OF THE

m

VOLUME 67

1980

NUMBER 1

FLORA OF PANAMA

1

BY Robert E. Woodson, Jr. and Robert W. Schery

and Collaborators

Part IX

Family 179. RUBIACEAE— PART 1

John D. Dwyer^

Trees, shrubs or herbs, occasionally vines. Leaves simple, entire, opposite,
occasionally ternate or whorled, the margin regular, usually petiolate; stipules
very variable, frequently deciduous, interpetiolar or intrapetiolar, often connate,
truncate or more usually with teeth or lobes, often connate to the petiole, leaflike
in the Rubieae, the leaves and stipules occasionally united as a whorl of four or

more segments. Infl

typically cymose, usually cymose-paniculate,

small dichasia frequent, the flowers rarely solitary; bracts and bracteoles fre-
quently deciduous. Flowers usually bisexual, regular, usually radially symmet-
rical, epigynous, usually 4-5-merous; hypanthium mostly oblong or rotund in
outline, the calycine cup usually well defined, the teeth or lobes usually present,

occasionally with interdental glands or with glands within the cup, the lobes equal
or unequal, occasionally with one or more lobes foliaceous; corolla usually sal-
verform, rotate or funnel shaped, the lobes usually 4-5, valvate, imbricate or
twisted, occasionally cucullate; stamens usually 4-5, epipetalous or rarely free
of the corolla, the anthers mostly oblong and longitudinally dehiscent, the con-
nective occasionally produced at the apex, the filaments usually slender, rarely
pubescent; ovary inferior, rarely half-inferior, usually surmounted by a conspic-
uous disc, 2(-4-8) loculed, the ovules 1 to many per locule, erect, pendulous or
horizontal, mostly adnate to peltate placentas affixed to the median septum, the
latter occasionally very thin, rarely evanescent; style simple, usually glabrous.

1 Assisted by National Science Foundation Grant DEB 77-04300. (Principal Investigator) W. G.
DArcy.

2 Missouri Botanical Garden, Post Office Box 299, St. Louis, Missouri 63166, and St. Louis
University, 221 North Grand, St. Louis, Missouri 63103.

Part 2 will appear in Vol. 67, no. 2 of the Annals of the Missouri Botanical Garden.

After this paper went to press, several novelties in the Panamanian Rubiaceae were received.
These will be published in a supplement in this journal.

Ann. Missouri Bot. Gard. 67: 1-256. 1980.

0026-6493/80/ 1 -256/$25.75/0

2 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

the Stigmas mostly 2, occasionally capitate. Fruits capsular, baccate or drupa-
ceous, occasionally a syncarp; drupes with 2 (occasionally 4-5) pyrenes, a fibrous
endocarp containing single seeds per carpel; cocci when present dehiscent or
indehiscent, the exocarp fleshy, deciduous or occasionally persistent; capsules
dehiscing septicidally or loculicidally; berries frequently large and often with a
fleshy pulp, the calyx persistent or deciduous, the ovarian disc occasionally con-
spicuous; seeds winged or not, rarely with a tuft of hairs, when many then usually
oriented vertically or horizontally.

The Rubiaceae is the largest family of dicotyledons in Panama with about 425
species in 87 genera; some new species were encountered after this paper went
to press. The family occurs at all altitudes and in all kinds of habitats, the aquatic
one excepted, although it may be argued that Lindcnia rivalis is a semi-aquatic.
Few genera are cultivated horticulturally in Panama, e.g., Ixora. Coffea (coffee)
is grown extensively as a food crop. Very few trees yield timber of commercial
value. Few medicinally important Rubiaceae occur in Panama, Cephaelis ipe-
cacahuanha being noteworthy. One species oi Cinchona, the source of quinine
bark, has been collected in Panama, and then only twice. Cinchona puhescens,
well distributed throughout tropical South America, is native to Panama, but it
has not been exploited commercially in Panama.

The Rubiaceae of the New World have been studied intensively in this century
by only a few workers, in contrast, for example, to the study of African Rubi-
aceae. From World War T to World War II, Dr. Paul Standley dominated the
study of tropical American Rubiaceae as is evident from the literature cited
throughout this paper. Another worker of special note is Dr. Julian Steyermark,
who was co-author with Standley of many New World species as well as authoring
many of his own. Following Standley's death in 1963, Steyermark became the
foremost student of tropical American Rubiaceae. In recent years he has devoted
much of his time to the Rubiaceae of the Guyana Highlands and Venezuela. In
recent years, several revisions or monographs of individual genera, some covering
many countries and others in the form of revisions restricted to Panama, have
appeared, e.g., Bouvardia (Blackwell); Cephaelis (Molina); Crusea (Anderson);
Galium (Dempster); Hamelia (Elias); Hoffmannia (Dwyer); Richardia (Lewis);

Rondel etia (Kirkbride).

The vast majority of Rubiaceae in Panama are trees or shrubs with the genus

Psychotria representing almost one-quarter of the species. Subshrubs or rarely

herbs like Boneria. Diodia. Spermacoce. etc., are particularly difficult to identify

as to species.

Despite its size, species of Psychotria in Panama are, for the most part, easier
to identify than those of the smaller genera, Hoffmannia and Randia. Hoffmannia
is taxonomically difficult, as its stipules are soon deciduous and these rarely
present on herbarium material. Added to this is the fact that the flowers and fruits
show little interspecific variability. Randia, like Hoffmannia except for a few
species, is poorly represented in herbaria and then usually only in fruit.

From a taxonomic viewpoint, the leaf characters which are especially note-
worthy, apart from the general shape, size, and texture of the blade, as well as
the apex and the base, are the number of lateral veins and whether or not these

■980] DWYER— FLORA OF PANAMA (Family 179. /?/<W«fPflf^) 3

unite submarginally to form what may be described as an undulate vein; blades
which are subsessile or sessile tend to be auriculate or cordate at the base.

The stipules of Rubiaceae are of great taxonomic value, although frequently
deciduous. Often it is difficult to decide whether the stipules are free or connate
and this may lead to an arbitrary decision.

There is a wide range of variability in the inflorescences of the Rubiaceae. In
terminal inflorescences, the occasional absence of a peduncle may bring the pri-
mary branches into a sessile position, thus creating the impression of several
independent inflorescences. The capitate inflorescence, while easy to describe in
terms of length or width, usually has a complex arrangement of internal bracts
and bracteoles.

The flowers of the Rubiaceae, despite obvious variabihty in size and texture
of the perianth parts, are rather uniform in general structure. Pollination mech-
anisms in the Rubiaceae are in need of intensive study.

The calyx usually possesses a cup which is occasionally truncate, but more
usually dentate or lobate. The usual 4-5 teeth or lobes are occasionally strongly
unequal, e.g., Rondeletia spp. Occasionally one calycine lobe dominates the
others in size, e.g., Calycophyllum. More than a quarter of the Panamanian
Rubiaceae have glands either within the cup or between the teeth or lobes, a
circumstance which is in need of investigation. A spathaceous calyx is rarely
encountered, e.g., Duroia or Psychotria boquetensis. In Morinda and Appunia,
the hypanthiums of adjacent flowers are connate and this fusion may extend to
the calycine cup.

As for the flowers of the Rubiaceae, the corolla is usually yellow or white.
Faramea is noteworthy as the corolla is usually blue. Occasionally collectors
confuse the color of subtending bracts or calycine lobes with that of the true
corolla.

The corolla varies in length from 2-3 mm to more than 80 mm in Tocoyena
and Posoqueria. Whether the lobes are valvate or imbricate is one of the most
important taxonomic characters in the Rubiaceae at the generic level. Many
species of Psychotria and Cephaelis have the lobes cucullate at the apex. The
lobation of each lobe in Joosia is unique among the Panamanian genera. The
coriaceous texture of the corolla of Pentagonia, Ladenhergia and Sitnira is note-
worthy.

Stamens of the Panamanian Rubiaceae are epipetalous except for Exostcma
and Coutarea. The fact that the anthers of these two genera are basifixed, an
uncommon situation in the Rubiaceae, suggests the possibility that these genera
belong to another family. The anthers are usually oblong, although in the Sper-
macoceae they tend to be round. Some genera have the anthers usually produced
at the apex, e.g., Bertiera, Haiuclia, and Hoffmannia spp. Ferdinandusa pan-
amensis has dimorphic or possibly trimorphic anthers. Isertia has anthers which
are transversely locellate, the endothecium exhibiting a tigriform pattern.

The filaments are usually slender, and are terete or plano-convex. Occasion-
ally they are crassate, e.g., in Chione and Simira. Filaments tend to be glabrous,
although occasional exceptions are encountered, e.g., Alseis has filaments com-
pletely pubescent. The point of filament attachment is usually near the middle of
the corolla tube or above the middle. Attachment position is related to the little

4 ANNALS OF THE MISSOURI BOTANICAL GARDKN [Vol. 67

understood phenomenon of heterostyly in the Rubiaceae. Genera with very elon-
gate corolla tubes tend to have the anthers on very short filaments at the orifice
of the tube, e.g., Ixora spp., Posoqueria, Lindcnia, Tocoyena. In describing
filament attachment position, it is important that observations be made on fully
developed flowers rather than buds, as the corolla tube may expand unevenly as
the bud approaches anthesis. References made by Anderson (1972) in his recent
monograph of Crusea concerning cleistogamy in the genus, as well as protandry
being the rule, are noteworthy. Croat (1979) made several references to pollina-
tion mechanisms.

The dioecious condition is only occasional in Panamanian Rubiaceae: Allhcr-
tia, Amaioua, Diiroia, Randia (some spp.), Stachyanhcna, Borojoa, and pre-
sumably in Nertcra. Frequently the number of flowers in the inflorescence of one
sex may differ greatly from the number characterizing the opposite sex. The
ovarian disc is rarely absent; it is variable in shape, e.g., pulvinate, annular,
mound shaped; it may be lobed or unlobed, glabrous or rarely pubescent. Secre-
tory cells are not evident in sections of the disc, although Anderson (1972) ob-
served nectar in young flowers of Crusea. The ovarian disc may persist in the
fruit, e.g., in Hamelia and Cosmibuena. The stigma or stigmas (stigmatic lobes)
are usually 2, but 3, 4, 5, or more are occasionally encountered. Dressleriopsis
has 8 radially disposed stigmas. Stigmas often adhere in the bud and expand later.

Most ovaries are bicarpellate with numerous ovules attached to "T-shaped'"
or peltate placentas, the latter connate with a relatively thick median septum.

The fruits of the Rubiaceae are much more versatile at the generic level than
are the flowers. They range in length from a few millimeters, e.g., Chimarrhis,
to several centimeters in Posoqueria, Randia, Borojoa, Siniira, Tocoyena, and
Stachyarrhena. Only two genera in Panama have aggregate fruits: Morinda and
Appunia. In some fleshy fruits the flesh is ephemeral, falling away to expose two
pyrenes, e.g., Psychotria and Cephaelis. GeophUa, however, retains the fleshy
portion enclosing the two pyrenes. Most fruits are rotund or oblong. The fruits
of Chiococca, Coutarea, and Declieuxia are strongly laterally compressed. Ga-
lium, Didymaea, and Psychotria acuminata have the didymous pericarps. Oc-
casionally fruits are markedly lenticellate, e.g., Coutarea and Exostema. The
calyx persists in many genera. In Coussarea and often in Faramea it is stiffly
cylindrical resembling a miniature pipe organ. Occasionally the calyx, when de-
ciduous, leaves a conspicuous annular scar, e.g., Condaminea, Amaioua, Ha-
melia, etc. In Hamelia the persistent ovarian disc may be prominent. Two genera
which can be usually identified readily by their fruits are Chionc, with the pericarp
usually ovate-oblong, rounded at the base and tapering at the apex, and Deppea,
being conspicuously turbinate and truncate at the apex. The achenelike alate
fruits of Allenanthus are unique in the Rubiaceae. Rarely are fruits wider than
long, most species of Faramea being an exception, although the common F.
occidentalis has rotund fruits. Elongate fruits tend to be dry; those which are
cigar shaped or elongate-turbinate usually suggest the tribe Cinchoneae, but are

not restricted to this tribe.

