subsessile; laminae elliptic to ovate or lanceo-

late, cystoliths dense and prominent on both surfaces of

dried specimens. Inflorescences racemose to thyrsoid,

mostly terminal, sometimes branched; flowers sessile or

pedicellate; bracts and bracteoles small and inconspic-

uous. Flowers with 5-merous calyx, calyx lobes equal;

corolla bilabiate, the upper lip narrow, the lower lip broad

and shallowly trilobate; stamens 2, exserted, the fila-

ments flattened, the anthers 1 -celled; style exserted, the

stigma 2-parted. Fruit a capsule, clavate, 4-seeded.

This is a Central American genus of three species.

Two of the species are endemic to Costa Rica, and

the third, R. spicata, is wide ranging over all of

Central America and Panama. Razisea is very

similar to the genus Kalbreyeriella of Panama and

Colombia, which differs in its beaked corollas prior

to expansion and its calyx lobes which are two to

three times longer than the bracts and bracteoles.

Razisea is also very similar to the genus Hanstei-

nia, which differs in its corolla tube, which is usu-

ally abruptly expanded above the ovary, and in

its paniculate inflorescences. Razisea is also dif-

ficult to distinguish from the genus Habracanthns.

The Costa Rican species of Habracanthus has a

blue corolla, but many of the South American

species have corollas of the same color as Razisea

species (i.e., red or yellow). It is quite possible that

a more comprehensive study of these genera will

bring them into synonymy.

Razisea citrina is known only from two collec-

tions, and R. wilburii, while relatively abundant

where found, is apparently restricted to the Carib-

bean lowlands of Costa Rica.

Key to Species of Razisea

la. Corolla lemon yellow; inflorescence a broad thyrse (cymose panicles) with pedunculate, compound

cymes R. citrina

Ib. Corolla bright red; inflorescence racemose with opposite sessile fascicles of 1-10 flowers 2a

2a. Corolla 4—6 cm long R. spicata

2b. Corolla 2.5-3.3 cm long R. wilburii

Razisea citrina D. Gibson, Fieldiana, Bot. 34: 78.

1972. Figure 14.

Suftrutescent plants to 2 m tall. Inter nodes between

leaf-bearing nodes, 2.5-4.5 cm long, 1.5-3 mm thick,

glabrous. Leaves petiolate, petioles 1.5-5.5 cm long, gla-

brous, leaves subtending inflorescence sessile or subses-

sile; laminae lanceolate, lanceolate-ovate or elliptic, 10-

19.5 cm long, 5-9 cm broad, apically acuminate, basally

acute to attenuate, glabrous on both surfaces, cystoliths

dense and prominent on both surfaces. Inflorescences

terminal, in broad thyrses (cymose panicles) 2-3-flow-

ered; pedicels 5-15 mm long, glabrous; bracts subulate,

3-5 mm long, glabrous, bracteoles similar but smaller.

Flowers with 5-merous calyx, calyx lobes long-subulate

7-9 mm long, glabrous; corolla lemon yellow, glabrous,

3.5-4.5 cm long, the tube ascending and curved outward,

basally 3 mm broad, expanding to 5-11 mm broad at

the mouth, upper lip erect, linear, minutely fimbriate,

lower lip when open, forming an oblique angle from the

upper lip, shallowly trilobate, the lobes ca. equal; sta-

mens exserted 12-15 mm beyond the tip of the upper

lip, anthers ca. 5 mm long. Fruits clavate, ca. 20-25 mm

long, 4-4.5 mm broad, 2.5 mm thick, glabrous, seeds 4,

ovate, 2-2.5 mm long, 1.5-2 mm broad, papillate.

I have seen only two specimens of this species:

the holotype, Jimenez 2475 (F), collected in Oc-

tober in cloud forest area between Agua Buena

70

FIELDIANA: BOTANY

and San Vito (San Vito de Java), Puntarenas Prov-

ince, 1200 m altitude, and Weaver et al. 1775

(DUKE), collected in August at 1000 m altitude,

where it was described as a common shrub along

a stream in the forest. Both specimens were in

flower.

Razisea citrina is recognized by its broad ter-

minal thyrses (cymose panicles) with lemon-yel-

low corollas. It is easily distinguished from other

species of Razisea by these two features.

Razisea spicata Oerst., Vidensk. Meddel. Dansk

Naturhist. Foren. Kjebenhavn 1854: 142. 1855.

Kolobochilus leiorhachis Lindau, Anales Inst.

Fis.-Geogr. Nac. Costa Rica 9: 188. 1898, and

in Pitt., Prim. fl. costaric. 2: 308. 1900. Figure

14.

Shrub to 2.5 m tall; internodes between leaf-bearing

nodes 2-9 cm long, 2-4 mm thick, quadrangular, gla-

brous to infrequently, but densely, pilose. Leaves peti-

olate, petioles 1.0-3.5 (8.5) cm long, glabrous; laminae

elliptic to ovate, 9.5-29 cm long, 3.5-8.5 cm broad, api-

cally acuminate, basally attenuate, acute, obtuse, or

rounded, margins entire to slightly crenate, glabrous

above, glabrous to sparingly puberulous along the costa

and veins beneath, cystoliths dense and prominent on

both surfaces. Inflorescences in terminal racemes to 30

cm long, rachis puberulous to subglabrous, flowers nu-

merous, solitary or in fascicles of 2-1 0 at opposite nodes,

pedicels 5-10 mm long, puberulous, bracts subulate, 5-

10 mm long, 1-2 mm broad, sparingly puberulous, brac-

teoles similar, but smaller. Flowers with 5-merous calyx,

calyx lobes linear, 4.5-9 mm long, ca. 0.5 mm broad,

puberulous; corolla bright red, funnelform, ascending,

slightly curved to one side, 4-6.2 cm long, glabrous, ca.

9 mm broad at the mouth, 1.5-2 mm broad basally,

upper lip erect, linear, to ca. 8 mm long, 4 mm broad

basally, apically rounded, the lower lip with 3 small,

acute lobes at the tip, the lobes equal, ca. 1 mm long,

the lower lip, when open, forming an oblique angle from

the upper lip; stamens exserted 10-20 mm beyond the

tip of the upper lip, the filaments to ca. 4.5 cm long,

glabrous, anther cell 3-4 mm long, the bases rounded.

Fruits clavate, 20-22 mm long, 3.5-4 mm broad, 3.0-

3.5 mm thick, glabrous; seeds suborbicular, ca. 2.5 mm

in diameter, tuberculate.

Found growing at the edges of light gaps and

clearings in lowland rain forests and cloud forests,

this species extends from Colombia to Guatemala.

It has been collected from all provinces in Costa

Rica at elevations of 50 to 1800 m. Flowering

collections have been made from June through

March.

Razisea spicata is recognized by its terminal ra-

cemes with pedicellate flowers either solitary or in

fascicles of two to 1 0 at opposite nodes, its con-

spicuous bright red corollas, which are 4 to 6.2 cm

long, funnelform, ascending, and slightly curved

outward from the rachis, and its two exserted sta-

mens with single-celled anthers. This species is

widespread and shows variation in number of

flowers per node, length of corolla, length of calyx

lobes (4.5 to 9 mm), and pubescence of young

stems. It may be distinguished from the similar

R. citrina, which has lemon yellow corollas and

less variation in calyx lobe length (4.5 to 9 mm

vs. 7 to 9 mm).

Razisea wilburii McDade, Syst. Bot. 7: 489. 1982.

Figure 14.

Shrubs to 1.5 m tall; internodes between leaf-bearing

nodes 1.5-5.5 cm long, 2-3 mm thick, puberulous to

pilose mostly along 2 lines, trichomes white to reddish

brown in a transverse banding pattern. Leaves petiolate,

petioles 1.5-1 1.5 cm long, pubescence that of the stem;

laminae ovate to elliptic or obovate, 12-23.5 cm long,

5-10 cm broad, apically acuminate, basally acute to ob-

tuse, margins entire to crenate, glabrous above, mod-

erately puberulous on costa and veins beneath, cystoliths

dense and prominent on both surfaces. Inflorescences in

terminal racemes borne singly or to 3, to ca. 30 cm long;

flowers numerous, borne singly or in fascicles of 2-3 at

opposite nodes, pedicels 2.5-6 mm long, sparingly pu-

berulous; bracts subulate, 5-6.5 mm long, ca. 1 mm

broad basally, ciliolate, bracteoles similar, but smaller.

Flowers with 5-merous calyx, calyx lobes linear-subu-

late, 4-5 mm long, 0.5-0.75 mm broad basally, glabrous

to sparingly puberulous; corolla bright red, funnelform,

ascending, slightly curved, 2.7-3.3 cm long, ca. 1.5 mm

broad basally, sparingly puberulous, upper lip erect, ovate,

4-6 mm long, 2-3 mm broad, the lower lip truncate,

shallowly trilobate, the lobes ca. 0.5 mm long, stamens

exserted 3-8 mm beyond tip of the upper lip, anthers 3-

3.5 mm long. Fruits clavate, 16-22 mm long, 3-4 mm

broad, 3-3.5 mm thick; seeds 4, suborbicular, 2-3 mm

in diameter, tuberculate.

This species is known mainly from the type lo-

cality, the Organization for Tropical Study Field

Station, La Selva, on the Rio Puerto Viejo near

the town of Puerto Viejo de Sarapiqui. Numerous

collections have been made at this locality, and

its is assumed that the species occurs elsewhere in

the Caribbean lowlands of northern Costa Rica.

Flowering collections have been made in January,

June, September, and October.

Razisea wilburii is recognized by its terminal

racemes with pedicellate flowers either solitary or

in fascicles of two to three at opposite nodes, its

conspicuous bright red corollas which are 2.7 to

3.3 cm long, and its calyx lobe length of 4 to 5

mm. It is most easily confused with R. spicata,

BURGER: FLORA COSTARICENSIS

71

which differs in its longer corolla (4 to 6.2 cm long)

and mostly longer calyx lobes (4.5 to 9 mm vs. 4

to 5 mm long).

Ruellia L.

REFERENCES— R. W. Long, A Biosystematic Ap-

proach to Generic Delimitation in Ruellia (Acan-

thaceae). Taxon 22(5/6): 543-555. 1973. R. W.

Long, Biosystematic Observation on Aphragmia

inundata (Acanthaceae) from Mexico. Rhodora 78:

17-24. 1976.

Perennial herbs or shrubs. Leaves mostly petiolate;

laminae elliptic, ovate, ovate-lanceolate, oblong-spatu-

late, apically acuminate or obtuse, basally attenuate or

acute, the margins entire or undulate, cystoliths present

and usually prominent on both surfaces. Flowers often

large and showy, borne either singly or in various inflo-

rescence forms which are either terminal or axillary or

both, regular, sometimes curved, pedicellate or sessile;

bracts 1 or absent; bracteoles 2 or absent. Flowers with

a 5-merous calyx with mostly equal segments; corolla

bluish, white, yellow, or red, funnelform, salverform, or

sometimes saccate, the tubes usually narrow and the

expanded portion often campanulatc. with 5 spreading,

obtuse lobes; stamens 4, didynamous, the anthers 2-celled,

basally muticous; stigma lobes unequal. Capsule oblong

or clavate, the dissepiment remains attached to the cap-

sule wall at maturity; seeds flattened and, when moist,

rimmed by sticky, mucilaginous hairs, appearing gla-

brous when dry.

This is the second largest genus of the Acantha-

ceae, with approximately 250 species distributed

worldwide. Most species are found in the tropics,

but a few extend into temperate regions.

Key to Species of Ruellia

la. Bracts, calyx, young leaves, and branches of inflorescence all with numerous glandular hairs; calyx

segments unequal in length 2a

Ib. Bracts, calyx, young leaves, and branches of inflorescence not all with numerous glandular hairs;

calyx segments equal in length 5a

2a. Capsules clavate; calyx with 1 segment conspicuously longer than the others . . R. inundata

2b. Capsules elliptic or cylindric; calyx segments unequal, but none conspicuously longer than

the others 3a

3a. Leaves and bracts gland-dotted R. golfodulcensis

3b. Leaves and bracts lacking gland-dots 4a

4a. Stamens slightly exserted; calyx segments 9-1 1 mm long R. paniculata

4b. Stamens included; calyx segments 1 6-20 mm long R. nudiflora

5a. Corolla 6 cm or longer; capsules 22 mm long or greater 6a

5b. Corolla 5.5 cm long or less; capsules 20 mm long or less lOa

6a. Inflorescences spicate; laminae and bracts gland-dotted; calyx lobes 3.5-4 mm long . . 7a

6b. Inflorescences paniculate or flowers solitary to clusters of 5 in the axils of uppermost leaves;

laminae and bracts eglandular; calyx lobes 9-35 mm long 8a

7a. Young stems puberulous; laminae puberulous only on costa and veins beneath

R. tubiflora var. tetrastichantha

7b. Young stems subtomentose; laminae pilose on both surfaces . . . R. tubiflora var. hirsuta

8a. Plants cultivated; inflorescences paniculate; bracts 4-10 mm long R. malacosperma

8b. Plants native; flowers axillary and solitary to clusters of 5; bracts 18-40 mm long .... 9a

9a. Flowers sessile, stamens included; bracts sericeous plus glandular hairs R. praeclara

9b. Flowers on pedicels 2-4 mm long, stamens exserted to just beyond the mouth; bracts glan-

dular-puberulous R. palustris

lOa. Leaves and bracts gland-dotted R. golfodulcensis

lOb. Leaves and bracts lacking gland-dots 1 la

1 la. Plants cultivated for large showy flowers; capsules 25-30 mm long R. malacosperma

1 Ib. Plants native; capsules 6.5-20 mm long 12a

1 2a. Capsules elliptic; younger stems with glandular hairs R. nudiflora var. puberula

1 2b. Capsules clavate; younger stems lacking glandular hairs 1 3a

1 3a. Flowers in compact cymes (often appearing fascicled) at the ends of long unbranched peduncles;

corollas puberulous-glandular R. biolleyi

72

FIELDIANA: BOTANY

13b. Flowers in spikes, spicate panicles, dichotomously branching panicles, or flowers axillary and

solitary to clusters of 2-4; corollas eglandular 1 4a

14a. Flowers axillary and solitary to clusters of 2-4 and some of these occasionally on peduncles to no

more than 5 cm long 1 5a

14b. Flowers in spikes, spicate panicles or dichotomously branching panicles 18a

1 5a. Corollas 4.5-7 cm. long; capsules 14-20 mm long R. jussieuoides

1 5b. Corollas 2-3.5 cm long; capsules 6.5-1 3 mm long 1 6a

16a. Capsules 6.5-8 mm long; leaves sessile or with petioles to 4 mm long R. geminiflora

16b. Capsules 1 1-13 mm long; leaves with petioles 5-15 mm long 17a

1 7a. Corolla puberulous; uppermost bracts 2-4 cm long; capsule minutely puberulous

R. metallica

1 7b. Corolla glabrous; bracts less than 1 5 mm long; capsules glabrous R. barbillana

18a. Corollas 2.8-3.2 cm long; capsules 15-20 mm long; calyx in fruit 6-10 mm long

R. stemonacanthoides

18b. Corollas 2.2-2.8 cm long; capsules 12-15 mm long; calyx in fruit 2-5 mm long 19a

1 9a. Capsule and calyx lobes puberulous plus glandular hairs; peduncle angled, but not winged

R. cooperi

1 9b. Capsule and calyx lobes puberulous to glabrous, but eglandular; peduncle angled and slightly

winged 20a

20a. Peduncles mostly less than 10 cm long, glabrous; younger stems glabrous; bracts elliptic-

oblong R. tonduzii

20b. Peduncles mostly more than 10 cm long, sparingly puberulous; younger stems puberulous;

bracts oblong-spatulate R. pittieri

Ruellia barbillana Cufodontis, Arch. Bot. Sist. 10:

47. 1934.

