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Full text of "Flora Costaricensis : Family #200, Acanthaceae by L.H. Durkee, Family #201, Plantaginaceae by William Burger"

UNIVERSITY OF 

ILLINOIS LIBRARY 

AT URBANA-CHAMPAIGN 

MATURAL HIST. SURVEY 





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 










114 












36 






























































144 












146 












































































i 




























. 

101 !i 

13 ; 

142 II 

47 .! 









FLORA COSTARICENSIS 

Family #200 Acanthaceae 
Family #201 Plantaginaceae 



Other Fieldiana: Botany Titles Available 
in the Flora Costaricensis Series, William Burger, Editor 



Family #40, Casuarinaceae; Family #41, Piperaceae. By William Burger. 1971; 227 pages, 14 illus. 

Publication No. 1140, $18.25 

Family #42, Chloranthaceae; Family #43, Lacistemaceae; Family #44, Salicaceae; Family #45, Garry- 
aceae; Family #46, Myricaceae; Family #47, Juglandaceae; Family #48, Bataceae; Family #49, Be- 
tulaceae; Family #50, Fagaceae; Family #51, Ulmaceae; Family #52, Moraceae; Family #52a, Can- 
nabaceae; Family #53, Urticaceae. By William Burger, Luis D. Gomez, Donald E. Stone, and John 
G. Furlow. 1977; 291 pages, 30 illus. 

Publication No. 1270, $23.25 

Family #15, Gramineae. By Richard W. Pohl. 1980; 608 pages, 224 illus. 

Publication No. 1313, $45.00 

Family #54, Podostemaceae; Family #55, Proteaceae; Family #56, Olacaceae; Family #57, Opiliaceae; 
Family #58, Loranthaceae; Family #59, Aristolochiaceae; Family #60a, Hydnoraceae; Family #60b, 
Rafflesiaceae; Family #6 1 , Balanophoraceae; Family #62, Polygonaceae; Family #63, Chenopodiaceae; 
Family #64, Amaranthaceae; Family #65, Nyctaginaceae; Family #66, Phytolaccaceae; Family #67, 
Aizoaceae; Family #68, Portulacaceae; Family #69, Basellaceae; Family #70, Caryophyllaceae. By 
William Burger, Job Kuijt, Kerry Barringer, Luis D. Gomez, and Richard Baker. 1983; 255 pages, 42 
illus. 

Publication 1350, $25.50 



Order by publication number, or ask for a free copy of our price list. Address all requests to: 

FIELD MUSEUM OF NATURAL HISTORY 

Division of Publications 
Roosevelt Road at Lake Shore Drive 
Chicago, Illinois 60605-2496, U.S.A. 



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 



1986 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 




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 




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 
3.84.0 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 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 14 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 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 24, 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, 24.5 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, 34 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 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 2.54 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 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, 34 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, 45 mm broad, 3.54 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 24 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. 44.5 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 3840 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, 34 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 46 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 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 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 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 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, 1.44 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, 24 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 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 2.44.5 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, 3.54 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, 44.5 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 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 3.04.5 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 24 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 



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