Raphides are abundant in all genera of the Rubioideae except Bertiera. Fruits
with deciduous calyx are occasional, e.g., Cosmibuena, Amaioua. Very thick-

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 5

walled fruits are found in a few genera, e.g., Borojoa, Posoqueria, Simira, To-
coy ena. The majority of tribes have fruits with two seeds. In Coussarea and
Faramca (tribe Coussareae) the seeds are solitary. Eight tribes of Panamanian
Rubiaceae: Cinchoneae, Condamineae, Gardenieae, Hamelieae, Mussaendeae,
Naucleeae, Oldenlandieae, and Rondeletieae, have numerous seeds. The seeds
may be embedded in a gelatinous pulp. In addition to 13 genera of the Cinchoneae
(Panama), the following genera have alate seeds: Appimia, Arcytophyllum, Bath-
ysa, occasionally Rondeletia spp., Mitracarpus, Simira, and Uncaria. In Hillia,
the body of the seed has a unipolar tuft of hairs. Capsules may dehisce septici-
dally, e.g., Hillia and Ladenbergia, loculicidally, e.g., Condaminea or Rustia,
or circumscissilly, e.g., Mitracarpus. Whether the capsule dehisces from apex
to base or vice versa is of paramount importance in segregating Cinchona from
Ladenbergia, A few genera have the carpels separate with each carpel (mericarp)
retaining the seed, e.g., Machaonia and Richardia. In Diodia one mericarp de-
hisces while the other does not. The valves of Joosia twist at dehiscence in
conspicuous fashion. The twisting is even more exaggerated in Lindenia,

Illustrations for this treatment were all prepared from material at Missouri
Botanical Garden (MO).

Literature:

Anderson, W, R. 1972. A Monograph of the Genus Crusea (Rubiaceae). Mem.
New York Bot. Card. 22: 1-128.

Blackwell, W. J. 1968. Revision of Bouvardia (Rubiaceae). Ann. Missouri Bot.
Gard. 55: 1-30.

Bremekamp, C. 1966. Remarks on the position, the dehmitation, and the sub-
division of the Rubiaceae. Acta Bot. Neerl. 15: 1-33.

Croat, T. 1979. Flora of Barro Colorado Island. Stanford.

Darwin, S. 1976. The Genus Lindenia (Rubiaceae). J. Arnold Arbor. 57: 426-
449.

DeCandolle, A. 1845. Prodromus Systematis Naturalis Regni Vegetabilis 4: 341-
622. Paris.

Dempster, L. T. 1973. The Fleshy-fruited Galiums (Rubiaceae) of Mexico and
Central America. Brittonia 25: 15.

Duke, J. 1968. Darien Ethnobotanical Dictionary. Battelle Mem. Inst., Colum-
bus, Ohio.

. 1970. Manual Dendrologico Para 1000 Especies Arboreas en la Republica

de Panama (Organizacion de Las Naciones Unidas para Agricultura y la AI-
imentacion), pp. 1-325. Panama.

Dwyer, J. 1968. Borojoa and Tocoyena (Rubiaceae) in Panama. Phytologia 17:
445_449.

. 1969, The genus Hoffmannia (Rubiaceae) in Panama. Ann. Missouri

Bot. Gard. 56: 269-286.

& M. V. Hayden. 1966. Notes on Coussarea (Rubiaceae) especially the

Panamanian species. Ann. Missouri Bot. Gard. 53: 368-374.
Escobar, N. 1972. Flora Toxica de Panama, pp. 1-279. Panama.
Hayden, M. V. 1968. Systematic morphological study of New World rubiaceous

6 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

seeds (Rubioideae sensu Bremekamp). Unpublished Ph.D. Dissertation, St.

Louis University, St. Louis, MO.

& J. D. Dwyer. 1969. Seed morphology in the tribe Morindeae. Bull.

Torrey Bot. Club 96: 704-710.
Johnston, L M. 1949. The Botany of San Jose Island (Gulf of Panama). Sargentia

8: 279-287.
Kirkbride, J., Jr. 1968. A revision of the Panamanian species of Rondeletia

(Rubiaceae). Ann. Missouri Bot. Gard. 55: 372-391.
. 1977. Index to the Rubiaceae by Julian A. Steyermark in the Botany of

the Guayana Highland by B. Maguire and Collaborators. Phytologia 36: 324-

366.
Lewis, W. H. & R. L. Oliver. 1974. Revision of Richardia (Rubiaceae). Brittonia

26: 271-301.

The Pharmacopeia of the United States of America. 18th revision. 1970. Mack

Pub. Co., Easton, Pennsylvania.

Pfl

156.

M

Panama), pp. 1-468. London.

Standley, P. 1916. Tardavel, a valid generic name to replace Borreria. Contr,

U.S. Natl. Herb. 18: 122.
. 1916. New species of Rubiaceae of several genera chiefly from Panama.

Contr. U.S. Natl. Herb. 18: 133-142.

. 1918. Rubiaceae, Part I. N. Am. Flora 32: 1-86.

. 1928. Flora of the Panama Canal Zone — Rubiaceae. Contr. U.S. Natl.

Herb. 27: 351-365.

. 1930. Rubiaceae of Colombia. Field Mus. Nat. Hist., Bot. Ser. 7: 1-75.

. 1932. Rubiaceae, Part II. N. Am. Flora 32: 87-158.

. 1933. Flora of Barro Colorado Island, Panama. Contr. Arnold Arbor. 5:

148-150.

1934a. Rubiaceae, Part III. N. Am. Flora 32: 159-228.
1934b. Rubiaceae, Part IV. N. Am. Flora 32: 229-300.

Mu

18(4): 1264-1380.

& L. O. Williams. 1975. Rubiaceae in Flora of Guatemala, Fieldiana:

Botany, Vol. 24, Part XI, nos. 1-3: 1-274.
Steyermark, J. A. 1965. Rubiaceae, Part VI. in B. Maguire and Collaborators,

The Botany of the Guayana Highland. Mem. New York Bot. Gard. 12: 178-285.
. 1967. Rubiaceae, Part VII. in B. Maguire and Collaborators, The Botany

of the Guayana Highland. Mem. New York Bot. Gard. 17: 230-436.

. 1972. Rubiaceae, Part IX. in B. Maguire and Collaborators, The Botany

of the Guayana Highland. Mem. New York Bot. Gard. 23: 227-832.

. 1974. Flora de Venezuela — Rubiaceae, primera parte, 9: 1-593; secunda

parte, 9: 603-1101; tercera parte, 9: 1111-2070. Caracas.
Verdcourt, B. 1958. Remarks on the classification of the Rubiaceae. Bull. Jard.
Bot. Belg. 28: 209-290.

1980] DVVYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 7

KEYS TO THE TRIBES AND GENERA OF THE RUBIACEAE

a. Ovules more than I in each cell (subfamily Cinchoneae).
b. Fruits fleshy, berrylike.

c. Corolla lobes valvate in bud 1. Mussaendeae

cc. Corolla lobes imbricate or contorted in bud.

d. Corolla lobes imbricate in bud 2. Hamelieae

dd. Corolla lobes contorted in bud 3. Gardenieae

bb. Fruits dry, capsular.

e. Flowers arranged in dense, compact, spherical heads 8. Naucleeae

ee. Flowers variously disposed, never in spherical heads.

f. Seeds winged or appendaged, vertically imbricate 4. Cinchoneae

ff. Seeds not winged, or if winged, horizontal.

g. Corolla lobes imbricate or contorted in bud 6. Rondeletieae

gg. Corolla lobes valvate in bud.

h. Seeds horizontal, usually very numerous; stipules entire or

bifid; trees or shrubs 5. Condamineae

hh. Seeds imbricate, vertical, usually few; stipules often setose-

laciniate: subshrubs or herbs 7. Oldenlandieae

aa. Ovules solitary in the cells (subfamily Rubioideae. except Naucleeae).

i. Seeds pendulous, the radicle superior; shrubs, trees, or woody vines.

j. Flowers in compact globose heads 8. Naucleeae

jy. Flowers never in globose heads.

k. Stamens inserted in the throat of the corolla 9. Guettardeae

kk. Stamens inserted at the base of the corolla tube or on a disc .

ii. Seeds ascending, the radicle inferior; plants woody or herbaceous.

10. Chiococceae

I. Corolla lobes contorted in bud; shrubs or trees 11. I:

1. Corolla lobes valvate in bud: habit various.

xoreae

m. Ovules basally attached; mostly shrubs or trees.

n. Ovary 7-8 celled 14. Dressleriopsideae

nn. Ovary 1-2 celled.

o. Ovary 1-ceIled or 2-celled with a thin septum: fruits 1-seed-

ed 12. Coussareae

oo. Ovary 2-celled, or the cells rarely more numerous, the septum

thick; fruits 2 or more seeded.

p. Stamens usually inserted in the throat of the corolla;

flowers perfect 13. Psychotrieae

pp. Stamens usually inserted at the base of the corolla tube;

flowers often unisexual 15. Anthospermeae

mm. Ovules lateral, attached to the septum of the cell; habit various.

q. Stipules neither foliaceous nor setiferous; trees or large shrubs;

flowers congested by the hypanthium to form a dense head; fruit a

syncarp, the base of the hypanthiums united 16. Morindeae

qq. Stipules either foliaceous or setiferous; herbs or small shrubs; flow-
ers never congested by the hypanthium: carpels free, not united.

r. Stipules setiferous; fruit not a syncarp 17. Spermacoceae

rr. Stipules foliaceous, similar to the leaves 18. Rubieae

1. MUSSAENDEAE
a. Leaves opposite.

b. Leaves with intervenal areas lineolate.

c. Petioles not tumescent or swollen.

d. Leaves large, !4-l m long, often pinnatifid-lobed; calycine cup with rows

of conspicuous glands within 65. Pentagonia

dd. Leaves small, less than 'i m long, not pinnatifid-lobed: calycine cup with-
out rows of glands within 79. Sominera

cc. Petioles tumescent, swollen.

bb. Leaves with intervenal areas not lineolate 47. Hippotis

e. Inflorescences terminal.

f. Plants drying black: leaves and flowers succulent: inflorescence capitate

^ 77. SchraJcra

8 ANNALS or THE MISSOUR[ BOTANICAL GARDLN [Vol. 67

aa.

ff. Plants not drying black; leaves chartaceous, occasionally stiffly so: inflo-
rescence spreading (except subcapitate in Amphidaysa).
g. Stipules with conspicuous fimbriate processes 5. Amphidaysa

Stipules without fimbriate processes.

h. Inflorescences spikelike; shrubs 42. Gonzala^unia

hh. Inflorescences cymose-corymbose or cymose paniculate; trees.

i. Corolla 3 or more cm long; anthers transversely locellate

49. Iscrtia

ii. Corolla less than 1.5 cm long; anthers not transversely lo-

llate ^9- Raritehe

ce

ee. Inflorescences axillary.

j. Plants prostrate; fruits bright china blue 21. Covcocypselum

jj. Plants erect or climbing; fruits not bright china blue 76. Sahicca

Leaves alternate 31. Diclymochlamys

2. HAMELIEAE

a. Ovary 4-5 celled; inflorescences regularly terminal; stamens with anther connectives reg-
ularly produced at the apex.

b. Corolla lobes contorted in bud; raphides absent from leaves 10. Bertiera

bb. Corolla lobes imbricate in bud; raphides present in leaves 44. Hamelia

aa. Ovary 2-3 celled; inflorescences always axillary; stamens with the anther connective not
produced at the apex (few species of Hoffmannia excepted).

c. Ovary 2 celled; calyx with lobes ca. 8 mm long; corolla lobes valvate

87. Xcrococciis

cc. Ovary 2-3 celled; calyx with lobes less than 5 mm; corolla lobes imbricate

48. Hoffmannia

3. GARDENIEAE

a. Plants armed with spines on stems and short shoots 69. Randia

aa. Plants unarmed.

b. Inflorescences not spikelike; fruits not gourdlike nor on an elongate curved pedicel.

c. Flowers perfect.

d. Inflorescences with flowers several to numerous, flowers disposed in a

candelabra array.

e. Flower buds curved at the apex; seeds with testa cells more than

twice as long as wide 66. Posoqueria

ee. Flower buds not curved at the apex; seeds with testa cells less than

twice as long as wide 83. Tocoyena

dd. Inflorescences with flowers few or solitary, not disposed in a candelabra

array.

f. Leaves drying black 40. Gcnipa

ff. Leaves not drying black.

cc. Flowers unisexual.

g. Fruits oblong; terminal stipules forming a conic cap.

h. Pistillate flowers capitate or cymose; fruits not densely hirsute

4. Aniaioua

hh. Pistillate flowers usually solitary; fruits densely hirsute 34. Duroia

gg. Fruits rotund; terminal stipules not forming a conic cap.

i. Fruits subtended by several persistent bracts, as large as a child's

hand, superficially rough 11- Borojoa

ii. Fruits not subtended by several persistent bracts, to 2.5 cm wide,

superficially smooth ^ 1. Alihertia

bb. Staminate inflorescences spikelike; fruit a gourdlike capsule on an elongate curved

pedicel 81. Stacliyarrhcna

4. CINCHONHAE

a. Vines 57. Manettia

aa. Shrubs or trees.

b. Stamens with anthers dimorphic or trimorphic 38. Fcrdinandusa

bb. Stamens with anthers all alike.