Herbs to 40 cm tall. Stem quadrangular, bearing nu-

merous punctiform cystoliths, retrorse hairs at the angles.

Leaves on petioles 1-1.5 cm long; laminae ovate-lanceo-

late, to 10 cm long, 2.5 cm broad, apically acuminate,

hasally attenuate, margins subrepand or undulate, gla-

brous above, glabrous to appressed-hirsute on the costa

and veins beneath, cystoliths visible on both surfaces.

Inflorescences of clustered cymes borne in the axils of

the upper leaves, bracts and bracteoles small. Flowers

sessile, with 5-merous calyx, calyx 5 mm long, segments

subulate; corolla pale purple, ca. 3 cm long, glabrous, 2

mm broad hasally, 6 mm broad at the throat, the limb

spreading, the lobes ovate, rounded, ca. 8 mm long, 6

mm broad; stamens included. Fruits glabrous, seeds 6.

This species is known only from the type col-

lection which was made on the bank of Rio Bar-

billa near Finca Waldeck in Limon Province. The

above description was adapted from the original.

Ruellia barbillana is recognized by its inflores-

cences of clustered cymes borne in the axils of

upper leaves, its sessile flowers with bracts less

than 15 mm long, its purplish, glabrous corollas

about 3 cm long, and its glabrous capsules. It is

most easily confused with the more common and

widespread R. metallica, which differs in its longer

bracts (2 to 4 cm long) and its puberulous corollas

and capsules.

Ruellia biolleyi Lindau, in Anales Inst. Fis.-Geogr.

Nac. Costa Rica 9: 188. 1898, and in Pitt., Prim,

fl. costaric. 2: 301. 1900. Ruellia longissima D.

Gibson, Fieldiana, Bot. 34: 79. 1972. Figure 15.

Herbs to 1 m tall, stems erect; internodes between leaf-

bearing nodes 1-14 cm, 1-3.5 mm thick, quadrangular,

yellow pilosulous. Leaves petiolate, petioles 8-22 mm

long, yellow pilosulous; laminae elliptic, 3.5-14 cm long,

2-5 cm broad, apically acuminate, basally attenuate,

margins undulate, glabrous above, puberulous (often yel-

low) along costa and main veins beneath, cystoliths dense

and prominent on both surfaces. Inflorescences in axil-

lary cymes, peduncles 10-23 cm long, quadrangular,

slightly winged, sparingly puberulous; flowers sessile to

pedicels ca. 2 mm long, often in compact fascicles, lower

bracts leaflike, elliptic to spatulate, to 1 5 mm long, upper

bracts similar, but smaller. Flowers with 5-merous calyx,

calyx segments long-subulate, 4-5 mm long, glandular-

puberulous; corollas violet or purplish, 20-24 mm long,

unexpanded portion of tube ca. 1 0 mm long, ca. 1 mm

broad basally, 5.5-7 mm broad at mouth, puberulous

plus some glandular hairs outside; stamens included, an-

thers basally rounded. Fruits clavate, 13-17 mm long,

3.4-4.0 mm broad, 1 .4-1 .5 mm thick, glandular-puberu-

lous, more so at the stipe; seeds 4-8, orbicular, 1.8-2

mm in diameter.

Found in lowland rain forest areas of Panama

and Costa Rica. In Costa Rica it has been collected

in mature or recently cut lowland rain forests from

near sea level to 300 m elevation in Heredia and

BURGER: FLORA COSTARICENSIS

73

Limon provinces. Flowering collections have been

made in Costa Rica in January, February, March,

July, and August.

Ruellia biolleyi is recognized by its long, pedun-

cled ( 1 0 to 23 cm long) inflorescences bearing com-

pact, near-sessile flowers in compact cymes. The

flowers often appear fascicled at the ends of these

long peduncles. Further, it is recognized by its vi-

olet or purplish, puberulous-glandular corollas (20

to 24 mm long) with included stamens, glandular-

puberulous calyx segments (4 to 5 mm long), and

4- to 8 -seeded glandular-puberulous capsules,

which are 1 3 to 17 mm long. It resembles R. coop-

eri, which differs in the more extensive branching

of its inflorescence, thus lacking the often-fascicled

flowers, a white corolla (which may turn purplish

with age) that is slightly longer (2.5 to 2.8 cm long),

and a puberulous calyx and fruit (vs. glandular-

puberulous).

I have placed R. longissima D. Gibson in syn-

onymy here because all of its characteristics lie

within the range of those described for the more

widespread and variable R. biolleyi.

Ruellia cooperi Leonard, Publ. Field Mus. Nat.

Hist., Hot. Ser. 18: 1251. 1938. Figure 15.

Herbs to ca. 1.5m tall, stems erect; inter nodes between

leaf-bearing nodes 3-12 cm long, 2-2.8 mm thick, te-

trangular, subglabrous. Leaves on puberulous petioles 1-

3.5 cm long; laminae elliptic, 5-16 cm long, 1.6-6 cm

broad, apically acuminate, basally attenuate, margins

undulate, glabrous above, puberulous on veins beneath,

cystoliths dense and prominent on both surfaces. Inflo-

rescences of peduncled, axillary, dichotomously branch-

ing panicles, the peduncles 8-30 cm, but mostly more

than 15 cm long, sharply quadrangular, glabrous, the

branches frequently curving; flowers borne singly or in

pairs at the axils of the branches or at their terminus;

bracts elliptic to spatulate, lower bracts leaf] ike. 20-25

mm long, 6-8 mm broad, upper bracts much smaller,

all glabrous. Flowers with 5-merous calyx, lobes 2-5 mm

long, puberulous plus occasional glandular hairs; corolla

white or pale lavender, 2.3-2.5 cm long, sparingly pu-

berulous, unexpanded portion of tube ca. 1 cm long,

7-8 mm broad at the mouth, 1.5 mm broad basally;

stamens included. Fruits clavate, 13-14 mm long, 3-3.6

mm broad, ca. 2 mm thick, puberulous plus occasional

glandular hairs.

Found at lower altitudes of Panama and Costa

Rica. Only one collection, Burger, Visconti and

Gentry 10643 (GRI) from near Quepos, Puntarenas

Province, has been seen. This was collected in Feb-

ruary on shaded slopes above a stream in rain

forest.

Ruellia cooperi is recognized by its inflores-

cences of dichotomously branching panicles on

long peduncles (8 to 30 cm long), capsules which

are puberulous with occasional glandular hairs,

and white to pale lavender corollas which are spar-

ingly puberulous. It most closely resembles R.

biolleyi, which differs in its inflorescences, with

cymose flowers that are short-pedicelled and often

fascicled, and its violet or purplish corollas which

are puberulous with some glandular hairs.

Ruellia geminiflora Kunth in H.B.K., Nov. gen.

sp. 2:240. 1817. Figure 15.

Ascending herbs to ca. 50 cm tall; internodes between

leaf-bearing nodes 1.5-5.5 cm long, 1-1.5 mm thick,

quadrangular, appressed pilose mostly at the angles.

Leaves sessile to petioles 4 mm long; laminae ovate to

ovate-elliptic, 4-7 cm long, 0.9-2.2 cm broad, apically

acute, basally attenuate, margins entire to crenulate, ma-

ture leaves pilose beneath, young leaves and those sur-

rounding the flowers densely puberulous to tomentose,

cystoliths densely distributed over both surfaces where

visible. Pedicels axillary, borne singly or in pairs, ca. 1

mm long. Flowers with 5-merous calyx, lobes equal,

subulate, 5-7 mm long, 0.8-1 mm broad basally, puberu-

lous, ciliate; corolla mauve, funnelform, 2. 1-3.5 cm long,

puberulous, unexpanded portion of tube 8-13 mm long,

ca. 1 mm broad, 6-9 mm broad at the mouth, lobes

equal, suborbicular, 6-10 mm broad; stamens included,

the filaments glabrous; ovary and style puberulous. Fruit

clavate, 6.5-8 mm long, 3.3-3.7 mm broad, 2-2.8 mm

thick, puberulous; seeds 4, orbicular, flattened, 2-3 mm

in diameter.

This species is found in open grassland areas of

Central America, the West Indies, and northern

South America. In Costa Rica it has been collected

in the Pacific watershed areas of Alajuela and

Guanacaste provinces often in secondary succes-

sion following burning. Collections in Costa Rica

have been made at altitudes of 150 to 500 m.

Flowering collections have been made in January,

February, March, May, and June.

Ruellia geminiflora is recognized by its rela-

tively small, subsessile leaves (4 to 7 cm long), its

axillary, short-pedicelled flowers borne singly or

in pairs with mauve corollas (2.1 to 3.5 cm long),

and its short, stout, clavate, puberulous capsules

(6.5 to 8 mm long). It may be confused with R.

metallica, which has larger, petiolate, often lus-

trous leaves (to 15 cm long), inflorescences with

sessile, axillary flowers in fascicles of three to four,

linear to linear-elliptic bracts (2 to 4 cm long) which

intergrade with leaves basally, and its clavate cap-

sules (11 to 13 mm long).

74

FIELDIANA: BOTANY

Ruellia golfodulcensis Durkee, sp. nov. Figure 15.

Herbae suffruticosae, usque ad 1 m altae, caulibus

erectis, glabris. Folia petiolata, laminis ellipticis, 7-15

cm longis, 1.4-4 cm latis, apice acuminatis, basi longe

attenuatis, subglabris et glanduloso-punctatis, cystolithis

supra prominentibus. Inflorescentiae terminales, race-

mosae, angustae, vel in specimenibus junioribus flores

axillares solitarii. Flores corollis albis infundibulifor-

mibus, ca. 4 cm longis, staminibus inclusis. Fructus non

visi.

Suflrutescent plant to 1 m tall, stems erect; internodes

between leaf-bearing nodes 1 .7-8 cm long, 1-2 mm thick,

tetrangular, glabrous, uppermost internodes gland-dot-

ted. Leaves on glabrous petioles 5-1 5 mm long; laminae

elliptic, 7-15 cm long, 1.4—4 cm broad, apically acu-

minate, hasally long-attenuate, margins entire, sparingly

ciliolate, subglabrous and gland-dotted on both surfaces,

gland-dots more prominent in younger leaves, cystoliths

visible on both surfaces but more prominent above. In-

florescences of narrow, terminal racemes in older spec-

imens, with axillary flowers borne singly in younger spec-

imens; bracts lanceolate, 1 5-26 mm long, 3-6 mm broad,

gland-dotted, margins ciliolate. Flowers on glandular-

puberulous pedicels 3-5 mm long; calyx 5-merous, lobes

linear, subequal, 13-20 mm long, 0.75-1.0 mm broad,

glandular-puberulous; corolla white, funnelform, ca. 4

cm long, sparingly glandular-puberulous, tube 2.8 cm

long, unexpanded portion of tube 1.5 cm long, lobes

rounded, 17 mm long, 6 mm broad; stamens included.

Fruits not observed.

TYPE— Costa Rica; Puntarenas Prov., Golfo

Dulce and Rio Grande de Ten-aba, elevation 1 600

m, undershrub in forest, December 1947, Skutch

5304 (holotype, us). Additional Collections: Pun-

tarenas Province, Golfo Dulce area, vicinity of

Esquinas Experiment Station, elevation sea level,

common, April 26, 1949, Allen 5266 (us). Flow-

ering collections were made in April and Decem-

ber.

Ruellia golfodulcensis is recognized by its short-

pedicellate flowers with a conspicuously glandu-

lar-puberulous calyx with subequal, linear lobes

( 1 3 to 20 mm long); white, glandular-puberulous

corollas (ca. 4 cm long) borne singly at leaf or bract

axils; leaves and bracts which are gland-dotted. It

most closely resembles the much wider-ranging R.

metallica of similar habitats, which differs in its

leaves which lack glandular dots, shorter calyx lobes

(5 to 7 mm long) which are subulate, and shorter

corolla (2.3 to 2.7 cm long) which is puberulous

only.

Ruellia inundata Kunth in H.B.K., Nov. gen. sp.

2:239. 1817. Figure 15.

Herbs sometimes suffrutescent, to 1.5 m tall; inter-

nodes between leaf-bearing nodes 2-7.5 cm long, 2.1-

2.8 mm thick, subquadrangular, pilose. Leaves on pilose

petioles 1-8.5 cm long; laminae ovate to ovate-lanceo-

late, 4-15 cm long, 1.2-8 cm broad, apically acuminate,

basally attenuate, margins denticulate, pilose above and

beneath, young leaves glandular-puberulous, cystoliths

common, but more visible above. Inflorescences of as-

cending axillary dichasia which often form dense pani-

cles, pedicels 8-30 mm long, glandular-pubescent; floral

bracts paired, lanceolate, often slightly unequal, 3-5 mm

long, 0.8-1.4 mm broad, puberulous-glandular, lower

bracts similar, but larger. Flowers with 5-merous calyx,

segments linear, 10-14 mm long, 1 segment conspicu-

ously longer (ca. 2 mm) than the others, glandular-pu-

berulous; corolla pink or mauve, narrowly funnelform,

the tube slightly curved, 2.2-2.5 cm long, 1-1.2 mm

broad basally, 3-4 mm broad at the mouth, glabrous to

sparingly puberulous, the 5 lobes equal, rounded 4-5.5

mm long, 3-3.5 mm broad; stamens included. Fruits

clavate, 8-10 mm long, 2.4-3.1 mm broad, 1.2-2 mm

thick, glabrous; seeds 2-4, flattened, orbicular, 2.0-2.5

mm in diameter.