19801 DWYHR— FLORA OF- PANAMA (Family 179. Ruhiaccne) 9

c. Calyx with one lobe expanded as a blade.

d. Inflorescence cymose or corymbiform; stamens equal and exserted

14. Calycopliyllum

dd. Inflorescence spicate; stamens unequal, not exserted 86. Wittmavkantluis

cc. Calyx with all lobes similar.

e. Plants epiphytic; leaves fleshy, the lateral veins often evanescent.

f. Flowers pink or red, the corolla inflated 70. Rnvnia

ff. Flowers while or yellow.

g. Seeds winged terminally 24. Cosnuhncna

Seeds with a tuft of hairs at one pole 46. Hillia

ee. Plants terrestrial; leaves not fleshy, mostly drying papery to coriaceous,

veins evident to the margin,
h. Inflorescences spikelike: stamens with filaments villose, attached at

the base of the corolla tube 3. Alseis

hh. Inflorescences various but not spikelike; stamens attached above the

base of the corolla tube (except Exostema and Coutarea).
i. Corolla lobes valvate in bud.

j. Flowers 4-merous; capsule rotund 13. Boiivanlia

jj. Flowers 5-merous; capsule oblong.

k. Capsule splitting from below toward the apex

20. Cinchona

kk. Capsule splitting from above toward the base.

I, Corolla lobes split at the apex; fruits forming 4
coiled valves 51. Joosia

II. Corolla lobes not lobed at the apex; fruits not

forming coiled valves.

m. Capsule loculicidal 56. Macrocncnuiin

mm. Capsule septicidal 52. Ladenhcr^ia

ii. Corolla lobes imbricate or contorted in bud.

n. Stamens conspicuously exserted; corolla not inflated;

fruits neither compressed nor lenticellate 36. Exostema

nn. Stamens not conspicuously exserted; corolla inflated;

fruits compressed, strongly lenticellate 26. Coutarea

5. CONDAMINEAE
a. Leaves glandular-punctate, the dots pellucid; anthers opening by terminal pores

aa. Leaves not glandular-punctate; anthers longitudinally dehiscent.

75. Rustia

b. Flowers ca. 3 mm long; inflorescences axillary 16. Chimarrhias

bb. Flowers more than 3 mm long; inflorescences terminal.

c. Calycine lobes unequal, one expanded into a large colored limb ____ 65. Pogonopus
cc. Calycine lobes equal or nearly so 23. Comlaminea

6. RONDELETIEAE

a. Plants woody.

b. Corolla lobes imbricate in the bud.

c. Calycine lobes unequal, often one expanded into a foliose blade.

d. Calyx with foliose blade a flaming red color 85. Warsccwiczia

dd. Calyx with foliose blade white 73. Rondeletia (some spp.)

cc. Calycine lobes equal or nearly so.

e. Fruits large, nutlike, dehiscing septicidally; seeds winged: wood on ex-
posure to air turning a purple red; filaments geniculate at point of attach-

ix\QX\\, '^^- Simira

ee. Fruits small, dehiscing loculicidally or septicidally; seeds not winged; wood

on exposure to air not turning red; filaments not geniculate at point of

attachment.

f. Calycine lobes often unequal: capsule with valves not woody, not bifid

at the apex 73. Rondeletia

ff. Calycine lobes usually equal; capsule with valves woody, bifid at the

apex 9. Bathysa

10 ANNALS OF THL MISSOURI BOTANICAL GARDEN [Vol. 67

aa

bb. Corolla lobes contorted in bud.

g. Trees, terrestrial; corolla very short, the tube shorter than the lobes; fruits with

valves not coiled at maturity.

h. Corolla glabrous within, 4-merous 29. Dcppea

hh. Corolla villous within, 5-merous 35. Elaea^nla

. Shrubs, growing in water or on banks of streams; corolla elongate, the tube at

least 5 times the length of the lobes, fruits with valves coiled at maturity

54. Lindenia

. Plants herbaceous 53. Limnosipanca

7. OLDENLANDIEAE

a. Plants herbaceous, often prostrate or subprostrate, not cricoid in habit; seeds angular or

winged 62. Oldcnlandia

aa. Plants subshrubs, cricoid in habit, usually upright; seeds carinate or plano-convex

8. Anytophyllum

8. NAUCLEEAE

Vine with recurved spines and inflorescences axillary and globose 84. Uncaria

9. GUETTARDEAE

a. Plants armed with rigid spines on twigs and stems.

b. Corolla lobes imbricate in bud; fruits dry, splitting into 2 indehiscent mericarps or

cocci 55. Machaonia

bb. Corolla lobes valvate in bud; fruits fleshy, not splitting into 2 indehiscent mericarps

19. Chomelia

aa. Plants unarmed.

c. Corolla lobes imbricate in bud; leaves with intervenal areas not lineolate-striate.

d. Fruits dry, separating into 2 indehiscent mericarps or cocci; flowers not secundly

disposed 55. Machaonia

dd. Fruits fleshy; flowers secundly disposed.

e. Fruits covered with a fine feltlike pubescence 43. Guettarda

. Fruits glabrate 6. Antirhea

cc. Corolla lobes valvate in bud; leaves of some species with intervenal areas Imeolate-

striate 19. Chomelia

10. CHIOCOCCEAE

a. Flowers 4-merous; fruits winged 2. Allcnanthus

aa. Flowers 5-merous; fruits not winged.

b. Corolla lobes imbricate in bud; fruits not strongly compressed; shrubs (Panama)

18. Chione

bb. Corolla lobes valvate in bud; fruits strongly compressed; trees, shrubs, or woody

vines -._-_ 17. Chiococca

II. IXOREAE

a. Bractlets connate, calyx-like or involucral; inflorescences axillary; cultivated as a tree

crop 22. Coffea

aa. Bractlets distinct, not calyx-like; inflorescences terminal; some species cultivated as or-
namentals 50. Ixora

12. COUSSAREAE

a. Ovules connate, borne on a common basal column; seeds vertical; flowers usually white

25. Coussarea

aa. Ovules separate in a one-celled ovary; seeds horizontal; flowers often blue 37. Faramea

13. PSYCHOTRIEAE

a. Plants prostrate, woody herbs; leaves cordate 41. Gcophila

aa. Plants erect subshrubs, shrubs or trees (except Declieuxia. a low woody herb [Panamal);

leaves various at the base, not cordate.

b. Lobes of calyx, if present, subequal, not polymorphic at the apex.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 1 1

c. Inflorescence an involucrate head, the heads solitary, sessile or pedunculate, or

with 3 heads arising from a common peduncle 15. Cephaelis

cc. Inflorescence bracts not congested into an involucre, flowers not congested into

a head.

d. Herbs: fruits laterally compressed 28. DecVieuxia

dd. Trees, shrubs, or subshrubs, rarely epiphytes; fruits terete in cross section.

e. Seeds with an incurved ventral surface: stipules pectinate

74. Ruilgea

ee. Seeds not with an incurved ventral surface; stipules rarely pectinate,
if so, membranous.

f. Corolla tube short, to 0.7 cm long (P. chiapensis and P. ostaurea
excepted), not swollen at the base; flowers white, green, or yel-
low; stipules of twiglets deciduous or persistent 68. Psychotria

ff. Corolla tube elongate, more than 0.7 mm long, usually swollen
at the base: flowers regularly orange and purple; stipules of twig-
lets regularly persistent 64. PalUourea

bb. Lobes of calyx unequal, polymorphic at the apex 59. Montamans

14. DRESSLERIOPSIDEAE
Ovary and fruit 7-8-carpellate 33. Dresslcriopsis

15. ANTHOSPERMEAE

Plants forming prostrate mats

6 1 . Nertera

16. MORINDEAE

a. Plants usually drying black, or at least fruit drying black; fruits succulent, 1 cm or more in

diam. 60. Morinda

aa. Plants not drying black; fruits dry, less than 1 cm in diam 7. Appunia

17. SPERMACOCEAE

a. Fruits with circumscissile dehiscence 58. Mitrcuarpas

aa. Fruits longitudinally dehiscent or separating into open or closed valves or cocci, lacking

circumscissile dehiscence.

b. Cocci 3-5(-6) 72. RUhardia

bb. Cocci 2.

c. Stipules with sheath with more than 3 setae; plants not fetid.

d. Cocci indehiscent.

e. Calyx lobate; cocci separating from a central axis or carpophore

27. Crusea

ee. Calyx dentate; cocci not separating from a central axis or

carpophore 32. Diodia

dd. Cocci dehiscent, often only I of the 2 opening.

f. Cocci opening only at the base; flowers axillary 45. Hemidiodla

ff. Cocci opening at the apex; flowers axillary or terminal.

g. Fruits with cells similar, both dehiscent 12. Borreria

Fruits with cells dissimilar, only one opening 80. Spermacoce

cc. Stipules with sheath with only 3 setae; plants fetid 82. Tahagoa

18. RUBIEAE

a. Flowers with the hypanthium immediately subtended by an involucre of leaflike bracts

71. Relhunium

aa. Flowers with the hypanthium not immediately subtended by an involucre of leaflike bracts.

b. Leaves opposite 30. Didymaea

bb. Leaves verticillate 39. Galium

12

ANNALS OF THE MISSOURI BOTANICAL GARDEN

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1980] DWYER— FLORA OF PANAMA (Family 179. W/z/j/rtccrtc) 19

1. ALIBERTIA

Alibertia A. Rich, in DC, Prodr. 4: 443. 1830. type: A. edulis (L. C. Rich.) A
Rich, in DC.

Cordiera A. Rich, in DC, Prodr. 4: 445. 1830. type: C. triflora DC.

Shrubs or trees. Leaves usually coriaceous, petiolate; stipules connate at the
base. Inflorescences terminal, the staminate flowers several, fasciculate, the pis-
tillate flowers usually solitary. Flowers dioecious, usually sessile; hypanthium
usually globose; calyx short or tubular, truncate or dentate; corolla tube salver-
form, stiffly carnose, the lobes contorted, imbricate; stamens 4-8, the anthers
linear; ovary 2-8-celled, the ovules few to numerous. Fruits baccate, 2-8-celled,
the placentas pulpy, the seeds several, large, compressed.

Alibertia is found throughout tropical America and the West Indies. It has
about 35 species.

a. Leaves with secondary veins 6-15; corolla tube 1-3 cni long at anthesis, 0.5-0.8 cm wide;

anthers 10-13 mm long _._ 1. A. edulis

aa. Leaves with secondary veins ca. 6-19: corolla tube less than 0.8 cm long at anthesis, ca.

0.25 cm wide; anthers ca. 3 mm long 2. A. garapativa

1. Alibertia edulis (A. Rich.) A. Rich., DC, Prodr. 4: 443. 1830.— Fig. 1.

Genipa edulis A. Rich., Acles Soc. Hist. Nat. Paris I; 107. 1792. type: not seen.

Gardenia edulis (A. Rich.) Poir. in Lam., Encyc. Meth. Bot. Suppl. 2: 708. 1812.

Garapatico edulis (A. Rich.) Karst., Fl. Colombia 1: 57. 1859.

Cordiera edulis (A, Rich.) Kuntze, Rev. Gen. PI. 279. 1891.

Sahicea edulis (A. Rich.) Seem., B. D. Jackson, Index Kew. 772. 1895.

Alibertia longistipulata Riley, Kew Bull. 1927: 122. type: Panama, Riley 145 (K, not seen).

Alibertia panaiuensis Riley, Kew Bull. 1927: 123. type: Panama, Rdey III (K).

Shrubs or small trees, presumably dioecious, the branchlets terete or often
ultimately angular, smooth, glabrous, stiffly ascending. Leaves narrowly oblong,
falcately oblong, oblong, ovate or rarely obovate-oblong, 6-20 cm long, 2-8 cm
wide, deltoid at the apex, acuminate, the acumen to 0.1 cm long, usually acute,
deltoid to obtuse at the base, the costa plane to prominulous above, prominent
beneath, to 0.2 cm wide, glabrous, the lateral veins 6-15, arcuate, to 2 cm apart,
thin-coriaceous, lustrous, glabrous; petioles 0.5-1.0 cm long, rarely absent, gla-
brous; stipules connate or free, deltoid, narrowly deltoid or obovate-oblong, ca.
1.5(-2.0) cm long, 0.4-0.6 cm wide, attenuate-acuminate, the acumen often longer
than the corpus, scarious, striate-venose, glabrous. Inflorescences of 1 to several
terminal flowers subtended by broadly ovate bracts shorter than or exceeding the
hypanthium in length, the buds with the apex of the corolla acute and dilated.

F

Flowers sessile, unisexual, the pistillate flowers solitary, the staminate flowers
in a cluster of about 8; hypanthium and calycine cup ca. 0.5 cm long, carnose,
glabrous, the teeth 5(-4), deltoid to linear, to 1.8 mm long, acute to obtuse;
corolla white, the tube 1-3 cm long, glabrous or villosulose outside, subcoria-
ceous, the lobes 4-5, imbricate, oblong, to 2 cm long, often exceeding the tube;
stamens usually (4-)5(-8), the anthers narrowly oblong, 5-13 mm long, ca. 13

mm long in male flowers, ca. 5 mm long in female flowers, 0.8-1.0 mm wide,
often minutely apiculate, dorsifixed, the filaments short, ca. 0.5 cm long; ovarian

20

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 67

Figure L Alibertia eclulis (A. Rich.) A. Rich.— A. Flowering branch (xVi). [After Uladik
49)j_B. Flower buds {xi/2). [After L^^iiv.v et al. 1866.]— C. Fruit (xVi). [After Lao c6 Holdridge
244.]

disc doughnut-shaped, occasionally scalloped, 1.0-1.8 mm long, 0.75-1.50 mm
wide, the style expanding toward the apex, the stigmas 3-4, crassate, oblong, to
1.8 mm long, the ovary 4-celled. Fruits terminal, sessile, solitary, rotund to com-
pressed-rotund, to 3 cm in diam., the persistent calyx coroniform and cylindrical,
to 0.5 cm long, rarely to 1 cm long, ligneous, smooth, glabrous; seeds numerous
in a slimy pulp, suborbicular, ca. 0.5 cm in diam., striate.

Alibertia eclulis is one of the most common shrubs in Panama. It ranges from
Mexico to the Amazon Basin, although Steyermark does not include it in his
Flora de Venezuela. It occurs in the West Indies, The fruits are scarcely edible.
The names 'Trompo'' and ^Trompito" refer to the fact that tops are made from
the fruits. 'Xargartillo''; '^Madrono''; ''Madrono de Comer'\ ^'Trompo
'Trompito^^ ''Wild Guava.

*t ^

^ ^

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhuucac) 21

BOCAS DEL TORo: Bocas, Lazor ct al. 2415 (MO). Columbus Island, Wcdel 32 (MO). Water
Valley, Wedel 798 (MO). Chiriqui Lagoon, Wedel 131 1 (MO). Old Bank Island, Wcdel 2009 (MO).
Isia Colon, Wedel 2474 (MO). Water Valley, Wedel 2672 (MO). Johns Creek, Wedel 2773 (MO).
CANAL zone: Juan Mina, Barflett & Lasser 1552 (MO). Cruces Trail and Madden Dam Road, Bartlett
& Lasser 16342 (MO). Juan Mina, Bartlert & Lasser 16563 (MO). Cocoli, Croat 15145 (MO). Pan-
american Highway near La Chorrera, DArcy 9439 (MO). Fort Kobbe, Duke 3841, 4201, 4725 (all
MO). Albrook, Dwycr 6746 (MO); Dwyer & Rohyns 119 (MO). Empire, Harden 58 (MO). Fort
Sherman, Hayden 97, 107 (both MO). 6 km W of Gamboa, Nee 7208 (MO). Road C-29, 6 km E of
Gamboa, Nee 8970 (MO). Howard Air Force Base, Oliver <i MacBryde 1894 (GH, MO, US). Balboa,
Standley 26050 (MO). Farfan, Stern & Chambers 153 (MO). Rio (ilocoli. Road K-9, Stern et al. 38
(MO). Rio Indio, Stexermark & Allen 17439 (MO). Curundu, Tyson 1316 (MO). Chiva-Chiva Trail,
Tyson 1357 (MO). Miraflores, White 129, 138 (both MO). Cocoli, Wilbur et al. 12864 (MO), barro
COLORADO island: Aviles 23 (MO); Croat 4226, 5167, 5358, 5574, 5724, 6006, 6128, 6142, 6328, 6429,
6878, 7390, 7454, 9249, 9512, 15263, 15397 (all MO): Dnyer 1434, 1438. 8467 (all MO); Ebinger 261
(MO); Hayden 3 (MO); HIadik 49 (MO); Shattuck 147, 643 (both F, MO), 896, 967 (both MO);
Wetmore & Abbe 78 (MO), 78 A (F, MO); Woodson & Sehery 977 (MO); Zetek 4295 (F, MO).
chiriqui: Boquete, Davidson 829 (MO); Dnyer & Hayden 7625 (COL, MO, UC, VEN); Kirkbride
93 (MO); Stern et al. 1200 (MO). Puerto Armuelles, Woodson & Sehery 847 (MO), cocle: Penonome,
Dwyer 2012 (MO). 2 mi E of Ri'o Hondo, Gentry 2919 (MO). La Pintada, Hunter & Allen 519 (MO).
El Valle, Lee 12 (MO), colon: Salud, Lao & Holdridge 244 (MO). Nuevo Chagres, Lewis et al.
1866 (GH, MO), darien: Mouth of Rio Yape, Allen 340 (MO). Quebrada Venado and Peje Swamp,
Bristan 1048 (MO). Rio Canclones and Rio Chucunaque, Duke 5118 (MO). Cerro Pirri, Duke 6584
(MO). Perrecenico River, Duke & Bristan 236 (MO). Rio Tuira, Duke & Kirkbride 14056 (MO).
Yaviza, Gentry & Mori 13504 (MO), Between Cana and Boca de Cupe, Stern et al. 613 (MO), i os
SANTOS: 1-2 mi W of Candelaria, Duke A-12433 (MO). Los Asientos, Wandehake 31 (MO). Panama:
Nuevo Emperador, Carrera I (MO). Cerro Campana, Castillo 22 (MO). Rio Pita, Duke 4770 (MO).
EI Llano, Duke 5520, 5824 (both MO), Isia del Rey, Duke 9518, 10411 (both MO). Coronado Beach,
Duke 11817 (MO). Isia de La Bayoneta, Dwxer 1736 (MO). Ri'o Pasiga, Gentry 2277 (MO). San Jose
Island, Johnston 486 (MO). Chiman, Lewis et al 3314 (MO, UC, VEN). Chorrera, Rodriguez 51
(MO). Bella Vista, Standley 25354 (MO). Isia de Espiritu Santo, Tyson 5562 (MO). 5 mi W of Chepo,

Tyson 6725 (MO). Pacora, Woodson et al. 743 (MO), san blas: Eslogandi, Duke 10196 (MO).
VFRAGUAs: Rio Canazas headwaters, Allen 188 (F, MO). Sona, Allen 1073 (MO). 5 mi N of Santiago,

Blum & Tyson 622 (MO). La Mesa, Blum & Tyson 640 (MO). 32 mi W of Santiago, Croat 10732
(MO). San Francisco, Dwyer 1257 (MO). Mouth of Rio Concepcion, Lewis et al. 2821 (COL, DUKE,

MICH, MO, UC). Laguna La Yeguda, Luteyn 1460 (MO). Quebrada just S of Santa Fe, Nee 8035
(MO). 5 mi E of Santiago, Tyson et al. 4272 (MO).

2. Alibertia garapatica (Karst.) Schum. in Mart., Fl. Bras. 6(6): 384. 1889.

Garapatica edulis Karst., Fl. Colomb. 1: 57. 1858, not A. edulis A. Rich, type: not seen.

Shrubs or small trees, to 4 m tall, the twiglets slender, often puberulent, the
nodes usually swollen, the leaf scars large. Leaves oblong or oblong rotund, 6-
15 cm long, 2.3-5.0(-6.0) cm wide, rounded or tapering widely into an obtuse or
deltoid apex, acuminate, the acumen 0.5-2.0 cm long, obtuse to cuneate at the
base, the lateral veins 6-10, thickly chartaceous to subcoriaceous, smooth, gla-
brous to sparsely golden-puberulent; petioles stiff, 0.5-1.0 cm long; stipules free?,
triangular, to 3 mm long, acute, puberulent, usually deciduous. Inflorescences
terminal, sessile or subsessile, the flowers of the female inflorescence ca. 5, oc-
casionally solitary?, those of the male 7-10, radially disposed. Flowers (only
male seen) ca. 1 cm long, the buds acute, the hypanthium ca. 1.5 mm long, the
calycine teeth absent; corolla white, the tube cylindrical, to 8 mm long, to 2.5
mm wide, stiffly carnose, puberulent, the lobes 5, triangular, to 2.5 mm long;
stamens 4(5?), narrowly oblong, ca. 3 mm long. Fruits subsessile or sessile,
solitary, rotund or subrotund, occasionally obovate-rotund, to 2.5 cm long, slight-
ly wider than long, drying black, minutely rugulose, glabrous, the persistent ca-
lycine cup crateriform-cylindrical, to 3 mm long.

22 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

Alihertia garapatica is known from Mexico, Panama and Colombia. The
species should be studied in the field as more than one species may be involved.
Stern et aL 1719 shows 4 very immature fruits in one fructescence terminating a
twiglet, while another twiglet shows 4 flowers in bud which measure about 1.2
cm in length and are not as acute as typical buds of the species. One flower
dissected proved to be male with 4 subfacate sessile stamens and a fifth stamen
which was abnormally plump. The 4 normal stamens measured 0,5 mm in width,
with an acumen to 0.8 mm long. The abnormal stamen measured 0.8 mm in
diameter. The unopened anther sacs of the normal stamens were linear and the
connective laminar as one sees in the stamens of the Himantandraceae. Allen
1616 showed a style 6.2 mm long, which was dilated basally and minutely bifid
at the apex; no ovary was evident. Dressier & Williams 3969 has leaf blades only
4.0-6.5 cm long. Several leaves of Liesner 334 are almost rotund, being 7.0-7.5

even though it seems out of range.