This is a common species growing in open or

partially shaded areas that have been recently dis-

turbed, such as roadsides and trails. Its range ex-

tends from Mexico to much of South America. In

Costa Rica it has been collected only on the Pacific

watershed at altitudes of 50 to 900 m in Alajuela,

Guanacaste, Puntarenas, and San Jose provinces.

Flowering collections have been made from No-

vember through March.

Ruellia inundata is recognized by its ascending

axillary dichasia that often form dense panicles;

pink or mauve, narrowly funnelform corollas (2.2

to 2.5 cm long, excluding lobes); small, paired,

lanceolate bracts (3 to 5 mm long) which are often

slightly unequal in length; and a glabrous, clavate

capsule (8 to 10 mm long). Bracts, calyx, pedicels,

and young leaves all have numerous glandular hairs

which give fresh specimens a fetid odor. It is most

easily confused with R. paniculata, which can be

distinguished by its cylindric capsules (10 to 13

mm long) and slightly exserted stamens.

Long (1976) presents evidence suggesting that

R. inundata belongs in the genus Aphragmia on

the basis of hybridization experiments as well as

morphological comparisons with R. tuberosa, the

type for the genus, and R. caroliniensis. However,

Long does not mention comparisons between R.

inundata and R. paniculata with which it is often

confused because of morphological similarities.

Ruellia paniculata is found in similar habitats and

extends over the same wide range from Mexico to

Brazil. Ruellia paniculata differs from R. inundata

in ways that suggest closer affinity to R. tuberosa,

with its bluish corolla and cylindrical capsule (vs.

red or pink corolla and clavate capsule), suggesting

BURGER: FLORA COSTARICENSIS

75

that R. paniculata does belong in the genus. Until

the relationships among R. inundata, R. panicu-

lata, and other species clearly regarded as within

the genus Ruellia are more thoroughly investigat-

ed, I have decided to recognize R. inundata as

Ruellia.

Ruellia jussieuoides Schlecht., Linnaea 6: 370.

1830. Ruellia achimeniflora (Oerst.) Hemsl.,

Biol. Centralbl. 2: 503. 1882. Arrhostoxylum

achimeniflorum Oerst., Vidensk. Meddel. Dansk

Naturhist. Foren. Kj0benhavn 1854: 131. 1881.

Figure 15.

Herbs or suffnitescent plants to 2.5 m tall; internodes

between leaf-bearing nodes 3-14 cm long, 2-4 mm thick,

tetrangular, puberulous mostly at the angles. Leaves pet-

iolate, petioles 1-3 cm long, puberulous; laminae, ellip-

tic, elliptic-ovate or lanceolate, 5-14.5 cm long, 2-5.5

cm broad, apically long-acuminate, basally attenuate,

margins entire to crenulate, sparingly pilose to pilose

above, pilose beneath, veins and veinlets conspicuous

above in dried specimens, cystoliths dense and promi-

nent on both surfaces. Inflorescences axillary, flowers

sessile and solitary to clusters of 2-3 at the ends of puber-

ulous peduncles 0.25-5 cm long; bracts lanceolate-ellip-

tic, 2-4 cm long, 5-10 mm broad, puberulous, bracteoles

similar though smaller. Flowers with 5-merous calyx,

calyx lobes subequal, long-subulate, 1 lobe often slightly

longer than the rest, 6-20 mm long, 0.8-1 mm broad

basally, ciliolate, puberulous; corolla lavender or purple,

funnelform, frequently curved, puberulous outside, the

tube 3.3-5.5 cm long, ca. 1 mm broad basally, expanding

to 5-10 mm broad at the mouth, the lobes subequal,

suborbicular, 10-18 mm in diameter; stamens included,

anthers ca. 3 mm long. Fruits clavate, 1 4-20 mm long,

3.5-5 mm broad, ca. 2.6 mm thick, puberulous plus

minute glandular hairs; seeds 10-14, elliptic, flattened,

ca. 2.5 mm long, 2 mm broad, puberulous, when wet,

margins covered with dense, mucilaginous hairs.

Ranging from southern Mexico to Costa Rica,

this species is found from 300 to 1 500 m elevation

in Costa Rica where it has been collected mainly

in openings and along trails in rain forests around

San Jose and Cartago provinces, although several

collections have been seen from the vicinity of San

Isidro of Puntarenas Province. Flowering collec-

tions have been made from November through

April.

Ruellia jussieuoides is recognized by its axillary

inflorescences with sessile flowers solitary to clus-

ters of two to three at the ends of peduncles 0.25

to 5 cm long, corollas lavender or purple with

tubes 3.3 to 5.5 cm long, stamens included, fruits

14 to 20 mm long, puberulous. It resembles R.

tubiflora var. tetrastichantha, which differs by its

flowers borne in terminal or lateral spikes resulting

in a conical rachis, bracts and calyx lobes that are

gland-dotted, and white corollas which are 7 to 8

cm long. Ruellia malacosperma also resembles R.

jussieuoides, but differs with its dichotomously

branched cymose inflorescences and its pedicellate

flowers. I have placed Ruellia achimeniflora in

synonymy with R. jussieuoides because I can find

no characters that differ consistently between these

two taxa. Ruellia jussieuoides is much wider rang-

ing, and it appears that R. achimeniflora is a pop-

ulation which differs mainly in wider leaves, but

this variation is inconsistent.

Ruellia malacosperma Greenm., Proc. Amer. Acad.

Arts 34: 572. 1899.

Herbs to 60 cm tall; internodes between leaf-bearing

nodes 1.5-8.5 cm long, 1.8-2.2 mm thick, quadrangular,

sparingly pilose. Leaves on glabrous petioles 0.5-2 cm

long; laminae narrowly elliptic 5-15 cm long, 1-3.4 cm

broad, apically acute or obtusish, margins undulate to

entire, glabrous above and beneath, cystoliths visible on

both surfaces. Inflorescences of ascending axillary di-

chasia that branch once or twice, peduncles quadrangular

to 9 cm long, glabrous pedicels 5-1 5 mm long, sparingly

glandular-puberulous; bracts linear 4-10 mm long, 1-

1.5 mm broad, glabrous. Flowers with 5-merous calyx

with lobes long-subulate, 10-12 mm long, ca. 1.25 mm

broad basally, glandular-puberulous; corolla mauve or

purple, 4.5-5.5 cm long, cylindrical portion ca. '/a the

length, 2.3-3 mm broad, the throat funnelform, ca. 3 cm

broad at the mouth, the lobes rounded, ca. 10 mm long,

1.5-1.8 mm broad; the stamens included. Fruits nar-

rowly clavate, 2.5-3 cm long, 6 mm broad, glabrous;

seeds 8-20, discoid, ca. 3 mm in diameter, silky with

white, appressed hairs when dry.

This species is native to Mexico, but is culti-

vated throughout much of tropical America, in-

cluding Costa Rica, and especially the central val-

ley area. The only Costa Rican collection I have

examined was from Parque Bolivar of La Hon-

dura, San Jose Province. It was collected in flower

in June.

Ruellia melacosperma is recognized by its sim-

ple panicles with one or two dichotomous branch-

es on peduncles to 9 cm long, its large, showy,

mauve or purple corollas (4.5 to 5.5 cm long), and

its large, narrowly clavate, glabrous capsules (2.5

to 3 cm long). It has been confused with R. ste-

monacanthoides, which has smaller corollas (3 to

4 cm long) and shorter capsules (15 to 20 mm

long). It also may be confused with R. paniculata,

which has shorter corollas (3 to 4 cm long) and

shorter, cylindric, glabrous to sparingly puberu-

lous capsules.

76

FIELDIANA: BOTANY

Ruellia metallica Leonard, Publ. Field Mus. Nat.

Hist., Bot. Ser. 18: 1253. 1938. Figure 16.

Suffrutescent herb to 1 m tall; internodes between leaf-

bearing nodes 1-9.5 cm long, 1-3 mm thick, subqua-

drangular, sparingly puberulous mostly at the angles.

Leaves on glabrous petioles 5-13 mm long, glabrous.

Laminae elliptic to elliptic-ovate, varying from broadly

to narrowly so, 4-17 cm long, 1-8 cm broad, apically

acuminate, basally attenuate, margins undulate, glabrous

above, sericeous on the costa and veins beneath, cys-

toliths prominent above and beneath, upper surface often

faintly lustrous or purplish, the lower surface often pur-

plish or violet purple beneath, especially in younger leaves.

Inflorescences axillary or terminal with sessile flowers in

fascicles of 3-4; uppermost bracts linear to linear-ellip-

tic, 2—4 cm long, lower bracts intergrading with leaves.

Flowers with calyx segments equal, subulate, 5-7 mm

long, glabrous except scant puberulence at the margins;

corollas lavender, pink, or white, funnelform, 2.3-2.7

cm long, puberulous, cylindrical portion ca. 1.5 mm

broad, 10 mm broad at the mouth, lobes suborbicular,

5-7 mm in diameter; stamens included. Fruits clavate,

1 1-13 mm long, 2.5-3.2 mm broad, 2.2-2.6 mm thick,

minutely puberulous; seeds 2, suborbicular, 2.5-2.8 mm

in diameter.

This species occurs in and adjacent to lowland

rain forest areas ranging from Guatemala to Pan-

ama. In Costa Rica it has been collected in moist

areas of both watersheds, excluding Guanacaste

and San Jose provinces. Flowering collections have

been made in January, February, March, May,

June, September, and December.

Ruellia metallica is recognized by its terminal

spikelike inflorescences with axillary fascicles of

three to four sessile flowers subtended by bracts

which are linear to linear-elliptic (2 to 4 cm long)

near the apex, but intergrading with leaves basally,

its lavender, pink, or white corollas (2.3 to 2.7 cm

long), its clavate, minutely puberulous capsules

(11 to 13 mm long), and its leaves which are often

purplish or violet purple beneath, lustrous or pur-

plish above. It is most easily confused with R.

geminiflora, which differs with its smaller (4 to 7

cm long) sessile to short petiolate leaves, its in-

conspicuous bracts, its inflorescences with pedi-

cellate, axillary flowers in fascicles of one to two,

and its clavate capsules (6.5 to 8 mm long).

Ruellia nudiflora (Engelm. & Gray) Urb., Symb.

antill. 7: 382. 1912. Dipteracanthus nudiflorus

Engelm. & Gray, Boston J. Nat. Hist. 5: 229.

1845. Ruellia nudiflora (Engelm. & Gray) Urb.

var. puberula Leonard, J. Wash. Acad. Sci. 17:

517.1 927. Ruellia puberula (Leonard) Tharp &

Barkley, Amer. Midi. Naturalist 42: 17. 1949.

Figure 16.

Herbs to 30 cm tall; internodes between leaf-bearing

nodes 1-8 cm long, 1.4-2.7 mm thick, quadrangular,

glandular-puberulous plus occasional larger pilose hairs.

Leaves on glandular-puberulous petioles 0.4-3.2 cm long;

laminae elliptic to obovate, 3.5-11.5 cm long, 1.3-3.7

cm broad, apically acute, obtuse or rounded, basally at-

tenuate, margins undulate, puberulous above mixed with

occasional pilose hairs, similar beneath, but densely pu-

berulous, cystoliths dense, more visible above, obscured

by hairs beneath. Inflorescences of axillary dichasia, pe-

duncles 2-8 cm long, glandular-puberulous, pedicels 0.5-

2 cm long, glandular-puberulous, bracts oblong, 7-9 mm

long, glandular-puberulous. Flowers with calyx lobes lin-

ear-subulate, equal, 16-20 mm. long, glandular-puber-

ulous; corolla light blue or lavender, funnelform, 3.5-

4.5 cm long, tube ca. 2 mm broad basally, expanding to

8-15 mm broad at the mouth, puberulous, the lobes

suborbiculate, 9-1 1 mm broad; stamens included. Fruits

elliptic, 16-18 mm long, 2.5-4 mm broad, puberulous;

seeds ca. 12, ovate, basally asymmetrical, ca. 2.5 mm

long, 2.0 mm broad.

Found in dry savannas and pastures, this species

survives burning and is then conspicuous for its

showy flowers appearing close to the ground after

growth is resumed. This variety ranges from Oa-

xaca to central Panama. In Costa Rica it is found

at elevations of 25 to 100 m in the lowlands of

Guanacaste. Flowering collections have been made

in January, February, May, and June.

Ruellia nudiflora is recognized by its puberu-

lous, elliptic to obovate laminae, its axillary di-

chasia on glandular-puberulous peduncles to 8 cm

long bearing pedicellate flowers with light blue or

lavender corollas 3.5 to 4.5 cm long with stamens

excluded, and its puberulous, elliptic capsules 1 6

to 1 8 mm long. It may be confused with R. pan-

iculata, which differs in its elliptic to ovate leaves,

its corollas with stamens slightly exserted, and its

glabrous, cylindric capsule (10 to 13 mm long).

Ruellia palustris Durkee, sp. nov. Figure 16.

Herbae suffruticosae, usque ad 1 m altae, caulibus

erectis, junioribus pilosis-adpressis. Folia petiolata, lam-

inis ellipticis vel elliptico-obovatis, 5.5-15.5 cm longis,

2.2-8.5 cm latis, apice acuminatis, basi acutis vel parum

attenuatis, maximam partem glabris, cystolithis abun-

dantibus. Inflorescentiae fasciculatae, axillares et ter-

minates, 1-5 floribus, bracteis 2 in quoque flore. Flores

corollis albis infundibuliformibus, tubis 6-7 cm longis

et lobis 1 .6-2 cm longis, staminibus paulo exsertis. Fruc-

tus clavati, ca. 23 mm longi.

Suffrutescent herb to 1 m tall; stems erect; internodes

between leaf-bearing nodes 3-1 5 cm long, 2-4 mm thick,

appressed-pilose. Leaves petiolate, petioles 6-18 cm long,

BURGER: FLORA COSTARICENSIS

77

appressed-pilose; laminae elliptic to elliptic-obovate, 5.5-

15.5 cm long, 2.2-8.5 cm broad, apically abruptly acu-

minate, basally acute to slightly attenuate, margins entire

to slightly undulate, glabrous on both surfaces except for

appressed hairs on costa and main veins, cystoliths abun-

dant and visible on both surfaces. Inflorescences in ax-

illary and terminal clusters of 1-5 flowers, flowers borne

on short pedicels 2-4 mm long, strigose; bracts 2 per

flower, shape variable from broadly elliptic to obovate

or oblanceolate, 3-4 cm long, 0.6-2.3 cm broad, apically

acute, glandular-puberulous. Flowers with a 5-merous

calyx, calyx lobes equal, long-elliptic to lanceolate, 25-

30 mm long, 4-5 mm broad, glandular-puberulous; co-

rolla white, funnelform, the tube 6-7 cm long, cylindrical

portion 4.5-5 cm long, 3.5-4 mm broad, funnelform

portion 1.5-2 cm long, 10-16 mm broad at the mouth,

glandular-puberulous, the lobes equal, suborbicular, 1 .6-

2 cm long, 2-2.5 cm broad; stamens exserted to just

beyond the mouth, anthers 4-5 mm long, ca. 0.8 mm

broad, filaments attached at ca. the beginning of the

funnelform portion of the corolla tube. Fruits clavate,

ca. 23 mm long, 8 mm broad, 8 mm thick, puberulous;

seeds 8.

TYPE— Costa Rica: Heredia Prov. Collected at

swamp's edge in mature rain forest at Finca La

Selva on Rio Puerto Viejo, elev. ca. 100 m, 27

July 1980, Hammel 9341 (Holotype, DUKE). Ad-

ditional Collections— All from the same vicinity

and similar habitat, Grayum 1467, 2428 (DUKE);

Hammel 10480 (DUKE), McDade 364 (DUKE); Da-

mon Smith 614 (DUKE). Flowering collections have

been made in July, August, and November.

Ruellia palustris is recognized by its large white

corollas (6 to 7 cm long) with two-thirds of the

tube cylindrical, the two large, mostly elliptic bracts

per flower and with axillary and terminal flowers

in clusters of one to five. It most closely resembles

R. praeclara of similar habitats in western Panama

and southeastern Costa Rica, which differs in its

shrubby habit, longer corollas (to 9 cm), and linear

bracts and calyx lobes.

Ruellia paniculata L., Sp. pi. 2: 635. 1753. Figure

16.

Herbs to suffrutescent plants; internodes between leaf-

bearing nodes, 2-6.5 cm long, 1.8-3.5 mm thick, qua-

drangular, glandular-puberulous. Leaves on petioles 0.5-

4 cm long, glandular-puberulous; laminae ovate to el-

liptic, 3-10(17) cm long, 1 .5-4.5 (6.5) cm broad, apically

acute to obtuse, basally abruptly attenuate, margins un-

dulate, glandular-puberulous plus occasional pilose hairs

on both surfaces, but more dense and prominent above,

less so beneath. Inflorescences in axillary, branching di-

chasia forming dense clusters in mature plants, the

branches glandular-puberulous; floral bracts narrowly el-

liptic to obovate, 3-5 mm long, 0.5-0.7 mm broad, glan-

dular-puberulous, lower bracts larger intcrgrading with

the leaves. Flowers with calyx segments elongate-subu-

late, unequal, 9-1 1 mm long, basally 0.7-0.8 mm broad,

glandular-puberulous; corolla light blue, funnelform,

puberulous, 2.5-4 cm long, unexpanded portion of tube

ca. 1.3 cm long, 1-1.2 mm broad, lobes suborbiculate,

6-7 mm broad; stamens slightly exserted. Fruits cylin-

dric, 12.5-15 mm long, 2-2.5 mm in diameter, glabrous

to minutely puberulous apically; seeds 8, flat, ovate, ca.

2 mm long, 1.5 mm broad.

This species is wide ranging from Mexico to

Brazil where it grows in open disturbed habitats

with a dry season at elevations of sea level to 200

m. It is not frequently collected, either because it

is uncommon or, as I suspect, it is mistaken for

the much more common R. inundata of similar

habitats. Only two Costa Rican collections have

been observed from the Bagaces and Canas areas

of Guanacaste, both in flower— one in January,

the other in April.

Ruellia paniculata is recognized by its dense ax-

illary dichasia, with most parts bearing a dense

glandular-puberulence, blue corollas (2.5 to 4 cm

long) with slightly exserted stamens, and cylindric

capsules (12.5-15 mm long). It is most often con-

fused with R. inundata of similar habitat, which

differs in its pink or mauve corollas, included sta-

mens, and clavate capsules (8 to 10 mm long).

Ruellia pittieri Lindau, Bull. Herb. Boissier 5:655.

1 897. Ruellia longissima var. glabra D. Gibson,

Fieldiana, Bot. 34: 80. 1972. Figure 16.

Herbs to suffrutescent plants to 2 m tall; internodes

between leaf-bearing nodes, 2.8-1 1 .5 cm long, 3-3.8 mm

thick, quadrangular, puberulous. Leaves on petioles 2-

6.5 cm long, puberulous; laminae ovate to ovate-elliptic,

7-12 (19) cm long, 2.5-7 (8.5) cm broad, apically acu-

minate, basally acute to obtuse, margins undulate, gla-

brous above, puberulous on costa and veins beneath,

cystoliths dense and prominent on both surfaces. Inflo-

rescences in axillary, lax, spicate panicles that are often

dichotomously branching; peduncles 1 2-2 1 cm long, pe-

duncles and branches sparingly puberulous and slightly

winged; nonfloral bracts oblong-spat ulate, 15-35 mm

long, glabrous; floral bracts elliptic, 4-8 mm long, 1-2

mm broad, glabrous. Flowers sessile to subsessile, with

calyx segments subulate, 2-3 mm long at anthesis, 3.5-

5 mm long at fructescence, ca. 0.5 mm broad basally,

sparingly puberulous; corolla light blue or pale violet,

funnelform, 2.3-2.8 cm long, puberulous, unexpanded

portion of tube 8-10 mm long, 1-1.5 mm broad, 5-10

mm broad at the mouth, lobes equal, obtuse, 5.5-7 mm

long, 5-7.5 mm broad; stamens included. Fruits clavate,

12-15 mm long, 3-4 mm broad, 1 .9-2.2 mm thick, spar-

ingly puberulous to glabrous; seeds 4, 6, or 8, suborbicu-

lar, 2.2-2.8 mm in diameter.

78

FIELDIANA: BOTANY

This species is commonly found along streams

and other openings in wet forests of Costa Rica

and Panama. In Costa Rica it is found in southern

Puntarenas Province at elevations of 30 to 270 m.

Most collections have been made in the Golfo

Dulce area and on the Burica Peninsula. Flowering

collections have been made in December, March,

and April.

Ruellia pittieri is recognized by its lax, dichot-

omous, spicate panicles, with slightly winged, te-

trangular peduncles (and branches) which usually

exceed 10 cm in length; its oblong-spatulate bracts,

its funnelform corollas (2.3 to 2.8 cm long), and

its clavate capsules (1 2 to 15 mm long). It is easily

confused with R. stemonacanthoides, which differs

in its longer corollas (2.8 to 3.2 cm long), its longer

calyx lobes at anthesis (4 to 7 mm long vs. 2 to 3

mm), and its longer capsules (15 to 20 mm long).

Ruellia pittieri is also easily confused with R. ton-

duzii, which differs in its shorter peduncles, (most-

ly less than 1 0 cm long), its oblong-elliptic bracts,

and while the size ranges overlap, its generally

shorter leaves (6 to 13.5 cm long vs. 7 to 19 cm).

Ruellia praeclara Standley, Publ. Field Colum-

bian Mus., Hot. Ser. 4: 263. 1929. Figure 16.

Suffrutescent plants to 2 m tall; internodes between

leaf-bearing nodes 2.4—4.5 cm long, 2.3-3.0 mm thick,

quadrangular, appressed pilose. Leaves on petioles 1.5-

3 cm long, glabrous to appressed pilose; laminae oblan-

ceolate, 5.5-30 cm long, 1.6-9 cm broad, apically acu-

minate, basally acute to attenuate, margins entire to un-

dulate, glabrous to sparingly strigose at the costa and

veins, the cystoliths prominent on both surfaces. Flowers

axillary, sessile, solitary to clusters of 3; bracts linear to

linear-oblanceolate, 18-30 mm long, 2-3 mm broad,

sericeous plus glandular hairs. Calyx segments linear to

linear-lanceolate, 9-35 mm long, 1.5-2 mm broad, se-

riceous with numerous glandular hairs; corolla white, 8-

10 cm long, funnelform, puberulous-glandular, unex-

panded portion 4-5.5 cm long, 3 mm broad, expanding

to 9-14 mm broad at the mouth, lobes orbicular 15-20

mm in diameter; stamens included, but extending to just

below the mouth. Fruits not seen.

Found in the shade of lowland rain forests of

western Panama (Caribbean side) and one site in

the Osa Peninsula of Costa Rica. Three flowering

collections have been made, two in January by

Burger & Liesner 7202 (F, NY), 7329 (F), and one

in March by Burger & Gentry 8915 (us), all from

near Rincon de Osa of Puntarenas Province. The

Costa Rican population is at the lower range of

measurements for leaves, bracts, and calyx lobes.

Also, bracts and calyx lobes are linear-oblanceo-

late and linear-lanceolate, respectively, in the Cos-

ta Rican population as opposed to linear for both

in the Panamanian population.

Ruellia praeclara is recognized by its sessile,

axillary, solitary flowers with white corollas (8 to

10 cm long), its large lanceolate leaves (to 30 cm

long), and its linear to linear-lanceolate calyx lobes

(9 to 35 mm long) with both sericeous and nu-

merous smaller glandular hairs. It may be con-

fused with R. tubifora var. tetrastichantha or var.

hirsuta, both of which differ in their terminal and

subterminal spicate inflorescences, the numerous

glandular spots on the lower surfaces of leaves and

bracts, and much shorter calyx lobes (3.5 to 4 mm

long vs. 9 to 35 mm long).

Ruellia stemonacanthoides (Oerst.) Hemsl., Biol.

Centralbl. 2: 507. 1882. Arrhostoxylum stemo-

nacanthoides Oerst., Vidensk. Meddel. Dansk

Naturhist. Foren. Kjobenhavn 1854: 130. 1855.

Figure 17.

Herbs to suffrutescent plants 1 m tall; internodes be-

tween leaf-bearing nodes, 3-10.5 cm long, 1.3-3.2 mm

thick, quadrangular, puberulous. Leaves on petioles to

3.5 cm long, puberulous to glabrous; laminae ovate-el-

liptic, 4-12 cm long, 2-5.7 cm broad, apically acuminate,

basally acute and often slightly attenuate, margins un-

dulate, glabrous above, frequently drying to a purplish

color, puberulous beneath on costa and veins, cystoliths

dense and visible on both surfaces although occasionally

obscure. Inflorescences in axillary panicles, often di-

chotomously branched, peduncles to 1 1 cm long, qua-

drangular, often slightly winged, glabrous; bracts elliptic,

10-27 mm long, 2-6 mm broad, glabrous. Flowers ses-

sile to pedicles 3 mm long, puberulous, with calyx seg-

ments linear-subulate, 4-7 mm long at anthesis, 6-10.5

mm long at fructescence, sparingly puberulous; corolla

pale violet, funnelform, 2.8-3.2 cm long, puberulous,

unexpanded portion of tube 9-11 mm long, 1 . 1-1 .5 mm

broad, 10-12 mm broad at the mouth, lobes equal, ob-

tuse, 6-9 mm long, 5-8 mm broad; stamens included.

Fruits clavate, 15-20 mm long, 3-5 mm broad, 2.25-

4.5 mm thick, sparingly puberulous; seeds 8-10, subor-

bicular, 2.5-3.2 mm in diameter.

This species is commonly found in or near re-

cently cut forests or in new pastures in premontane

forest areas of Costa Rica to Guatemala and west-

ern Mexico. In Costa Rica it has been collected

mainly from the Pacific watershed areas of San

Jose, Alajuela, and Guanacaste provinces, with

one collection from the Guapiles area of Limon

Province, all at elevations of 300 to 1000 m. Flow-

ering collections have been made in September

through November, January, February, April, and

July.

BURGER: FLORA COSTARICENSIS

79

Ruellia stemonacanthoides is recognized by its

paniculate inflorescences which are often dichot-

omously branched on peduncles to 1 1 cm long, its

pale violet corollas (2.8 to 3.2 cm long), its calyx

lobes (4 to 7 mm long at anthesis), and its sparingly

puberulous, clavate capsules ( 1 5 to 20 mm long).

It is easily confused with R. pittieri and R. ton-

duzii. See the description of R. pittieri to distin-

guish these differences.

Ruellia tonduzii Lindau, Anales Inst. Fis.-Geogr.

Nac. Costa Rica 9: 1 88. 1 898, and in Pitt, Prim,

fl. costaric. 2: 203. 1900. Figure 17.

Erect herbs to ca. 50 cm tall; internodes between leaf-

bearing nodes 2.5-7 cm long, 1 .5-2.5 mm thick, quadran-

gular, slightly winged, glabrous. Leaves on petioles 2-3

cm long, puberulous; laminae elliptic to ovate, 3.5-12.5

cm long, 1.1-5.2 (7) cm broad, apically acuminate, ba-

sally acute and frequently attenuate or obtuse, margins

undulate, glabrous above, puberulous on costa and veins

beneath, cystoliths dense and prominent on both sur-

faces. Inflorescences in axillary, dichotomously branch-

ing panicles on glabrous, quadrangular, slightly winged

peduncles 5-9.8 cm long; bracts elliptic-oblong, 8-13

mm long, 1.5-3 mm broad, glabrous. Flowers borne sin-

gly on glabrous pedicels to 3 mm long; calyx segments

subulate, 2.7-4 mm long, 0.5 mm broad basally, gla-

brous to subglabrous; corolla lavender, funnelform, 2.2-

2.8 cm long, puberulous, unexpanded portion of tube

ca. 9 mm long, 1.5 mm broad, 7 mm broad at the mouth,

lobes equal, obtuse, 5-7 mm long, 3.5-5.5 mm broad,

stamens included. Fruits clavate, 12-15 mm long, 3-4

mm broad, 2.5 mm thick, sparingly puberulous; seeds

8, suborbiculate, 2.4-2.6 mm in diameter.