(MO) from Mexico

CHiRiQuf: Quebrada Punta de Piedra, 0-100 m. Liesner 334 (MO), cocle: Aguadulce, McDaniel
& Cooke 14762 (MO), colon: Santa Rita Ridge, E of Transisthmian Highway, 300-500 m, Gentry
6589 (MO). Santa Rita Road, Dressier & Williams 2969 (MO). Santa Rita Road, Mori i& Kallunki
2787 (MO), herrira: Ocu, limestone area, Stern et oL 33583, 33593 (both MO). Panama: Rio Las
Lajas, Allen 1616 (F). Villa Rosario, International Highway, Saldana 21 (MO). Guayabito, road to
Laguna, Smith <ft Smith 3463 (MO), veraguas: Isla de Coiba, Dwyer 1629 (MO).

2. ALLENANTHUS

Allenanthus Standley, Ann. Missouri Bot. Gard. 27: 344. 1940. type: A, eryth-
rocarpus Standley.

Trees. Leaves ovate elliptic, acuminate, the lateral veins few; petioles present;
stipules free, ovate or rotund, cuspidate. Inflorescences terminal, paniculate,
elongate pedunculate; bracts and bracteoles small. Flowers with calycine lobes
4; anthers short. Fritits compressed, drying red, the central seminiferous part
surrounded by a thick wing.

Allenanthus is found in Costa Rica and Panama. It is among the easiest of the
Rubiaceae to recognize as the fruit has the appearance of the achene of Rumex
of the Polygonaceae.

1. Allenanthus erythrocarpus Standley, Ann. Missouri Bot. Gard. 27: 344. 1940.
type; Panama, Allen 1999 (MO, holotype). — Fig. 2.

Chimarrhis decnrrens Steyermark, Ceiba 3: 18-19. 1952. typf: Allen 3566 (F, holotype).

Trees 10(-20) m tall, the branchlets smooth, terete, rubescent, ultimately
piano-compressed, glabrate, the nodes usually 4-5 cm apart. Leaves ovate ellip-
tic, 7-11 cm long, 3.8-5.0 cm wide, deltoid at the apex, acuminate, the acumen
to 1.5 cm long, obtuse and scarcely inequilateral at the base, the costa promi-
nulous above, prominent beneath, the lateral veins ca. 7, arcuate, usually with
one subparallel vein between 2 lateral veins, the intervenal areas mosaic-like,
chartaceous, scarcely discolorous, glabrous except puberulent on the costa and
the veins beneath; petioles slender, to 1.5 cm long; stipules deciduous, free,

19801

DWYER— FLORA OF PANAMA (Family 179. Ruhiaccac)

23

Figure 2. Allcnanrhits crythrocarpus Slandley. — A. Branch with leaves (x.75). — B. Inflores-
cence (X.75). [After Duke 5654,]

widely ovate or compressed rotund, 3-6 mm long, abruptly cuspidate, the cusp
as long as the body. Inflorescences terminal, solitary, paniculate, to 15 cm long,
to 10 cm wide, rounded terminally, the peduncles terete, 4,0-6.5 cm long, pu-
berulent, the branches opposite, the lowermost to 9 cm long, arcuate or strict,
strongly ascending, pubescent, the hairs often present only on part of the pedun-
cle and branches; bracts small, linear-subulate. Flowers with pedicels 1-5 mm
long; hypanthium turbinate, glabrous, to 2 mm long; calycine lobes 4, prominent,

24 ANNALS OF THE MISSOURI BOIANICAI. GAROF^N [Vol. 67

ovate to rotund, 0.5-0.9 mm long, glabrous; corolla tube to 4 mm long, pubescent
outside, sparsely pilose within, the lobes oblong, ca. 1.5 mm long, almost as
wide, rounded apically; anthers oblong, ca. 1 mm long, the filaments slightly
shorter than the anthers. Fruits strongly compressed, ca. 8 mm long, to 6 mm
wide, glabrous, appearing winged, drying red, the central seminiferous part to 2.5
mm long, 3-nerved, surrounded by a thick wing.

AUenanthiis erythrocarpus is known only from Panama and Costa Rica.

cocle: El Valle de Anton, Alh'ii 1999, 2713 (both MO), 3566 (F). Panama: Between El Llano
and Rfo Mamoni, Duke 5654 (MO).

3. ALSEIS

fl^

Infl

spiciform cylindrical, patulous. Flowers protogynous, the hypanthium obconical,
the calyx 5-lobed; corolla urceolate, inflated; stamens 5, exserted, the filaments
inserted at the base of the corolla; style recurved, the ovary 2-loculate, the septum
thin, the ovules numerous. Fruits capsular, narrowly oblong, lignose, dehiscing
from apex to base, 2-locular, the seeds many, narrowly fusiform.

Alsc'is has about 20 species ranging from Mexico through Central America to
Brazil and Peru.

a. Leaves with lateral veins ca. 25; fruits before dehiscence ca. 0.2 cm wide ]. A. hlackiana

aa. Leaves with lateral veins ca. 9; fruits before dehiscence 0.4 cm wide 2. A. claricnensis

1. Alseis blackiana Hemsl., Diag. PI. Nov. 30. 1879. type: Panama, Hayes 665
(K).— Fig. 3.

Trees to 75 ft tall; twigs terete, ultimately angular, the nodes well spaced or
crowded, glabrous. Leaves obovate oblong, to 36 cm long, to 9.5 cm wide, cu-
neate at the apex, often cuspidate, the cusp to 1 cm long, cuneate or rarely
truncate at the base, equilateral, rarely subinequilateral, the costa prominulous
above, prominent beneath, diffusely villose above, the hairs weak, these short
and long, occasionally aggregated in the axils, the secondary veins ca. 25, the
distal veins widely arcuate, the proximal veins perpendicular, chartaceous, usu-
ally scarcely discolorous, glabrous except on the costa and the veins; petioles
rigid, to 3 cm long, to 0.2 cm wide, angular, the blade decurrent on the upper
part of petiole; stipules deciduous, subulate, to 1.5 cm long, ca. 0.12 cm wide,
scarious, glabrous. Infloreseences usually several in the axils of the terminal
leaves, spiciform-cylindrical, ascending, the peduncles unbranched, ascending,
often falcate, shorter than or exceeding the leaves, to 12 cm long, to 0.3 cm wide,
lignose, puberulent, usually lenticellate. Flowers with pedicels slender, pubes-
cent, to 0.2 cm long, the bracts basal, linear, about as long as the pedicel; hy-
panthium narrowly cuneate cylindrical, ca. 1.2 mm long and indistinguishable
from the pedicel, minutely puberulent, the calycine tube obsolete, the teeth 5,
erect, or spreading, oblong deltoid, subequal, to 0.5 mm long, obtuse; corolla
white, or pale yellow, inflated, urceolate, ca. 2.2 mm long, often slightly con-

1980]

DWYRR— FLORA OF PANAMA (Family 179. Riihiaccac)

25

Figure 3. Alscis hlackiana Hemsl.— A. Flowering branch (xVi). [After Croat 14824.]— B
Fruiting branch (xVz). [After Ciocil 15560.]

stricted medially, densely white-villose within, glabrate outside, petaloid, the
orifice wide, the lobes 5, much reduced, inflexed slightly, oblong, to 0.5 mm long,
obtuse: stamens 5, exserted, the anthers oblong, ca. 1 mm long, versatile, the
filaments subequal, ribbonlike, ca. 6 mm long, densely villose ca. 1 mm above
the base, attached at base of tube; ovarian disc prominent, mound-shaped, 2-3-
lobed, 0.3-0.6 mm long, the style slender, ca. 3 mm long, expanded apically, the
stigmas 2, coiled, ca. 1.3 mm long, glabrous, the ovary thin walled, the septum
thin, the placentas intrusive, thick, the ovules numerous. Fruits with pedicels to

26 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

0.2 cm long, before dehiscence narrow-oblong, 1.2 cm long, ca. 0.2 cm wide,
often falcate, penis-like or operculoid at apex, attenuate at base, lignose, drying
brown, delicately longitudinally veined, puberulent and farinose, dehiscing from
apex to base, the valves 2, drying lustrous yellow within; seeds numerous.

This species occurs only in Panama and Colombia, It fruits readily but despite
numerous collections from Panama, it has been rarely collected in flower. In his
original description Hemsley says that the stamens of the lowermost flowers are
long-exserted, with those of the upper flowers included. This is not the case in
the few flowering specimens examined. Croat 14829 notes that protandry occurs
in Alseis hlackiana.

CANAL zone: Salamanca Hydrographic Station, Rio Pequenf, Dod^e et aL 17981 (MO). Pipeline
Road, 4 km NW of Gamboa, Nee 7539 (MO). Road K-9, Stern et aL 39 (MO), barro Colorado
island: Croat 5389^ 6492. 8323, 9134, 9448, 9503, 10756, 14824, 14829, 16206, 15560, 16196, 16208,
16509 (all MO); Ih^yer 1439 (MO): Foster 863 (MO): Haxden 90 (MO); 1029 (GH. MO, US); HIaclik
415 (MO); Shattiick 52, 822 (both MO); Wilbm' & Weaver 10792 (MO); Zetek 5086, 5600 (both MO).
darien: Rio Tuquesa below Quebrada Venado, Bristan 1087 (MO). Rfo Ucurgantf, Bristan 1168
(MO). S of hi Real, Duke 5063 (MO). Line CC, Duke 5236 (MO). Quebrada Nigua below Santa Fe,
Duke 8825 (MO). Rio Pucro, below Pucro, Duke 13120 (MO). Paya, vie Rio Paya, Stern et al. 187
(MO). El Real, Ri'o Tuira, Stern et al. 284 (MO). Campamento Buena Vista, Rio Chucunaque, Stern
et al. 848 (MO). Cerro Piriaque, Tyson et al. 3856 (MO).

2. Alseis darienensis Dwyer.^ type: Panama, Mori 7001 (MO, holotype).

Trees, much branching, the twiglets terete, the bark thin, scaling off, rubes-
cent, glabrous, the nodes moderately crowded on new growth. Leaves ascending,
oblong or lanceolate, 13 cm long, 2.0-5.5 cm wide, deltoid or acute at the apex,
acuminate, the acumen widely triangular, to 1 cm long, ultimately obtuse, cuneate
or acute at the base, moderately inequilateral, the costa slender, drying reddish
above, somewhat prominulous beneath, the lateral veins ca. 9, arcuate, disap-
pearing near the margin, not forming a conspicuous submarginal vein, with sev-
eral smaller veins between a pair of laterals, these leaving the costa at right
angles, soon disappearing, the smallest veins forming a reticulate mosaic beneath,
the margin delicately callose, thin chartaceous, concolorous, beneath appressed
or spreading hirsute on the costa, the hairs at right angles and lateral to the costa
in and near the vein axils; petioles wiry, stiff, to 45 cm long, ca. 0.75 mm wide;
stipules triangular or ovate triangular, to 7 mm long, acute, longitudinally striate,
scarious, drying red, glabrous. Inflorescences terminal and axillary, cymose spi-
cate, to 1 1 cm long, to 0.1 cm wide, rubescent, glabrous. Flowers (calyx only)
with the calycine cup absent, the lobes 5, imbricate, oblong rotund, ca. 1 mm
long, about as long as wide, obtuse, venose, puberulent marginally. Fruits sessile,
narrowly oblong (immature), to 1.2 cm long, to 0.4 cm wide, somewhat com-
pressed, covered with a grey indumentum; seeds membranous, drying yellow,
fusiform, to 5.5 mm long.