This species is found in the shade of moist forest

areas of the Pacific coastal regions of Panama and

Costa Rica. In Costa Rica it has been collected in

Puntarenas and San Jose provinces at elevations

of 5 to 340 m. L. D. Gomez 2208 (F, MO, NY) was

not included in this description because of the

corolla color described as yellowish red and its

location at 2100 m elevation in Heredia, although

in other respects this specimen very closely resem-

bles R. tonduzii. Flowering collections have been

made from February through August.

Ruellia tonduzii is recognized by its axillary,

dichotomously branching panicles on glabrous,

quadrangular peduncles (ca. 5 to 9 cm long), flow-

ers borne singly on short pedicels (to 3 mm long),

elliptic-oblong bracts (8 to 13 mm long), calyx

lobes (2.7 to 4 mm long), and capsules (12 to 15

mm long). It is easily confused with R. pittieri and

R. stemonacanthoides. (See the description of R.

pittieri to distinguish these differences.)

Ruellia tubiflora Kunth in H.B.K., Nov. gen. sp.

2: 241. 1817. Figure 17.

Ruellia tubiflora H.B.K. var. tetrastichantha (Lin-

dau) Leonard, Contr. U.S. Natl. Herb. 31:91.

1951. R. tetrastichantha Lindau, Anales Inst.

Fis.-Geogr. Nac. Costa Rica 2: 300. 1900. Fl.

Costaricensis. 2: 300. 1900.

SufTrutescent plants to 3 m tall; internodes between

leaf-bearing nodes 4.5-1 1 cm long, 2.5-4.5 mm thick,

quadrangular, puberulous. Leaves on petioles to 5 cm

long, puberulous; laminae elliptic, 10-23 cm long, 3.5-

10 cm broad, apically acuminate, basally attenuate, mar-

gins undulate, glabrous above, puberulous on costa and

veins to glabrous beneath, gland-dotted beneath, occa-

sionally gland-dotted above, cystoliths visible above, ob-

scure beneath. Inflorescences in compressed, short, ter-

minal, and subterminal spikes with a rachis to 1.2 mm

long; bracts, pale green to greenish white, leafy, elliptic,

4-5.5 cm long, 12-18 mm broad, gland-dotted; brac-

teoles small, subulate, ca. 2 mm long. Flowers sessile;

calyx segments subulate, 3.5—4 mm long, 1 mm broad

basally, puberulous; corolla white or yellow, funnelform,

7.5-9 cm long, puberulous, often gland-dotted, unex-

panded portion of tube 3-3.5 cm long, 2.5-3.5 mm broad,

expanded portion 1 5-23 mm broad at the mouth, lobes

suborbicular, 9-15 mm long, 1 1-20 mm broad; stamens

included. Fruits clavate, 2.2-3 cm long, 7.4-8 mm broad,

6 mm thick, glabrous to minutely puberulous; seeds 8-

12, suborbicular, 4.5-5.5 mm in diameter.

Comparatively wide ranging, this species is found

in shaded areas of wet forests or thickets at lower

to middle elevations of Costa Rica, Panama, and

Colombia. In Costa Rica it has been collected in

Alajuela, Cartago, Limon, and Puntarenas prov-

inces at elevations of 450 to 900 m. Flowering

collections have been made in January through

March.

Ruellia tubiflora var. tetrastichantha is recog-

nized by its inflorescences of very short, terminal

spikes (1.2 cm or less long), its white, funnelform

corollas (7.5 to 9 cm long) which are often gland-

dotted, its leaves which are gland-dotted beneath,

and its pale green to greenish white bracts which

are also gland-dotted. It most closely resembles

R. tubiflora var. tubiflora and var. hirsuta. Variety

tubiflora is distinguished by its red corolla, and

var. hirsuta, by its subtomentose internodes in

younger stems and its leaves with relatively long

pilose hairs on both surfaces. It can also be con-

fused with R. praeclara, which differs in its axillary

flowers, its calyx segments (1 to 3 cm long) with

sericeous hairs combined with numerous shorter

glandular hairs, and its puberulous glandular co-

rolla.

80

FIELDIANA: BOTANY

Ruellia tubiflora H.B.K. var. hirsuta Leonard,

Contr. U.S. Natl. Herb. 31:93.1951. Figure 1 7.

This variety resembles R. tubiflora var. tetra-

stichantha most closely and differs mainly in its

hairiness. Leafy internodes are subtomentose, and

the leaves and bracts are pilose on both surfaces.

This variety is also found in central Colombia.

Only one collection, Allen 6316 (F, MO), has been

made in Costa Rica in the Esquinas Forest be-

tween Rio Esquinas and Palmar at 30 m. The

collector described it as very common in the forest.

This flowering collection was made in late No-

vember.

Sanchezia Ruiz & Pavon

REFERENCES— E. C. Leonard, Notes on the genus

Sanchezia, J. Wash. Acad. Sci. 16: 484-492. 1926.

E. C. Leonard and L. B. Smith, Sanchezia and

related American Acanthaceae. Rhodora 66: 3 1 3-

343. 1964.

Erect or climbing herbs or shrubs, mostly glabrous.

Leaves petiolate; laminae with numerous cystoliths on

both surfaces, glabrous. Flowers solitary or more often

fascicled, those that are fascicled subtended by bracts

which are often large, partly connate and showy; calyx

5-merous; corolla usually large and showy, corolla tube

cylindric. the lobes 5 and equal; stamens 2, usually

exserted, the anthers 2-celled, mucronulate basally, pub-

erulous, staminodes 2. Fruit an oblong capsule, 6-8 seeds;

seeds orbicular.

This genus contains about 59 species, all native

to tropical America and found mainly in wet forest

areas of the northern Andes with two exceptions:

S. pennellii Leonard extends into the Darien Prov-

ince of eastern Panama, and 5". parvibracteata is

found in Mexico and Central America, but prob-

ably escaped from cultivation.

Sanchezia parvibracteata Sprague & Hutchinson,

Bull. Misc. Inform. 253. 1908.

Shrubs to 2.5 m tall, stems subquadrangular, glabrous;

internodes between leaf-bearing nodes, 4-6 cm long, 4-

5.5 mm thick. Leaves with short, glabrous petioles to 3

cm long; laminae elliptic to obovate, 1 2-24 (35) cm long,

5-1 1 (1 5) cm broad, apically abruptly acuminate, basally

attenuate, margins undulate-serrate, glabrous, costa and

veins edged in yellow, cystoliths numerous and promi-

nent on both surfaces, ca. 0.5 mm long. Inflorescences

in sparingly branched terminal panicles to 20 cm long;

bracts loosely imbricate, ovate, 15-20 mm long, 7.5-10

mm broad, apically obtuse, glabrous; bracteoles oblong

ovate, ca. 1 5 mm long, 4 mm broad, apically rounded,

glabrous; peduncles to 5 cm long, glabrous; rachis gla-

brous. Flowers 3-5 per pair of bracts, sessile; calyx seg-

ments oblong, 2-2.5 cm long, 3.5-5 mm broad, apically

obtuse, minutely puberulous at apex outside, glandular-

puberulous inside; corolla red or ochre, puberulous with

yellowish appressed hairs, 4-5 cm long, 6-7 mm broad

at the throat, 5 mm broad just above ovary, lobes oblong,

reflexed, 5-7 mm long, 5 mm broad; stamens exserted

to 8 mm beyond the mouth of corolla, filaments sparsely

villous, anthers 6-7 mm long; staminodes 1 5 mm long,

sparingly pilose. Fruit not seen.

In Central America this species is usually found

in gardens. It is probably native to the Northern

Andes since collections have been made there

(Leonard, 1951). Only one Costa Rican collection

has been observed. This was a flowering collection

from a thicket in the Monteverde area, altitude

1 500 m, Puntarenas Province, Palmer 36769 (CR).

Sanchezia parvibracteata is recognized by its

flowers in fascicles of three to five loosely enclosed

by a bract, its large red or ochre, equally five-lobed

corolla, and its exserted stamens with two-celled

anthers which are basally spurred. It is not easily

confused with other Acanthaceae from Costa Rica.

Spathacanthus Baillon

Shrubs or small trees. Leaves petiolate; laminae usu-

ally large, oblong, or elliptic. Flowers large, few, borne

on small terminal branches; bracts small; calyx spatha-

ceous, valvate; corolla usually large, white, the tube sal-

verform, the limb 5-lobed, bilabiate, the lobes subequal;

stamens 4, didynamous; anther lobes parallel. Fruit a

capsule, large, glabrous.

There are five species in this genus of Mexico

and Central America. Because of the spathaceous,

two-lobed calyx, this genus is not easily confused

with other genera of the family.

Spathacanthus hoffmannii Lindau, Bull. Herb.

Boissier 3: 370. 1895. Figure 17.

Shrubs or small trees to 8 m tall; internodes between

leaf-bearing nodes 1.5-4.5 cm long, 2-3 mm thick, gla-

brous. Leaves subsessile to petioles 10 mm long, gla-

brous; laminae elliptic to elliptic-obovate, 7.5-27 cm

BURGER: FLORA COSTARICENSIS

81

long, 3-7.5 cm broad, apically acuminate, basally long

attenuate, margins undulate, glabrous on both surfaces,

cystoliths dense and prominent on both surfaces. Flow-

ers axillary, solitary, in clusters of 2 or 3 or in small

racemes, peduncles to ca. 2 cm long, glabrous; pedicels

6-10 mm long, glabrous; bracts subulate, ca. 2 mm long,

1 mm broad basally, bracteoles similar but smaller.

Flowers with a spathaceous calyx 2.2-3.5 cm long, gla-

brous, fused for ca. half its length and then separating

into 2 mostly equal lobes that are apically acute and

apiculate, lobes 8-12 mm broad; corolla white, 8-8.7

cm long, glabrous, ca. 2 mm broad basally, expanding

to 13-15 mm broad at the mouth, the lobes subequal,

obtuse, 12-15 mm long, 12-15 mm broad; stamens di-

dynamous, included, anthers 2-celled, ca. 4.5 mm long,

basally rounded. Fruits clavate, 7-8 cm long, 12-14 mm

broad, ca. 1 1 mm thick, glabrous; seeds 4, flattened,

suborbicular, 9-10 mm in diameter, 2-3 mm thick, sur-

face dark brown, rugose.

This species is found in moist habitats at middle

elevations in forest clearings most frequently along

streams and rivers. I have seen no collections of

it outside Costa Rica where it has been collected

at elevations of 750 to 1750 m, mostly in the

Caribbean watershed areas of Alajuela, Cartago,

and Heredia provinces. One collection, Standley

& Valeria 44745 (F) is known from the Pacific

watershed, Tilaran area, Guanacaste. Flowering

collections have been made in July through Oc-

tober and in January (the Guanacaste collection).

Spathacanthus hoffmannii is recognized by its

woody habit, its large elliptic to elliptic-obovate

leaves, its large, showy, white flowers with a five

equally lobed corolla, two large, lobed spathaceous

calyx and strongly didynamous stamens with two-

celled anthers which are rounded basally, and its

large, clavate fruit. With this combination of char-

acters, it is not easily confused with other Acan-

thaceae.

Streblacanthus Kuntze

Herbs to suffrutescent plants. Leaves petiolate. Inflo-

rescences of lax terminal and axillary spikes; bracts nar-

rowly oblong to filiform, long and conspicuous. Flowers

with a 4-merous calyx, segments narrow and equal; co-

rolla tube expanded basally, narrowed to a slender throat,

the limb bilabiate, spreading, the upper lip 2-lobed, the

lower lip 3-lobed; stamens 2, exserted, the filaments short,

attached to the throat of the corolla, anthers 2-celled,

the cells unequal and widely separated, 1 usually fertile,

calcarate basally, the other much smaller, rounded ba-

sally and often abortive; staminodes absent. Fruit a slen-

der clavate capsule; seeds 2-4.

This is a genus of four species with one species

each described from Mexico, Guatemala, Costa

Rica, and Bolivia.

Streblacanthus monospermus Kuntze, Revis. gen.

pi. 2: 498. 1891. Streblacanthus macrophyllus

Lindau, Anales Inst. Fis.-Geogr. Nac. Costa Rica

9: 188. 1898, and in Pitt., Prim. fl. costaric. 2:

306. 1900. Streblacanthus longiflorus Cufodon-

tis, Arch. Bot. (Forli). 10: 48. 1934. Figure 17.

Herbs to 1 m tall; internodes between leaf-bearing

nodes 3-5.5 cm long, 3-5.5 mm thick, tetrangular, pu-

berulous along 2 lines in younger portions to glabrous

in older. Leaves on petioles to 2.2 cm long, puberulence

of petioles that of the stem; laminae elliptic, 6-19 cm

long, 2.5-8 cm broad, apically acuminate, basally acute,

margins entire to sparingly crenulate, glabrous to spar-

ingly strigose on both surfaces, cystoliths visible on both

surfaces. Inflorescences in terminal and axillary spikes,

to 25 cm long, peduncle to 2.5 cm long, brownish pilose;

rachis brownish pilose-glandular; bracts narrowly ellip-

tic to oblong, 13.5-20 mm long, 3-4 mm broad, densely

pilose-glandular, bracteoles long-subulate, 6-10 mm long,

1-2 mm broad basally, pilose-glandular. Flowers with

4-merous calyx, lobes subulate, 4—4.5 mm long, 0.8-1

mm broad basally, pilose-glandular; corolla lavender or

white, salverform, puberulous, the tube 2-3 cm long, 1.5

mm broad expanding to 3 mm broad around the ovary,

lobes of the upper lip obovate, 9-10 mm long, 7-8 mm

broad, apically acute, lower lip very similar, lobes ca. 5

mm broad; stamens exserted just beyond the mouth with

a tuft of pilose hairs at the apex of each filament; anther

cells unequal and widely separated, the upper cell 1 mm

long, basally calcarate, the lower 0.5 mm long and pro-

ducing pollen. Fruits clavate, long-stipitate, 2-2.5 cm

long, 5.5-7 mm broad, ca. 2 mm thick, pilose-glandular;

seeds 4, orbicular, 5-6 mm in diameter.

This species is apparently endemic to Costa Rica

where it is found in lowland rain forests of the

Caribbean coastal region. Flowering collections

have been made in March, May, and September.

Since the type collection was made at the Rio Yor-

kin at the Panamanian border, this plant should

also be found in adjacent Panama.

Streblacanthus monospermus is recognized by

its white to lavender salverform corollas with two

exserted stamens on very short filaments and

widely separated anther cells, the upper cell cal-

carate and the lower cell much smaller and lacking

a spur. After examining the types of S. mono-

spermus, Kuntze s.n. (us) and S. macrophyllus,

Lindau 8547 (us), I have concluded that these are

the same species. The only difference between them

is in the length of the corolla lobes, and more

82

FIELDIANA: BOTANY

recent collections of S. monospermus show that

the original measurements were made from im-

mature corollas.

Teliostachya Nees

REFERENCES— C. G. Nees von Esenbeck, Acan-

thaceae. In DC, Prodr. 11: 262-264. 1847. C. E.

B. Bremekamp, Notes on the Acanthaceae of Suri-

nam. Recueil Trav. Bot. Neerl. 35: 149. 1938.

Herbs, ascending or decumbent. Leaves petiolate;

laminae ovate to lanceolate with numerous cylindrical

cystoliths, the margins entire. Inflorescences in dense

spikes either terminal or terminal and axillary, the spikes

either cylindrical or ovoid; pedicels arranged in whorls

of 3-7, the lateral flowers of each whorl subtended by a

1 - to 3-nerved bract; 2 bracteoles subtending the lateral

flowers, 1 -nerved. Flowers with a 5-merous calyx, seg-

ments unequal, divided nearly to the base, the posterior

lobe much larger than the rest, 3-nerved, the lateral lobes

1 -nerved, the anterior lobes 1- or 2-nerved, longer than

the lateral nerves; corolla about as long as the calyx, the

tube cylindric, 2-lipped, the upper lip obtuse, emarginate

or subentire, the lower lip 3-lobed, the lateral lobes ob-

tuse, the middle lobe emarginate or obtuse; stamens 4,

didynamous, the anthers 2-celled, unequal, the lower cell

and sometimes both basally apiculate; stigma 2-lobed,

the style curved toward the upper lip.

This is a genus of about 1 0 species occurring

throughout much of tropical America and in West

Africa. Only one species occurs in Central Amer-

ica. It closely resembles Lepidagathis, a genus en-

demic to the Old World. According to Breme-

kamp, Teliostachya differs in its definitely terminal

spikes, calyx segments which are almost entirely

free as opposed to at least partially united seg-

ments, its unequal anther cells as opposed to equal

cells, and its pollen grains which are entirely free

from reticulations as opposed to the finely retic-

ulate grains of Lepidagathis.

2.5-8 cm long, 1 .5-3.2 cm broad, apically acute to slight-

ly acuminate, basally attenuate, margins entire, gla-

brous above, strigose mostly on costa and veins beneath,

cystoliths dense on both surfaces. Inflorescences in dense

terminal or axillary cylindrical spikes to 5 cm long, 1.3

cm broad; flowers in sessile whorls of 3-5; 1 bract sub-

tending each flower, obovate lanceolate, 5-6 mm long,

1.5-2.2 mm broad, apically caudate, basally acute, thin,

translucent, veins prominent, 3-nerved, transparent when

wet, the margins ciliate, bracteoles 2, lanceolate linear,

ca. 5 mm long, 0.4 mm broad, translucent with prom-

inent veins, 1 -nerved, ciliate. Flowers with a 5-merous

calyx, segments unequal, the posterior pair oblanceolate,

4-5.5 mm long, the remaining segments lanceolate, 3-6

mm long, all venose, ciliate; corolla white to violet, tu-

bular, 3.5-4 mm long, glabrous, 2-lipped, upper lip ca.

2.5 mm long, 1 mm broad, bidentate, lower lip 2.5 mm

long, 3-lobed, lobes ca. 1 mm long; stamens exserted.

Fruits oblong, 3.3-3.7 mm long, 1-1.1 mm broad, 0.8-

1 mm thick, apically acuminate, basally acute, seeds 4,

near ovate, flattened ca. 1 mm long, 8 mm broad, basally

oblique, pilose when dry, hairs densely mucilaginous

when wet.

This species occurs in moist areas along streams

and trails in rain forests, in thickets, and in open

fields. Its range includes most of South America,

Central America, and the West Indies. Only one

Costa Rican collection has been seen: Weston et

al. 4292 (us) from near Repunta, 1 5 km south of

San Isidro, San Jose Province, altitude 600 m. This

flowering collection was made in January. Collec-

tions from neighboring countries have been made

from near sea level to 1400 m elevation.

Teliostachya alopecuroidia is a short, small-

leaved herb that is recognized by its dense, ter-

minal spikes with flowers containing a fi ve-merous

calyx with very dissimilar segments, the posterior

and anterior segments longer and broader than the

lateral segments, and its inconspicuous two-lipped

corolla with four didynamous anthers which are

two-celled and cells superposed. It is not easily

confused with other species of Acanthaceae.

Teliostachya alopecuroidea (Vahl) Nees in Mart.,

Fl. bras. 9: 72. 1 847. Ruellia alopecuroidea Vahl,

Eclogae americanae 2: 49. 1798. Lepidagathis

alopecuroides (Vahl) R. Br. ex Griseb., Fl. Brit.

W.I. 453. 1864. Figure 18.

Herbs to 25 cm tall; internodes between leaf-bearing

nodes 2-5.5 cm long, 1-1.5 mm thick, quadrangular,

puberulous mostly at the angles. Leaves on puberulous

petioles to 1 cm long; laminae ovate to ovate-elliptic,

Tetramerium Nees

REFERENCE— G. B. Happ, Monograph of Tetra-

merium and Henrya. Ann. Missouri Bot. Gard.

24: 501-582. 1937.

Herbs to suffrutescent plants; stems erect or declining,

much-branched, pubescence often arranged in two op-

posite lines, the old bark often exfoliating. Leaves sessile

or petiolate; laminae linear to ovate, glabrous or pubes-

cent. Inflorescences in terminal and axillary 4-sided

BURGER: FLORA COSTARICENSIS

83

spikes; bracts opposite, 4-ranked, conspicuous, often

closely imbricate with mucronate apex; pedicels 1-3 per

bract. Flowers with a 4-5-merous calyx of equal, ciliate

segments; corollas funnelform to tubular, bilabiate, the

upper lip entire, the lower lip 3-lobed, the lobes ca. equal;

stamens 2, included in the upper lip, the anthers 2-celled,

cells slightly unequal. Fruits clavate, apiculate, 4- or 2-

seeded; seeds flattened, tuberculate, or muriculate.

Twenty-three species have been recognized for

this genus, and they range from the southern United

States to Colombia and Ecuador.

Tetramerium nervosum Nees in Benth., Bot. voy.

Sulphur 148, pi. 48. 1844. T. hispidum Nees in

DC., Prodr. 11: 468. 1847. Figure 18.

Repent to upright herbs to ca. 30 cm tall; internodes

between leaf-bearing nodes 2.5-6.5 cm long, 1.25-1.5

mm thick, terete, sparingly pilose. Leaves on pilose pet-

ioles to 15 mm long; laminae ovate, 2-7 cm long, 0.8-

3 cm broad, apically acute to acuminate, basally rounded

to acute, margins entire, sparingly strigose on both sur-

faces, numerous cystoliths visible on both surfaces. In-

florescences in terminal and axillary spikes to 7.5 cm

long, 1.5 cm broad; bracts clearly 4-ranked, imbricate,

ovate to elliptic, 7-1 1 mm long, 3.5-7 mm broad, api-

cally mucronate, typically curved outward, basally ob-

tuse, mostly 5-nerved, puberulous plus longer pilose hairs,

ciliate; bracteoles narrowly lanceolate, 4-6.5 mm long,

ca. 1 mm broad, ciliolate; peduncle 1-4 mm long, puber-

ulous. Flowers with 4-merous calyx, calyx segments sub-

ulate, 3-3.5 mm long, ca. 0.5 mm broad, ciliolate; corolla

white or slightly yellowish, bilabiate, 16-17 mm long,

the tube narrowly funnelform, 6.5-7.5 mm long, ca. 1.25

mm broad at the throat, 1 mm broad basally; the upper

lip of one lobe, the lower lip 3-lobed, all lobes oblan-

ceolate, 7-9 mm long, 2.5-3.5 mm broad, the lobes of

lower lip at the narrower range; stamens exserted to ca.

the length of the upper lip, attached to base of middle

lobe of lower lip, cells of the anthers slightly unequal,

ca. 1.5 mm long, gray or bluish. Fruit clavate, 5-6 mm

long, 2.25-2.5 mm broad, 1.75-2.25 mm thick, puber-

ulous; seeds 4, rhomboid, 1 .75-2 mm long, 1.5-1.75 mm

broad, surface tuberculate.

This species is found in secondary growth pas-

tures, along roadsides and trails throughout trop-

ical America, and extends into the southwestern

United States. In Costa Rica it is found from near

sea level to 500 m elevation. It has been collected

from Guanacaste and the Pacific Coastal area of

Alajuela. Because this is a weedy species and omit-

ted in collecting, it is of much more common oc-

currence in Costa Rica than these collections would

suggest. Flowering collections have been made from

December through March.

Tetramerium nervosum is recognized by its

dense, terminal and axillary spikes with common-

ly outcurving, imbricate, mucronate bracts that

are clearly four-ranked, its bilabiate, four-parted

white or yellowish corollas with the two exserted

stamens attached at the base of the middle lobe

of the lower lip, and its two-celled anthers that are

gray or bluish. It is sometimes confused with Ble-

chum brownei, but can be distinguished from it

by its four-merous calyx and corolla (vs. five-mer-

ous) and its two stamens (vs. four that are didyn-

amous).

Thunbergia Retz.

REFERENCE— C. E. B. Bremekamp, Delimitation

and subdivision of the Acanthaceae, Bull. Bot.

Surv. India 7: 21-30. 1965.

Herbaceous to suffrutescent vines or shrubs. Leaves

petiolate with hastate, rhomboid, or ovate laminae, cys-

toliths absent. Flowers large, peduncled, solitary in the

axils or in terminal or axillary racemes; bracts 2, folia-

ceous, large; calyx short, cupuliform, truncate, or toothed;

corolla 5-merous; stamens 4, didynamous, attached near

the base of the corolla tube, anthers 2-celled with an

apiculate connective; ovary fleshy, the style, apically di-

lated; ovary subtended by a conspicuous, fleshy nectary.

Fruit a 2-chambered capsule with a globose base ex-

tending abruptly into a flattened beak; seeds 2 in each

locule, borne on papilliform funicles, semiglobose to

ovoid, ventral side conspicuously excavated.

This is a genus of about 200 species mainly, if

not exclusively, from tropical Africa and Asia.

Key to Species of Thunbergia

1 a. Petioles winged; corollas orange or yellow, often with a dark purple throat (rarely white) T. alata

Ib. Petioles not winged; corollas blue, scarlet, white, or yellow 2a

2a. Flowers solitary in the leaf axils; calyx reduced to 1 0-20 teeth 3a

2b. Flowers in terminal or axillary racemes; calyx reduced to a basal ring or annulus 4a

3a. Erect shrub; laminae basally acute T. erecta

84

FIELDIANA: BOTANY

3b. Vine; laminae basally hastate to cordate T. fragrans

4a. Corolla with blue lobes, yellowish or white throat; bracts irregularly oblong, 3-3.5 cm long

T. grandiflora

4b. Corolla scarlet, tube yellow; bracts ovate, 2-2.5 cm long T. mysorensis

Thunbergia alata Bojer ex Sims, Bot. Mag. 52: pi.

2591. 1825. Figure 18.

Herbaceous trailing or climbing vine; internodes be-

tween leaf-bearing nodes 4.5-13 cm long, 1-1.25 mm

thick, strigose. Leaves with petioles to 6.5 cm long, pet-

ioles winged ca. 1 mm broad on each side, hirsute; lam-

inae deltoid to deltoid-ovate, 3.5-7.5 cm long, 2.5 cm

broad, apically acute, basally hastate to cordate, margins

undulate, sparingly to moderately hirsute on both sur-

faces. Flowers axillary and solitary, peduncles to 8.5 cm

long, hirsute; bracts 2 per flower, deltoid-ovate, 18-20

mm long, 9-10 mm broad, apically acute hirsute; calyx

toothed with ca. 15-17 subulate lobes ca. 2 mm long,

glandular-puberulous; corolla yellow or orange with dark

purple throat; salverform, ca. 4 cm long, tube 2 cm long,

lobes 5, ca. 2 cm long, apically emarginate. Fruits densely

puberulous, 16-18 mm long, base ca. 7 mm in diameter,

beak ca. 1 1 mm long, 5 mm broad, 2 mm thick; seeds

4, ca. 3.5 mm in diameter, honeycombed.

This species is native to eastern Africa and is

widely cultivated in tropical America. It frequent-

ly escapes from cultivation and is found in fields

and along trails and the edges of thickets. In Costa

Rica it has been collected from near sea level to

about 1000 m elevation. Flowering collections have

been made from November through March and

in July.

Thunbergia alata is recognized by its herba-

ceous, viny habit, its deltoid to deltoid-ovate lam-

inae with winged petioles, and its yellow or orange

salverform corolla with five equal emarginate lobes

and a dark purple throat. It is most easily distin-

guished from other species of the genus by its leaves

with winged petioles. There is a variety with white

corollas and another with a white throat and or-

ange lobes, but I have seen no collections of these

from Costa Rica. Thunbergia alata most closely

resembles T. fragrans, which has similar though

white corollas and lacks winged petioles.

Thunbergia erecta (Benth.) T. Anderson, J. Linn.

Soc., Bot. 7: 18. 1864. Meyenia erecta Benth.,

Niger. Fl. 476. 1849.

Shrub to 3 m tall; internodes between leaf-bearing

nodes 2.5-6.5 cm long, 1.25-2.5 mm thick, quadran-

gular, glabrous. Leaves with glabrous petioles to ca. 5

mm long; laminae elliptic, 1.5-6.5 (8.5) cm long, 0.8-

2.6 (4.3) cm broad, apically acute, basally acute, margins

undulate, glabrous to subglabrous on both surfaces.