^Alseis darienensis Dwyer, spec. nov. Arhores ramuUs glabris. Folia oblonga vel hmceolata, 4-
13 cm longa, 2.0-5.5 cm lata, venis lateralibus ca. 9, tenui-chartacea subtus in costa hirsuta; petiolis
ad 45 cm longis; stipulis triangularibus vel ovato-triangularibus, ad 7 mm longis. acutis glabris. Inflo-
rescentiae terminales vel axillares, cymoso-spicatae, ad 11 cm longae, glabrae. Flores lobis calycis
5 imbricatis oblongo-rotundis. ca. 1 mm longis, obtusis. Fructus angusto-oblongi, ad 1.2 cm longi,
conspicue griseo-puberuli.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 27

Alseis darienensis is known only from Panama. The plant bears little resem-
blance to any Rubiaceae I have encountered in preparing the family for the Flora
of Panama. But because of the cymose-spikelike inflorescence I have elected to

place it \n Alseis, but its placement is quite uncertain.

darien: Rio Tuquesa, middle Tuquesa Mining Co. camp called Charco Peje, 250 m, Mori 7001

(MO).

4. AMAIOUA

Amaioua Aubl., Hist, PI. Guiane 13. 1775. type: A. gidanensis Aubl.

Trees or shrubs. Leaves usually crowded at the apex of the twiglets; stipules
interpetiolar, connate, often irregularly cleft, finally caducous. Inflorescences
terminal or with short lateral axes, fasciculate, capitate or corymbose, the pe-
duncle present or absent. Flowers dioecious, 5-6-merous, the calycine cup tu-
bular, the teeth reduced or conspicuous; corolla cylindrical, sericeous on the
outside, the lobes contorted; stamens included (male flowers), the anthers sessile,
inserted near the middle or on the lower half of the tube; female flowers with the
ovary 2-celled, the ovules numerous in each cell, horizontally and biseriately
arranged. Fruits baccate, 2-celled; seeds numerous, horizontally disposed, tri-
angular and rounded marginally, the testa fibrous,

Amaioua occurs in Central and South America. There are about 25 species
in the genus.

a. Flowers sessile or subsessile.

b. Fruits oblong rotund, to 1.5 cm long; leaves with 6-7 lateral veins \. A. corymhosa

bb. Fruits rotund, 1.8-2.2 cm long; leaves with ca. 14 lateral veins 2. A. magnicarpn

aa. Flowers pedicellate, the pedicels to 1.5 cm long 3. A. pediceUata

1. Amaioua corymbose H.B.K., Nov. Gen. Sp. 3: 419. 1820. type: Colombia,
Humboldt & Bonpland (B, not seen, photo MO). — Fig. 4.

Gardenia hexandra Willd. ex R. & S., Syst. Veg. 5: 243. 1819. type: America Merid., Humboldt &

Bonpland, not seen.
Amaioua fagifolia Desf., Mem. Mus. Hist. Nat. 6: 14. 1820. type: French Guiana, Martin (C, not

seen, photo MO).
Amaioua guianensis Hemsl., Biol. Cenlr. Amer., Bot. 2: 39. 1881, not A. gidanensis Aubl. 1775.
Nomen nudum.

Shrubs or trees to 40 ft tall, the branches terete, the nodes well spaced;

branchlets angular, the leaves often crowded at the apex. Leaves oblong, often

widely so, the large blades about twice as long as wide, 9-23 cm long, 2.5-12.5
cm wide, obtuse at the apex, shortly acuminate, the acumen to 1.2 cm long,

basally inequilateral or equilateral, truncate, rounded, cuneate or acute, the costa
plane to prominulous above, often appressed villose proximally, appressed villose
beneath, the principal secondary veins 6-7, arcuate, prominulous above, prom-
inent beneath, subcoriaceous, usually concolorous, lightly appressed villose on
costa and secondary veins, often sparsely pubescent on margins, the intervenal
area glabrous; petioles to 1.5 cm long, stout, to 0.3 cm wide, often arcuate;
stipules oblanceolate, 1.5-2.5 cm long, ferrugineous-sericeous outside, glabrous
and glandular within. Inflorescences terminal, few-flowered cymes, the peduncles

28

ANNALS OF THT. MISSOURI BOTANICAL GARDEN

[Vol. 67

Figure 4. Amaioua corvmhosa H.B.K.— A. Flowering branch (x»/2).— B. Opened flower
(x2.5). [After Croat 16214.]

arising in a fan, each usually unbranched, slender, stiff, angular, puberulent.
Flowers staminate, pistillate or bisexual (functionally so?), the staminate flowers
slightly larger, sessile or on pedicels shorter than the hypanthium, the buds ru-
gous, turgid below the middle, tapering to an obtuse apex; hypanthium com-
pressed cylindrical ca. 1 mm long, the calycine cup to 5 mm long, to 2.5 mm
wide, carnose, puberulent outside, glabrous to puberulent and eglandular within,
truncate, the calycine teeth 5, subulate, to 1 mm long, obtuse; corolla white, the

1980] DWYF-R— FLORA OF PANAMA (Family 179. Ri/hiaceae) 29

tube cylindrical, somewhat inflated, 5-7 mm long, 1.5 mm wide, carnose, densely
white villose outside, glabrous within, the lobes 5, lanceolate, 5-7 mm long,
glabrous within, minutely white-puberulent adaxially; stamens 5-6, the anthers
linear oblong, 4-4.5 mm long, distinctly petaloid rostrate for 1 mm, the filaments
very short, inserted near middle of tube; ovarian disc doughnut shaped, 1-2 mm
long, 0.9-1.8 mm wide, 0.5 mm high, the style slender, 4-9 mm long, the stigmas
2, subulate, 0.3-0.6 mm long. Fruits compactly clustered, sessile, oblong rotund,
to 1.5 cm long, pruniform, marcescent, sparsely sericeous, often the vestige of
the calyx forming a conspicuous annular scar; seeds compressed, angulate, ca.
2 mm long, shiny red, the surface with a series of sinuous, densely packed some-
what parallel ridges.

Amaiona corymhosa ranges from Mexico to northern South America. It also
occurs in Cuba. The flowers should be studied critically under field conditions.
Plants have been described as dioecious, but some dissected flowers have func-
tional anthers, a well-defined style and a pair of stigmas. The ovary is not apparent
in these flowers. One collection, Croat 11532. has fruit more oblong in shape and
does not have the appearance of a small dried prune so typical of A. corymhosa.
The pericarp is smoother than the typical. Croat noted on the collection that the
fruits were old, which may account for the difference in appearance.

CANAL zone: Fort Kobbe, near beach, Duke 4719, 4724 (both MO). Albrook, Dwycr & Rohyns
43 (MO). Headquarters J.W.T.C, Johnston 1574 (MO). W of Gamboa, Nee 6772 (MO), barro
COLORADO island: Croat 6072. 15164, I62I4 (all MO): Foster 1337 (MO). CHiRiguf: San Lorenzo,

5 m, Lao 598 (MO), cocle: La Mesa, 900-930 m, Croat 37462 (MO), colon: Salud, Lao & Hohlrid^e
225 (MO), darien: Pico Piriaque, Duke 8137 (MO). Piiias, Duke 10598 (MO). Baia de Pinas, Stern

6 Chambers 192 (MO). Mouth of Rfo Lara, Tyson et al. 3864 (MO). Piriaque Camp at River, Tyson
et al. 4741, 4746, 4754 (all MO). Panama: Taboga Island, Allen 134 (BM, MO). Cerro Azul to Cerro
Jefe, Austin 24 (MO): Blum et al. 18 (MO); Croat 11532 (MO). Panama Viejo, Duke 5736 (MO). San
Jose Island, Duke 12503 (MO). Cerro Azul to Cerro Jefe, Dnyer 1487. 1501, 1871, 207L 2776 (all
MO), 8041 (GH, MO); Duyer & Gentry 10243 (MO); Guerra 5 (MO); Lewis et al. 268 (GH, K, MO,
UC, US); Porter et al 4079 (MO, SCZ, US); Reece 5 (MO). Villa Rosario, International Highway,
Saldana 21 (MO). San Jose Island, Stimson 5328 (MO). Cerro Azul to Cerro Jefe, Tyson 2089 (MO);
Tyson et al. 4061, 4314, 5381 (all MO), vlraguas: La Yeguada, 600 m, Correa 2217 (MO). Isla de
Coiba, Dwxer 1629 (MO).

2. Amaioua magnicarpa Dwyer.^ type: Panama, Dwyer 9552 (MO).

Trees 6-15 m tall, the branchlets ultimately angular, the older bark cracking,
ashen grey when dry, glabrous, the nodes somewhat swollen, moderately crowd-
ed. Leaves oblong lanceolate or oblong rotund, 11.5-17 cm long, 5-9 cm wide,
deltoid to obtuse at the apex, acuminate, the acumen short, 0.4-1 cm long, obtuse
or acute, often falcate, basally cuneate obtuse, scarcely inequilateral, the costa
slender above, prominent beneath, the lateral veins ca. 13, at first strict, then
arcuate toward the margins, the secondary veins perpendicular, subcoriaceous,
concolorous, glabrous except appressed-white ciliate on the costa and veins, the
minute raphides often abundant above, the margin lightly revolute; petioles to 1.5

"^Amaioua ma^niearpa Dwyer, spec. nov. Arbores 6-15 m altae. Folia oblongo-lanceolata, vel
oblongo-rotunda, 11.5-17 cm longa, 5-9 cm lata, venis lateralibus ca. 13 subcoriacea glabra praeter
costam venasque appresso-albido-citiatas: petiolis ad 1.5 cm longis. Fructus rubri et cerasini rotundi,

1.8-2.2 cm longi, apice saepe umbonati.

30 ANNALS OF THE MISSOURI BOTANICAL GARDHN [Vol. 67

cm long, to 2.2 mm wide, rigid, densely appressed villose. Flowers not seen.
Fruits red, cherry-like, rotund, 1,8-2.2 cm long, obtuse at the apex and base, often
umbonate at the apex, puberulent, with the scars of the floral parts scarcely
elevated, ca. 0.5 cm in diam.

The new species is known only from Panama. It is distinguished from the
relatively common A. corymbosa (Willd.) H.B.K. by having much larger fruit
which resemble cultivated commercial cherries.

colon: Santa Rita Ridge, ca. 2000 ft, Dwycr 9552 (MO). 9.4 km N of Goofy Lake, Folsom el

al. 1976 (MO). Panama: El Llano-Carti 10-12 km from junction Inter-American Highway, Mori <i

KaUunki 2880 (MO). El Llano-Carti Road, 1.7 km from Inter-American Highway, Mori, Gentry &
Kalhinki469l (MO).

3. Amaioua pedicellate Dwyer.^ type: Panama, Croat 25937 (MO, holotype; NY,
isotype).

Trees 5-10 m tall, to 22 d.b.h., the branchlets terete, glabrous, the bark dis-
colorous, the nodes well spaced. Leaves ovate-oblong, 6-19 cm long, 2.3-10 cm
wide, widely deltoid toward the apex, attenuate-acuminate, the acumen to 1 cm
long, falcate, ultimately acute, basally acute or cuneate, rarely obtuse, the costa
prominulous above, prominent beneath, the lateral veins 12-14, rigid, especially
prominent beneath, subcoriaceous, discolorous, above dark brown, yellow brown
beneath, golden pilose with the veinlets pinnately disposed; petioles rigid, woody,
to 1.5 cm long, the hairs golden, spreading or appressed; stipules not seen. In-
florescences terminal (in fruit) radially umbelliform, to 5 cm long, the peduncle
absent, the pedicels to 1,5 cm long, rigid, puberulent. Flowers not seen except
for a fragment of the calyx; calycine cup cylindrical, to 5 mm long, ca. 4 mm
wide, truncate, ciliate without and within, the hairs golden, appressed, the teeth
5, triangular-subulate, to 0.3 mm long, these arising from a thin keel on the body
of the cup. Fruits ovate-oblong or oblong-rotund, to 1.5 cm long, black when
dry, purple brown, lightly puberulent.

Amaioua pedicellata is known only from Panama. It is readily distinguished

by the long-pedicellate fruits and presumably flowers disposed in the form of an
umbel.

cocle: Lumber camp at Alto Calvaro, 7 km N of El Cope, 900 m, Folsom 1290 (MO), veraguas:
Road between Fscuela Agricola Alto Piedra and Ri'o Dos Bocas, 730-770 m, Croat 25937 (MO). NW
of Santa Fe, 8.8 km from Escuela Agricola Alto Piedra, Mori et al. 4001 (MO).

5. AMPHIDAYSA

Amphidaysa Standley, Field Mus. Nat. Hist,, Bot. Ser. 11: 180. 1936. type
Sahicea amhigua Standley = Amphidaysa amhi^ua (Standi.) Standi.

^Amaioua pedicellata Dwyer, spec. nov. Arhores 5-10 m altae, ramulis glabris. Folia ovato-
oblonga, 6-19 cm longa, 2.3-10 cm lata, apicem versus lato-deltoidea attenuato-acuminata acumine
ad I cm longo, subcoriacea venis lateralibus 12-14; petiolis rigidis, ad 1.5 cm longis. Inflorescentiae
terminales radiate umbelliformes ad 5 cm longae, pedunculo nullo pedicellis ad 1.5 cm longis. Flores
cupula calycis cylindrica, ad 5 mm longa, ca. 4 mm lata, truncata dentibus ad 0.3 mm longis. Fructus
ovato-oblongi vel oblongo-rotundi, ad 1.5 cm longi.

19801 DWYHR— FLORA OF PANAMA (Family 179. Ruhiaceac) 3 1

Pittierothamniis Steyermark Bol. Soc. Venez. Cienc. Nat. 23 (101): 92. 1962. type: Steyermark
86323 (holotype, VEN), 89361 (paratype, VEN), 89763 (paratype, VEN).

Herbs or subshrubs, the stems unbranched. Leaves often crowded at the
apices of the stems, petiolate, the stipules interpetiolar, large, entire or more
often the margin setose pectinate, the processes numerous, unequal. Inflores-
cences axillary or terminal, capitate to trichotomously cymose, the bracts entire
to multifid, often obscuring the flowers. Flowers with the hypanthium oblong or
turbinate, the calycine cup short, the lobes 5, unequal, somewhat foliaceous,
persistent; corolla tubular, 5-lobed, the lobes valvate; stamens 5, inserted in the
middle of the tube, the anthers linear, dorsifixed; ovary 2-celled, the ovules nu-
merous in each locule, the placentas axile, adnate to the axis. Fruits fleshy,

indehiscent, the persistent calyx conspicuous.

Amphidaysa is found in Panama, Colombia, Venezuela, and Brazil. The genus
includes 6-7 species.

Literature:

Steyermark, J. 1972. Amphidaysa. in The Botany of the Guayana Highland
Mem. New York Bot. Gard. 23: 316-322.

a. Lobes of the calyx 5-6, 1. 2-1. 5 mm wide 1. A. amhii^aa

aa. Lobes of the calyx 4, 2-4 mm wide 2. A. spathuhita

1. Amphidaysa ambigua (Standley) Standley, Field Mus. Nat. Hist., Bot. Ser.
11: 181. 1931.— Fig. 5.

Sahicca ambigua Standley, Field Mus. Nat. Hist., Bot. Ser. 7: 49. 1930. iypk: Colombia. Triana 1847,
not seen.

Subshrubs to Y^ m tall, the stem occasionally subprostrate or upright, lignose,
often rough with petiole scars, glabrous, the leaves aggregated terminally. Leaves
oblong, elliptic or occasionally obovate, 7-24 cm long, 2.2-8 cm wide, occasion-
ally subfalcate, deltoid or rounded at the apex, acuminate, the acumen short or
to 1 cm long, cuneate at the base, often slightly inequilateral, the costa grooved,
plane or prominulous above, prominulous beneath, 1.2-2.5 mm wide proximally,
the lateral veins 14-18, plane or prominulous above, prominulous beneath, often
opposite, the intervenal areas patulous-reticulate, chartaceous, often discolorous,
glabrous above, puberulent beneath, especially on the costa and veins, rarely
subsessile; petioles 2-6 cm long, to 0.2 cm wide, puberulent, often flexuous;
stipules free, appressed, the sheath rectangular, 4-5 mm long, or much larger
(fide Standley), the lobes 6-8 subulate and flabellately disposed, terminal, 2-3
times the length of the body, acute, densely puberulent. Inflorescences terminal
and axillary, the flowers densely crowded at the apex of the stem, the conglom-
erate shorter than the petioles of the large leaves. Flowers with the hypanthium
ca. 4 mm long, pubescent or glabrous, the calycine cup short, the calycine lobes
5-6, lanceolate, 5-12 mm long, 1.2-1.5 mm wide, acute, subcarnose, somewhat
villosulose or villose especially on the margins; corolla white, the tube cylindrical,
20-30 mm long, 0. 15 cm wide, petaloid to thickly carnose, puberulent on outside,
glabrous or with glandular fimbriate structures within, these to 0.8 mm long, the

32

ANNALS OF TMF MISSOURI BOTANICAL GARDEN

[VoL 67

Figure 5. Amphidaysa amhi^na (Standley) Standley. Habit (XV2). [After Dressier & Mahler

2444 . J

lobes 5, narrow lanceolate or narrowly oblong, 3-5 mm long, with glandular
fimbriate structures adaxially; stamens 3-5, the anthers oblong or ovate-oblong,
2-2.5 mm long, with a terminal petaloid appendage, subulate, 0.5-1 mm long,
acute, the filament short; style slender, to 28 mm long, ca. 0.35 mm wide, the
stigmas oblong, 4-5 mm long, 0.7 mm wide. Fruits oblong or oblong-rotund,
5-7 mm long; calycine lobes persistent, linear-subulate, unequal, to 17 mm long,
acute, ca. 3 times the length of the fruit, the latter truncate, the nectary scar
umbonate, the corolla tube scar annular, smooth or rugulose (the pressure of
seeds against wall on drying), glabrescent to densely pubescent.

1980J DWYKR— FLORA OF PANAMA (Familv 179. Rubiaccac) 33

Amphidaysa amhii^iia is found in Panama and in Colombia. It is of interest
that all Panamanian collections were made during the current decade except for
Terry & Terry 1489,

colon: Santa Rita Ridge, irAny & DArcy 6139 (MO); D\yycr8583 (MO), darikn: Cerro Pirre,
Brisran 511 (MO); Duke 5290, 6114, 13660 (afl MO). Cana-Cuasf Trail, Chepigana. Tcrr\ & Terry
1489 (MO). Panama: Road to Carti. 13 km N El Llano, Basey 385 (MO). Campo Tres, 5 km NE
Altos de Pacora, Basey 828 (MO). Road from El Llano to Carti'-Tupile, Kennedy et al. 2444 (MO).
La Eneida, Cerro Jefe, Maas & Dressier 682 (MO). Road from El Llano to Carti'-Tupile, Dressier &
Mahler 2444 (MO); Liesner 652 (MO), 1142 (MO); Mori, Genlrx & Kallunkl 4558 (MO); Mori &
Kallnnki 5135 (MO); Mori 7710 (MO). 7 km W of Santa Fe, ca. 2900 ft, Nee 11261 (MO), san blas:
Trail from El Llano to Carti-Tupile, Kennedy 2579 (MO). Colombia: choco: N of Alto Curiche,
Duke & Idroho 11228 (MO). Bahia Solano (Puerto Mutis), Gentry & Forero 7228 (MO).

2. Amphidaysa spathulata Dwyer, spec, nov.*' type: Panama, Duke 15615 (MO).

Subshrubs about V?* m tall, the stem erect, simple, terete, glabrous to gla-
brescent, the leaves crowded toward the apex. Leaves sessile or subsessile, ob-
long, 4.5-16 cm long, 1.8-8 cm wide, deltoid or obtuse at the apex, cuneate or
attenuate-acute at the base, the costa slender, prominulous above or subsulcate,
subprominent beneath, the lateral veins ca. 14, arcuate, forming an undulate vein
beneath the margin, the pair of lateral veins including 1-2 subparallel veins, stiffly
membranaceous, glabrous above, the raphides numerous, white, puberulent on
the costa and veins; stipules free, puberulent, the body 2 mm long, thin, provided
with glandular fimbriate structures within, with 6-8 linear setas, 0.4-0.6 mm long.
Inflorescences to 4 cm long, the cymes sometimes on reduced branches. Flowers
with the hypanthium narrow oblong, to 4 mm long, glabrous, the lobes of the
calyx, elliptic-spathulate or narrowly obovate oblong, to 11 mm long, 2-4 mm
wide, the blade 5.5-6.5 mm long, foliose, often 5-veined, acute, puberulent, the
claw rigid puberulent, often longer than the blade, scarcely alate; corolla white,
with the tube narrowly cylindrical, to 3 cm long, 2 mm wide, glabrous,

the lobes 5, lanceolate, to 6 mm long, acute; stamens 4, the anthers oblong
rotund, the acumen ca. 0.4 mm long; stigmas 2, subulate, ca. 2.5 mm long. Fruits
subsessile or on thick pedicels, puberulent, these provided with a stipuloid brac-
teole toward the apex, this with 5 setas, linear subulate, to 8 mm long, narrowly
oblong, to 1.5 cm long, 0.4 cm wide, 6-angled, lignose, drying black, with 8
slender ribs.

The new species is known only from Panama but it is to be expected in
Colombia. It is closely related to A. neblinae Steyermark from Brazil which has
more acute leaves and narrower calycine lobes.

darien: Cerro Campamenlo, S of Cerro Pirre. Dnke 15615 (MO), 15731 (US).

*' Siiffrutices foliis conferlis. Folia sessilia ve! subsessilia oblonga, 4.5-16 cm longa, 1.8-8 cm
lata, venis lateralibus 14, rigido-membranacea supra glabra; petiolis ad 0.5 cm longis; stipulis liberis,
corpore 2 mm longo, tenui intus fimbriis glandulis instructo setas 8 lineares ferente, eas 0.4-0.6 mm
longas. fnflorescentiae ad 4 cm longae. Flores lobis calycis 4 elliptico-spathulatis vel angusto-obovato-
oblongis, ad II mm longis, 2-4 mm latis, lamina 5.5-6.5 mm longa, foliosa: corolla tubo ad 3 cm
longo, 0.2 cm lato, glabro, lobis 5, ad 6 mm longis; staminibus 4, antheris acumine ad 0.4 mm longo,
Fruetiis subsessiles vel in pedicellis crassis versus apicem bracteola stipulanea praediti, hac 5 setis
lineari-subulatis ornata, angusto-oblongi, ad 1.5 cm longi. 6-anguIati costis 8 instruct!.

34 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

6. ANTIRHEA

Antirhea Commerson A, L. Juss., Gen. PI. 204. 1789 ('Antirhea'). type:
A. horhonica J, F. Gmelin.

Antirhoca Endlicher, Ench. 272, 692. 15-21 Aug. 1841 orth. mut.

Stenostomum Gaertn. Fruct. Sem. PI. 3: 69. 1806. type: 5". hicidum (Swartz) C. F. Gaertn.
Stunnia Gaertn. Fruct. Sem. PL pL 192. 1805, not Sturmia Hoppe (1799), not Stimnia Reichb. (1828).
typb: S. lucida Gaertn.

Trees or shrubs. Leaves often coriaceous, the stipules deciduous or persistent.

Inflorescences axillary, cymose, the peduncle when present with branches
spreading, the flowers secundly disposed, the bracteoles present or absent. Flow-
ers with the hypanthium ovoid or obovoid, the calyx lobes persistent; corolla
funnelform, the 4-5 lobes imbricate, 2 of them exterior; stamens 4-5, included
or scarcely exserted, the anthers slender, dorsifixed, often pubescent; style slen-
der, the stigma knoblike or 2-3-lobed, included, the ovary 2-10-celled, the ovules
solitary, pendulous. Fruits drupaceous, thin carnose, the shell ligneous or bony,
the seeds elongate, pendulous.

Antirhea, with about 40 species, occurs in the West Indies, Panama, Mad-
agascar, in tropical East Asia to Australia,

1, Antirhea trichantha (Griseb.) Hemsley, Biol. Centr. Amer., Bot. 2: 42.
1881.— FiG. 6.