Flowers solitary in leaf axils, peduncles to 2.8 cm long,

glabrous; bracts oblong-ovate, 10-20 mm long, 5-8 mm

broad, apically acute, basally obtuse, glabrous to mi-

nutely puberulous at the apex. Flowers with the calyx

reduced to 10-16 subulate teeth ca. 3 mm long, glan-

dular-puberulous; corolla all white or deep violet lobes

with a yellowish throat, broadly salverform, glabrous,

6-8 cm long, 6-8 mm broad basally constricting to 2-3

mm broad just above the ovary, expanding to 1 5-20

mm broad at the throat, lobes suborbicular, 15-25 mm

in diameter; cells of the anther unequal, the base and

margins of the anther cells puberulous. Fruit not ob-

served.

This species is a native of tropical West Africa,

but is widely cultivated. It has become naturalized

from cultivation in many tropical areas, but all

Costa Rican collections I have observed were from

cultivation. These were from near sea level in Li-

mon Province and from about 1000 m elevation

from San Jose. Flowering collections were made

in June and November.

Thunbergia erecta is recognized by its shrubby

habit and its all white or violet, lobed, yellow-

throated corollas from 6 to 8 cm long. The all white

corolla may be confused with that of T. fragrans,

and the violet, yellow-throated corolla may be

confused with that of T. grandiflora, but both of

these species have a viny habit and shorter corol-

las.

Thunbergia fragrans Roxb., PI. Coromandel 1: 47.

1795.

An herbaceous to somewhat woody vine; internodes

between leaf-bearing nodes 3-11.5 cm long, 1-2 mm

thick, quadrangular, strigose. Leaves with strigose peti-

oles to 2.5 cm long; laminae ovate to ovate lanceolate,

5-1 1.5 cm long, 2-6.5 cm broad, apically acute, basally

hastate to cordate, margins entire to slightly undulate,

strigose on both surfaces. Flowers solitary in the leaf

axils, pedicels to 8 cm long, strigose; bracts deltoid ovate,

1.3-2 cm long, 5-8 mm broad, apically acute, strigose.

Flowers with a calyx of 12-20 teeth, the teeth ca. 3 mm

long, puberulous; corolla white, salverform, pilose, the

tube 2-2.5 cm long, 3-5 mm broad, the lobes cuneate,

BURGER: FLORA COSTARICENSIS

nearly as long as the tube, crenate; anther cells ca. 3 mm

long, basally acute. Fruit ca. 2.5 cm long, the beak sub-

ulate, flattened, 1-1.5 cm long.

A native of India, this species is rather widely

cultivated, as it is in Costa Rica. Collections from

cultivation have been made from near sea level to

about 1 000 m elevation. Flowering collections were

made in October, February, April, and June.

Thunbergiafragrans is recognized by its twining

habit and solitary flowers with a toothed calyx and

a white corolla. It most closely resembles T. alata,

which differs in its winged petioles and yellow co-

rollas with a dark purplish throat.

Thunbergia grandiflora Roxb., Hort. bcngal. 45.

1814.

Suffrutescent climbing or twining vine; inter nodes be-

tween leaf-bearing nodes 6-19 cm long, 2.6-3.2 mm

thick, subquadrangular, glabrous. Leaves on scabrous

petioles to 6 cm long; laminae ovate to ovate-hastate,

8-16 cm long, 3-13.5 cm broad, apically acuminate,

basally hastate or rounded, margins often coarsely toothed

or lobed below the middle, scabrous mostly on costa and

veins on both surfaces. Inflorescences of elongated ter-

minal and axillary racemes with 2-4 flowers at each

node; peduncles to 4.5 cm long, quadrangular, glabrous

to sparingly scabrous; pedicels to 3.3 cm long, sparingly

scabrous; bracts irregularly oblong, 3-3.5 cm long, 17-

20 mm broad, apically acute, sparingly puberulous plus

numerous dark glandular appearing dots. Flowers with

a very much reduced annular calyx at the base of the

corolla, margin of annulus puberulent; corolla light or

dark blue with a white or yellowish throat, campanulate,

the tube 3.0—4.5 cm long, lobes 3-4 cm long; bases of

the anther cells of the posterior stamens each with spurs

to 3 mm long, only 1 anther cell spurred on each of the

anterior stamens. Fruit base ca. 1 6 mm in diameter, beak

flattened, ca. 2 cm long.

This species is native to Bangladesh and is widely

cultivated in tropical regions of the world. Only

collections from cultivation have been made in

Costa Rica in the Central Valley area from Ala-

jucla. Heredia, and San Jose provinces. Flowering

collections were made in September and March.

Thunbergia grandiflora is recognized by its

twining habit, its loose terminal and axillary ra-

cemes, its annular calyx, and its large, showy, cam-

panulate corolla with blue lobes and a white or

yellowish throat. It is not easily confused with oth-

er cultivated species of Thunbergia.

Thunbergia mysorensis (Wight) T. Anderson ex

Bedd. in Trans. Linn. Soc. London 25. 222: 1 865.

Hexacentris mysorensis Wight, PI. asiat. rar. Ill:

78. t. 871. 1832.

A woody vine; internodes between leaf-bearing nodes

2-8.5 cm long, 1-1.5 mm thick, subquadrangular, gla-

brous. Leaves with glabrous petioles to 1.6 cm long;

laminae lanceolate-oblong, 5-9.5 cm long, 1.2-2.3 cm

broad, apically attenuate, basally obtuse to subcordate,

margins undulate, mostly 3-nerved, glabrous on both

surfaces. Inflorescences in terminal and axillary racemes;

peduncles to 6 cm long, glabrous; pedicels to 2.5 cm

long, glabrous; bracts red, ovate, 2-2.5 cm long, 8-10

mm broad, apically acute, glabrous. Flowers with a very

much reduced annular calyx at the base of the corolla;

corolla zygomorphic, lobes scarlet, tube yellow, 5-5.5

cm long, throat opening obliquely with lower lobes re-

flexed and 2 upper lobes fused most of their length to

partially enclose the stamens and style, stamens with

anthers ca. 8 mm long, basally long-spurred, spurs ca. 4

mm long. Fruits not observed.

Native to India, this species is occasionally cul-

tivated in Costa Rica for its showy inflorescences.

I have observed only two Costa Rican collections,

Jimenez 447 (F) and Sanchez 4 (F). Both were in

flower, one collected in January, the other in March.

Thunbergia mysorensis is recognized by its lan-

ceolate-oblong leaves with obtuse to subcordate

bases, its showy flowers in terminal or axillary

racemes with red bracts, its annular calyx, and its

zygomorphic corollas with scarlet lobes and yellow

tubes. It is not easily confused with other Thun-

bergia species grown in Costa Rica.

Trichanthera H.B.K.

Shrubs or trees. Leaves petiolate; laminae ovate to

oblong, entire to undulate, numerous cystoliths visible

on the upper surface of dried specimens. Inflorescences

in compact terminal panicles (or corymbs). Flowers sub-

tended by 2 small, triangular bracteoles; calyx 5-merous

with sepals separate, equal and mostly obtuse; corolla

regular, 5-lobed, tomentose, campanulate; stamens 4,

exserted, didynamous, the anthers ciliate, with 2 equal

basally rounded cells. Fruit a 2-chambered capsule with

2 seeds per locule.

Trichanthera is a genus of two species ranging

from Central America to Brazil. It is most easily

confused with the genus Bravaisia, but can be dis-

tinguished from it by its basally rounded anthers

(vs. spurred) and one- to four-seeded capsules (vs.

eight-seeded).

Trichanthera gigantea (Humb. & Bonpl.) Nees in

DC, Prodr. 11: 218. 1809. Ruellia gigantea

86

FIELDIANA: BOTANY

Humb. & Bonpl., PI. aequinoct. 2: 75, pi. 102.

1809.

REFERENCE— T. B. Croat, Flora of Barro Colo-

rado Island, p. 790. Stanford Univ. Press. 1978.

Shrubs or small trees to 10 (17) m tall; internodes

between leaf-bearing nodes 4-6 mm thick, tetrangular,

brown tomentose. Leaves with petioles 1.5-6 cm long;

laminae ovate to elliptic, 10-26 cm long, 6-10 cm broad,

apically acuminate to acute, basally acute to obtuse or

oblique, margins entire to slightly undulate, the surfaces

glabrous except sparingly pilose on the costa of larger

leaves. Inflorescences in terminal, compact panicles

tending to be secund, to 1 5 cm long, 6 cm broad, branch-

es light brown tomentose. Flowers subtended by 2 tri-

angular bracteoles to 3 mm long; sepals 10-12 mm long,

tomentose, apically rounded to acute; corollas shiny yel-

low in the throat, brownish red in the expanded portion,

3-4 cm long, glabrous at throat, silky tomentose above,

silky tomentose outside, the tube 1-1.5 cm long, the limb

2-3 cm across; lobes 4-6 mm broad, oblong to oblong-

ovate; ovary brown tomentose, style 4-5 cm long. Fruits

1.5-2 cm long, 5 mm broad, apically obtuse, sericeous;

seeds flattened, lenticular, 3-4 mm in diameter.

This is a wide-ranging species of moist lowland

forests, frequently along streams and swampy areas.

While it is not abundant in areas where collected,

it extends from Costa Rica through Colombia and

Venezuela to the mouth of the Amazon in Brazil.

Although T. gigantea occurs in Costa Rica (Leon-

ard, 1938), this description is based upon Pana-

manian collections. On Barro Colorado Island,

Panama, Croat has observed flowering from Jan-

uary to April, especially in February and March.

Trichanthera gigantea is recognized by its ar-

borescent habit; paniculate inflorescences with to-

mentose sepals and corolla; more or less equally

five-lobed corollas 3 to 4 cm long; didynamous

stamens with two-celled anthers that are basally

rounded; sericeous capsules with one to four seeds.

It may be confused with Bravaisia integerrima,

which differs in its basally spurred anthers and

glabrous capsules that are eight-seeded.

PLANTAGINACEAE

By William Burger

Herbs (rarely subshrubs with few-branched woody

stems in island endemic species ofPlantago), annual or

perennial, stems usually very short above the ground

with closely congested internodes, xylem in discrete bun-

dles or forming a cylinder, a rhizome often present; stip-

ules absent. Leaves alternate (rarely opposite), simple

and usually from basal rosettes, petioles often not strong-

ly differentiated from the lamina, leaf-base clasping the

stem; laminae linear to broadly ovate, entire or with

small lobes or teeth, glabrous or puberulent, venation

palmate or parallel. Inflorescences solitary from each leaf

axil, flowers usually in dense spikes or capitula on a long,

unbranched peduncle, each flower subtended by a single

bract. Flowers small and sessile or subsessile, bisexual

(in ours) or unisexual, radially symmetrical and usually

4-parted (rarely 3-parted); sepals 4 (3), free or partly

united, imbricate in bud; petals united to form a short

tube, the 4 (3) corolla-lobes spreading or reflexed at an-

thesis, membranous to scarious, white to yellowish, often

persisting; stamens as many as the corolla-lobes and al-

ternate with them (1 or 2 in Bougueria), filaments borne

on the corolla-tube and inflexed in bud, anthers exserted

and versatile, 2-thecous, introrse and longitudinally de-

hiscent, pollen 4-20-porate; pistil 1 , ovary superior, usu-

ally 2-(3, 4)-locular in Plantago with 1 to many ovules

in each locule, 1-locular with a single basal ovule in

Bougueria and Littorella, style 1 , stigma 1 - or 2-lobed.

Fruit a thin-walled capsule opening at or below the mid-

dle circumscissilly (a pyxis) in Plantago, a small nut in

Bougueria and Littorella; seeds often lustrous, endo-

sperm present (absent in Littorella), embryo straight

(curved in Bougueria).

A family of three genera and about 250 species

in temperate, arctic, and montane habitats. Bou-

gueria nubicola is found in the high Andes of

southern South America. Littorella is a genus of

semiaquatic, freshwater habitats, with one species

in Europe, one in North America, and a third

species in southern South America. Plantago is

cosmopolitan in cooler or montane habitats and

the only genus found in Central America. While

its close relationships are obscure, the tubular co-

rolla and bilocular ovary with axile placentation

(in Plantago) has suggested an affinity with the

Scrophulariales.

Plantago Linnaeus

Herbs (in ours) or subshrubs, annual or perennials,

usually scapose plants with very short stems, clustered

basal leaves and erect, unbranched peduncles bearing

small, congested flowers. Leaves all basal and rosulate

in Central American species, alternate and tightly con-

gested (in ours), petioles present or absent, poorly dif-

ferentiated from the lamina, clasping the stem at their

base; laminae linear to broadly ovate, glabrous or more

often puberulent, entire or with short blunt lobes or teeth,

venation palmate or parallel. Inflorescences spicate or

capitate with long, erect, unbranched peduncles, flowers

sessile in the axils of bracts. Flowers small and bisexual,

sessile in the axils of bracts, sepals 4, free or partly united,

often unequal with 2 longer and 2 shorter, imbricate in

bud, margins scarius, petals united to form a short tube,

BURGER: FLORA COSTARICENSIS

the 4 lobes reflexed or spreading at anthesis, persistent

and sometimes forming a cover over the fruit; stamens

4 or 2, exserted at anthesis; ovary 2-locular or 3- or 4-

1 ocular by intrusion of the placentae, ovules 1 to several

in each locule. Fruit a thin-walled capsule with circum-

scissile dehiscence around the middle; endosperm pres-

ent, embryo straight.

A genus of worldwide range but restricted to

temperate, arctic, and montane environments; ab-

sent in the lowland tropics except on islands. A

number of island endemics have unusual woody

growth forms, but most species have a very short

aerial stem, leaves arising from near the ground

and erect scapose inflorescences. These plants can

be mistaken for sedges and other monocotyledons

because of the often narrow leaves with parallel

venation and the small, tightly clustered flowers

subtended by brownish bracts. The small flowers

with short corolla-tube, four scarious corolla-lobes,

long, slender filaments, exserted versatile stamens,

and fruit with circumscissile dehiscence easily dis-

tinguish these plants. Dr. Knud Rahn has revised

many New World species (see Nordic Journal of

Botany 3: 331-342, 1983, and included refer-

ences).

Key to Species of Plantago

la. Leaves becoming broadly ovate, lamina abruptly narrowed at the base (in larger leaves); corolla-

lobes 0.5-1 mm long; ovary with many ovules, seeds usually more than 8 P. major

1 b. Leaves elliptic to oblanceolate, laminae tapering gradually at the base to join the poorly differentiated

petiole; corolla-lobes 2-3 mm long; ovary with 6 or fewer ovules, seeds fewer than 7 2a

2a. Corolla-lobes persisting and erect in fruit, forming a pointed "cap" at the apex of the fruit; ovary

with 3-6 ovules, seeds 1-6; native and common P . australis

2b. Corolla-lobes not persisting as a cap above the fruit; ovary with 2 ovules; seeds 1 or 2; introduced,

rare . .P. lanceolata

Plantago australis Lamarck, Encycl. Meth. Bot.

Illust. Gen. 1: 339. 1793. P. hirtella H.B.K.,

Nov. gen. sp. 2: 187 or 229, t. 127. 1817. P.

hartwegii Decne. in DC, Prodr. 13, pt. 1: 724.

1 852. P. schiedeana Decne. in DC., loc. cit. 723.

P. sodiroana Pilger, Bot. Jahrb. Syst. 25: 733.

1898. P. ecuadorensis Pilger, loc. cit. 50: 232.

1913. P. australis ssp. hirtella (H.B.K.) Rahn,

Bot. Tidsskr. 60: 50. 1964. Figure 19.

Herbs, perennial, stems very short above the ground,

internodes congested and obscured by the leaf-bases.

Leaves usually in basal rosettes, quite variable in size on

different plants, petioles to 15 cm long; laminae 4-25

(35) cm long 0.5-5 (7) cm broad, lanceolate to oblan-

ceolate or narrowly elliptic-obovate, obtuse to acute at

the apex, gradually narrrowed to the base and continuous

with the petiole, margin entire or slightly undulate, very

sparsely (rarely glabrous) to densely puberulent on both

surfaces with slender septate and translucent hairs 0.5-

1 .5 mm long, drying thin-chartaceous and greenish, ve-

nation parallel with (3) 5 or 7 primary veins. Inflores-

cences 3-50 (100) cm long, peduncle sparsely to densely

whitish villous with hairs to 2 mm long, flowering por-

tion 3-50 cm long, flowers usually congested near the

apex, more separate near the base, bracts 1.6-4.2 mm

long, 0.6-1.4 mm wide, triangular to ovate, usually cil-

iate along the margin. Flowers with sepals 1.7-2.7 mm

long, ca. 1 mm broad, 1 often longer than the other 3,

usually puberulent along the keel and ciliate along the

margin distally, corolla-lobes 2-3.5 mm long, narrowly

ovate, usually erect and spread apart only at anthesis,

persistent and becoming stiff in fruit; anthers 1.2-1.6

mm long; ovary usually with 3 ovules. Fruit 2—4 mm

long, to 6 mm together with the persisting beaklike im-

bricate corolla-lobes, pale brown; seeds 1.2-2.4 mm long,

ca. 1 mm broad, oblong, flat and with a small circular

scar on the inner face, pale brown, smooth.

Plants of open, sunny sites in regions of wet,

evergreen, montane forest formations between

1500 and 3200 m elevation in Costa Rica; to as

low as 400 m in northern Central America. Inflo-

rescences are produced throughout the year, but

there may be a flowering peak in August. The

species ranges from Arizona and Mexico through

the Central Highlands of Central America and

western Panama through the highlands of northern

and central South America to Bolivia, Brazil, and

Argentina.

Plantago australis is our only native species of

Plantago and can be recognized by the stiff, erect

corolla-lobes that are persistent and form a "beak"

on the apex of the fruit. The corolla-lobes appear

to be open for a very short period during anthesis

and are otherwise erect and imbricate. As with

other species, the name Llanten is commonly used

for these plants. Rahn distinguishes a number of

subspecies; most of our material would appear to

belong to ssp. hirtella (H.B.K.) Rahn.

88

FIELDIANA: BOTANY

FIG. 19. Plantaginaceae: Plantago australis. A, Flower viewed from above; B, a small plant; C, seeds (1-mm

scale); D, fruit with bracts and perianth (5-mm scale); E, fruit (same scale as D); F, a large plant.

BURGER: FLORA COSTARICENSIS

Plantago lanceolata L., Sp. pi. 113. 1753.

Herbs, annual or becoming perennial, aerial stem very

short with congested internodes, secondary roots nu-

merous. Leaves erect or spreading laterally, petiole weakly

differentiated from the lamina, clasping the stem at the

base; lamina 4-28 cm long, 0.5-4 cm broad, lanceolate

to oblong-lanceolate, entire or with small teeth along the

margin, glabrous or puberulent, the veins usually strong-

ly impressed above. Inflorescences 1 5-60 (80) cm long,

peduncle longitudinally ridged, spike 1.5-8 cm long,

bracts 2.3-5 mm long, triangular. Flowers with sepals

2.4-3.5 mm long, the 2 anterior (abaxial) sepals almost

completely united, posterior sepals ovate, corolla-lobes

1.8-2.8 mm long, thin, translucent, reflexed; anthers ca.

1.8 mm long; ovary with only 2 ovules. Fruit ca. 5 mm

long, not covered by the persisting corolla-lobes; seeds

2.3-3 mm long, inner face concave.

Plantago lanceolata has been reported on the

slopes of Volcan Irazu and Volcan Turrialba

(Standley, Publ. Field Mus. Nat. Hist., Bot. Ser.

18: 1264, 1938), but we have seen no specimens

from Costa Rica or elsewhere in Central America.

The species is now cosmopolitan and is natural-

ized in many parts of the Andes; it can be a trou-

blesome weed of lawns and grassland.

Plantago major L., Sp. pi. 1 12. 1753.

Herbs, perennial from a short, thick rootstock, inter-

nodes tightly congested and hidden by the leaf-bases.

Leaves in basal rosettes, flat on the ground or erect,

petioles 2-1 5 cm long, expanded at the base and clasping

the stem; 10-30 (50) cm long 3-12 (18) cm broad, ovate

to elliptic-ovate or broadly elliptic, obtuse at the apex,

the larger laminae abruptly narrowed to the base and

obtuse to truncate, margins entire or irregularly short-

dentate, drying chartaceous, glabrous or with slender

septate translucent hairs 0.3-1 mm long, venation pal-

mate with 3-7 (1 1) primary veins. Inflorescences spicate,

1 5-35 (50) cm long, flowering portion 4-28 cm long and

6-8 mm thick (dry), peduncle puberulent in early stages

but becoming glabrous, flowers closely congested dis-

tally, bracts 2-3 mm long. Flowers with sepals 1.2-2

mm long, broadly ovate to obovate, with thin margins,

glabrous or minutely puberulent along the midvein abax-

ially, corolla-lobes 0.5-1 mm long, narrowly triangular,

obtuse or acute, rotate, not persisting as a covering on

the fruit. Fruit 2-3.8 mm long, ovoid, usually with 5-

many brown seeds; seeds ca. 1 mm long and 0.6 mm

thick, variable in shape but mostly oblong.

A cosmopolitan weed of open sunny sites such

as recently cleared land, roadsides, and stream

edges. It is naturalized in the Central Highlands

between 500 and 1 500 m elevation and appears

to flower throughout the year in Costa Rica. The

plants have been used medicinally, fresh or ground

as poultices; they are known by the name Llanten.

The broad leaves and very short corolla-lobes are

distinctive.

90

FIELDIANA: BOTANY

Index

The index includes all accepted names (in Roman type), synonyms (italics), common English names

(Roman), and vernacular names (italics). New species are in boldface, and the page numbers of illus-

trations are in boldface. Hyphenated words and multiple words are alphabetized by letter.

Acanthaceae 1

Acanthoideae 2

Acanthus 23

mollis 2, 23

montanus 23

Adhatoda carthaginensis 52

umbrosa 50

Amasonia integerrima 33

Aphelandra 23

araoldii 30

aurantiaca 3, 24

aurantiaca var. aurantiaca 3, 24

aurantiaca var. stenophylla 3, 25

campanensis 3, 25

deppeana 28

dolichantha 3, 25

golfodulcensis 4, 26

haenkeana 28

leonardii 4, 26

lingua-bovis 4, 27

pectinata 28

pulcherrima 28

repanda 25

scabra 3, 27

seiberti 4, 28

sinclairiana 4, 26, 28

storkii 4, 29

tonduzii 5, 29

tridentata 5, 30

Aphragmia 75

inundata 72

Arrhostoxylum achimeniflorwn 76

stemonacanthoides 79

Asystasia 30

gangetica 5, 3 1

Barleria 3 1

discolor 3 1

m icans 5, 31

pyramidata 32

Beloperone 48

brenesii 5 1

guttata 5 1

urophylla 6 1

variegata 53

medium 32

brownei 5, 32

brownei f. puberulum 32

costaricense 5, 32

dariense 32

pyramidatum 32

Bravaisia 33

integerrima 6, 33

Buceragenia 33

glandulosa 6, 34

Carlowrightia 34

arizonica 34

costaricana 34

Chaetochlamys 35

panamensis 35

Chaetothylax 35

leucanthus 6, 35

Chamaeranthemum 36

durandii 6, 36

tonduzii 36

cultivated genera 2

Dianthera candelariae 52

comata 53

Diapedium trifurcatum 39

Dicliptera 37

imbricata 6, 37

iopus 6, 38

pallida 7, 38

podocephala 7, 39

skutchii 7, 39

trifurca 7, 39

trifurcata 39

unguiculata 7, 40

Dipteracanthus nudiflorus 77

Dyschoriste 40

valeriana 7, 40

Ecbolium refractifolium 58

trichotomum 60

umbrosum 50

Elytraria 4 1

imbricata 7, 4 1

Eranthemum atropurpureum 68

cuspidatum 68

praecox 69

Glockeria blepharorhachis 43

sessilifolia 44

stricta 44

ventricosa 45

Graptophyllum 4 1

hortense 42

pictum 8, 42

Habracanthus 42, 70

silvaticus 8, 42

Hansteinia 43

blepharorhachis 8, 43

gracilis 44

sessilifolia 8, 44

stricta 8, 44

ventricosa 8, 45

Hemisandra aurantiaca 24

Henrya 45

scorpioides 9, 45

Herpetacanthus 46

panamensis 9, 46, 47

Hexacentris mysorensis 86

Hygrophila 47

conferta 47

costata 9, 47

Hygrophila guianensis 47

Hypoestes 47

phyllostachya 9, 47

Jacobinia 48

aurea 50

crenata 54

spicigera 59

tinctoria 59

umbrosa 50

Justicia 48

angustibracteata 9, 50

asymmetrica 60

aurea 9, 50

brandegeana 11, 51

brenesii 10, 51

candelariae 10, 51

carthaginensis 10, 52

chamaephyton 10, 53

comata 10, 53

costaricana 10, 53

crenata 11, 54

ephemera 54

gangetica 3 1

graciliflora 59

guttata 51

imbricata 41

macrantha 11, 55

metallica 11, 55

microphylla 50

oerstedii 11, 55

orosiensis 11, 56

parvibracteata 12, 56

pectoralis 12, 57

picta 42

pittieri 12, 57

pseudopolystachia 58

refractifolia 12, 58

refulgens 58

sarapiquensis 12, 58

scabra 28

secunda 61

skutchii 12, 59

spicigera 13, 59

tinctoria 13, 59

tonduzii 13, 60

trichotoma 13, 60

tubiformis 66

umbrosa 50

urophylla 13, 61

valerii 13, 61

Kalbreyeriella 70

Kolobochilus blepharorhachis 43

leiorhachis 1 1

Lepidagathis 83

alopecuroides 83

Llanten 88, 90

BURGER: FLORA COSTARICENSIS

91

Louteridium 62

costaricensis 14, 62

tamaulipense 62

Megaskcpasma 62

erythrochlamys 15, 62

Mendoncia 63

brenesii 14, 63

costaricana 14, 64

gracilis 63, 64

lindavii 14, 63, 64

littoralis 63

retusa 14, 65

tonduzii 14, 65

Mendoncioideae 1, 2

Meyenia erect a 85

Nelsonia 65,66

canescens 15, 66

Nelsoniodiae 1, 2

Odontonema 66

flagellum 66

strictum 66

tubiforme 15, 66

Onchyanthus speciosus 33

Pachystachys 2

Plantaginaceae 87

Plantago 87

australis 88, 89

australis ssp. hirtella 88

ecuadorensis 88

hartwegii 88

hirtella 88

lanceolata 90

major 90

schiedeana 88

sodiroana 88

Poikilacanthus 67

macranthus 15, 67

Pseuderanthemum 67

atropurpureum 15, 68

cordatum 34

cuspidatum 15, 68

pittieri 16, 69

praecox 16, 69

standleyi 16, 69

Razisea 70

breviflora 43

citrina 16, 70

spicata 16, 71

wilburii 16, 71

Rhytiglossa candelariae 51, 52

microphylla 50

Ruellia 72

achimeniflora 76

alopecuroidea 83

ha rhi liana 73

biolleyi 17, 73

blechum 32

caroliniensis 75

cooperi 17, 74

geminiflora 17, 74

golfodulcensis 17, 75

inundata 17, 75

jussieuoides 17, 76

longissima 73

longissima var. glabra 78

malacosperma 76

metallica 18, 77

nudiflora 18, 77

nudiflora var. puberula 77

palustris 18, 77

paniculata 18, 78

pittieri 18, 78

praeclara 18, 79

puberula 77

stemonacanthoides 19, 79

tonduzii 19, 80

tuberosa 75

Ruellia tubiflora 8 1

tubiflora var. hirsuta 19, 8 1

tubiflora var. tetrastichantha 19,

80

Sanchezia 8 1

parvibracteata 8 1

pennellii 81

Scrophulariaceae 1

Sericographis tinctoria 59

Sornia 32, 52

Spathacanthus 8 1

hoffmannii 19, 81

Standleyacanthus costaricanus 46

Streblacanthus 82

longiflorus 82

macrophyllus 82

monospermus 19, 82

Teliostachya 83

alopecuroidea 20, 83

Tetramerium 83

hispidum 84

nervosum 20, 84

scorpio ides 45

Thunbergia 84

alata 20, 85

erecta 65

fragrans 85

grandiflora 86

mysorensis 86

Thunbergioideae 1, 2

Thyrsacanthus flagellum 66

si rictus 66

Trichanthera 86

gigantea 20, 86

Tubiflora 4 1

squamosa 41

Zornia 32, 52

92

FIELDIANA: BOTANY

UNIVERSITY OF ILLINOIS-URBANA

30112028175997