Pittoniotis trichantha Griseb., Bonplandia 6: 8. 1858. typk: Duchaissuii> (? P, not seen).

Trees to 10 m tall, the twiglets round, smooth, drying black, glabrous or

puberulent. Leaves oblong to ovate-rotund, 6-21 cm long, 2.5-8.5 cm wide, sub-
equilateral, deltoid to rounded at the apex, often contracted into a narrow cusp

to 2 cm long, usually acute, occasionally obtuse, basally acute to cuneate, the
costa subplane above, prominulous to subprominent beneath, the lateral veins
ca. 12, arcuate, the intervenal areas irregular pinnatiform, chartaceous, discolo-

rous, densely strigillose above, densely pilose or villose beneath; petioles slender
or stout, to 3 cm long, 0,8-1.5 mm wide, pilose; stipules persistent or decidous,

appressed to the stem, triangular subulate, to 1.5 cm long, pilose. Inflorescences
several at apices of the twiglets, axillary, spreading; peduncles usually wiry, 4-
7 cm long, ca. 0.8-1.5 mm wide, strict or flexuous, with a few alternate flexuous
branches, these spreading, either with few flowers or occasionally with flowers
densely congested, the flowers secundly disposed. Flowers with hypanthium ob-
long rotund or oblong, ca. 1.5 mm long, densely pubescent, the calycine cup
short, the teeth 5, oblong rotund, to 0.4 mm long, obtuse, petaloid, pubescent on
the margin and outside; corolla white with the tube narrowly cylindrical or nar-
rowly funnelform, 2.5-3 mm long, ca. 0.6 mm wide, petaloid, glabrous within,
the lobes 5, oblong, about as long as the tube, densely villose above; stamens 5,
exserted, the anthers linear oblong, 1.6-2 mm long, villose, the filaments linear,
elongate, 0.5-2 mm long, attached at the corolla mouth; stigmas linear oblong,
Vi the length of the style, the style slender but dilated below stigmas. Fruits
drupaceous, drying red, fleshy, oblong, 0.5-0.8 mm long, usually twice as long

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

35

4^. c&(

''■■ '^

r

rj. ^-4. cx-^

Figure 6. Antirhea trichantha (Griseb.) Hemsley.— A. Habit (x.45).— B. Flower (x4.5).
C. Ovary, cross section (x4.5). [After Stern et al. 149.]

as wide, capped by a calycine cup to 0.8 mm long, drying black or brown, gla-
brate, delicately longitudinally veined, seed solitary ca. 5 mm long, unilaterally
grooved.

Antirhea trichantha is known only from Panama. Few collections have been
made outside the Canal Zone.

CANAL zone: Victoria Fill, Allen 1761 (MO), Chiva-Chiva Road, 1 mi E of Gaillard Highway,
Blum 468 (MO). Albrook Tower, vicinity TTC site, BInm 551 (MO), C-15 Road to TTC Albrook
Tower, Blum 2240 (MO). Vicinity of Rio Cocoli, Road K-9, Chambers et al. 37 (MO). Madden Forest
Preserve, Correa 146 (MO). Gaillard Highway near Penitentiary, Croat 9311 (MO). Road C 2 F, Vi
mi NW Summit Naval Radio Station, Croat 11033 (MO), Fort Kobbe, Duke & Mussell 6667 (MO).

36 ANNALS OF THR MISSOURI BOTANICAL GARDEN [VoL 67

Power Station near Fort Kobbe, Duyer 2668 (MO). US Army TTC, Albrook, Dwycr 6727 (MO).
Madden Dam, Duycr & Ellas 7477 (GH, MO, US). Madden Dam, Boy Scout Road, Dwyer &
Lalhithin 8579, 8824 (both MO). Madden Dam, Dwycr & Rohyns 24 (MO). Farfan Beach Road,
Gentry 4901 (MO). Navy Pipeline Preserve, 3 mi NW Gamboa, Haines 509 (MO). Madden Dam,
HoldrUliye 6248 (MO). Madden Dam, Boy Scout Road, Kirkhride 45 (MO). Farfan Beach, Kirkhride
& Elias 74 (MO). US Navy Ammunition Depot, 2 mi NW of Balboa, Luteyn S: Foster 930 (MO).
Howard Air Force Base, SE Kobbe Beach, Oliver & MacBrxde 1874 (MO). Madden Dam, Boy Scout
Road, Porter et at. 4015 (COL. DUKE, MO, SCZ, UC, VEN). Naval Ammunition Depot. Stern

et al. 38 (MO). Farfan Beach, Tyson 1824 (MO), barro Colorado island: Croat 4971 , 6336. 10391,
11973, 14255 (all MO). DAR!f:N:"Rio Pucro below village of Pucro, Duke 13134 (MO). El Punteadero
bridge, over Rio Chucunaque, Stern et al. 149 (MO). Panama: Mocambo, Croat 14906 (MO). Rio
Pita, Dnke 4800 (MO). Chilibre, Lao & Holdridi^e 8 (MO). 6 km SW Arraijan, old road to Bique,
Nee 6890 (MO),

7. APPUNIA

ifl'

B. & H.

Bellynkxia Muell.-Arg., Flora 48: 465. 1875. typf-: B. angalata Muell.-Arg.

Shrubs or small trees. Leaves usually lanceolate, opposite; stipules free, sub-
ulate acuminate. Inflorescences terminal or axillary, the flowers few, congested
into a small head at the end of the peduncle. Flowers with the hypanthium some-
what coherent, the calycine cup truncate, the teeth absent or scarcely visible,

funnelform or U-shaped, the lobes usually 5, valvate; ovary 4-celled, with an
ascending, solitary ovule in each cell. Fruits sessile, aggregate, but virtually free

at maturity, each fruitlet fleshy, drupaceous, with 4 pyrenes, each pyrene un-
equally 2-locellate, the ventral part empty; seeds exalbuminous.

Appunia is a genus of approximately 10 species in Central and South America.

Literature:

Hayden, M. V. & J. D, Dwyer. 1969. Seed Morphology in the Tribe Morindeae
Bull. Torrey Bot. Club 96: 704-710.

1, Appunia seibertii Standley, Ann. Missouri Bot. Card, 24: 208. 1937. type:
Panama, Seihert 624. — Fig. 7.

Shnths to 1,5 m tall, the branches exfoliate, subterete, smooth, red brown and
cracking when dry, glabrous or puberulent, the nodes thickened, the branchlets
slender. Leaves somewhat verticillate at the apex of the branchlets, narrow el-
liptic, rarely subrhombiform or obpyriform, acute or deltoid at the apex, attenuate
cuneate at the base, 5-17 cm long, 1-7 cm wide, membranaceous, scarcely dis-
colorous, often green when dry, glabrous or scattered granulose above, the in-
tervenal areas minutely granulose beneath, the costa slender, prominulous above,
subplane distally beneath, the lateral veins opposite or alternate, ca. 12, slender,

prominulous, widely arcuate, the veins white puberulent, the margins glabrous
or minutely puberulent; petioles 0.3-2 cm long, ca. 0.7 cm wide, puberulent;
stipules triangular, 2.5-5(-7) mm long, 2-4 mm wide, smooth, glabrous, entire.
Inflorescences 1-4, pseudoterminal, arising from the axils of the fasciculate
leaves, the peduncles several per axil, umbellately disposed, slender, 0.5-1,5 cm

1980]

DWYER— FLORA OF PANAMA (Family 179. Riihiaceac)

37

Figure 7. Appunia seihertii Standley.— A. Habit (x4.5). [After Oliver <& MacBryde /S%.]— B.
Fruit (X.09). (After Duke 4713.]

long, ca. 0.5 mm in diam., minutely red or white puberulent, the flowers aggre-
gated into a solitary terminal head ca. 4 mm in diam., with 10-11 flowers. Flowers
sessile; calyx cupuliform, ca. 1.5 mm in diam., the teeth absent or sometimes as
irregular points, to 0.5 mm long, fleshy, glabrous, the raphides many; corolla (in
bud) with the tube ca. 7 mm long, ca. 2 mm wide, glabrous outside, pubescent
within, the lobes 5, oblong to triangular, scarcely shorter than the tube, briefly
cucullate; anthers 5, oblong, to 2 mm long, the filaments attached near the apex
of the tube; ovary 4-loculate, the septa thick, the style short, the stigmas 2,

38 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

subulate ca. 1.25 mm long. Fruits sessile, usually 2-5, crowded at the apex of
the peduncle, the individual fruits turbinate, to 1 cm long, black and smooth when
dry, often inconspicuously 4-lobate, each pyrene unequally 2-locellate, the ventral
part empty, bulliform, the endocarp replete with raphides; seeds 4, oblong, ca.
5 mm long, slightly winged.

This species is known from Panama and Colombia. Additional notes may be
found in Dwyer & Hayden (1969), where there is an emended description of the
species.

In forming fruit often several flowers of the mass fail to mature.
The persistent calycine tube and floral disc, as well as the hypanthium, sim-
ulate a Morinda fruit in miniature.

CANAL zone: Fort Kobbe, Duke 3958, 4246, 4698, 4713 (all MO). Howard Air Force Base, SB
of Kobbe Beach, Oliver & MacBryde 1896 (GH. MO, US), darien: Without locality, Bristan 207
(MO). El Real to Pinogana, Duke 5126 (MO). Rio Pucro to Quebrada Maskia, Duke 13082 (MO). Rio

Morti\ Camp Morti Hydro, Duke Ml 18 (MO). Chepigana, Duke ct Bristan 267 (MO). Rio Tuquesa,
lower Tuquesa Mining Camp called Charco Chiva, Mori 6987 (MO). Periaque camp at river, Tyson

et al. 4767 (MO). Panama: Ri'o Bayano above confluence with Ri'o Chepo, Duke 4000 (MO). Cermeno,
Dwyer 5604 (MO). Arenoso, lower Ri'o Trinidad, Seihert 624 (MO). 2 mi E of El Llano, Tyson 1749
(MO). SAN Bi as: Isia Pino near Mulatupo, Elias 1718 (MO, UC, VEN).

8. ARCYTOPHYLLUM

Arcytophyllum Willd. ex. Schultes in R. & S., Syst. Veg. 3: 5. 1827. type: A.
hlacrioides Willd. ex. J. S. & J. H. Schultes.

Mullostonm Karst., Fl. Colomb. 2: 9. 1862. TVPt not indicated.

Rachicallis DC, Prodr. 4: 433. 1833. type; R. rupestris (Sw.) A. P. DeCandoIle, based on Hedyotis

rupestris Sw.
Pseudorluuhicallis Benth. & Hook., Gen. PI. 2: 60. 1873. typh not indicated.
Ereicoctis Kuntze, Rev. Gen. PL 281. 1891. typl not indicated.

Shrubs or subshrubs, erect or prostrate. Leaves small, crowded, often densely

imbricate, coriaceous; stipules entire, bifid or setose. Inflorescences cymose or

flowers solitary, axillary. Flowers with the hypanthium globose or turbinate, the

calyx 4-5 lobed, often with glands between the lobes; corolla funnelshaped or

salverform, the lobes 4; stamens with the anthers included or exserted, dorsifixed,

the filaments short and adnate to the tube in riblike fashion; style slender, the
stigmas 2, short, the ovary 2-celled, the placentas adnate to the septum. Fruits

capsular, turbinate to globose, 2-celled, usually septicidally dehiscent to the base;

seeds few, boatshaped or piano convex, oblong, punctate, winged.

Arcytophyllum, a genus of about 30 species, ranges from Mexico through
Central America, extending to the Andes of South America.

1. Arcytophyllum lavarum Schum. in Engl. & Prantl, Nat. Pflanzenfam. IV (4):
28. 1891. type: Panama, Maxon 5349 (US, holotype).— Fig. 8.

Mallostoma hivarufn (Schum.) Donnell. Smith, Enum. PI. Giiatem. 5: 36. 1899.

Suhshruhs, cricoid in habit, profusely branched, the branches lax, glabrous,
drying silver tan or purple black, ridged, the nodes well spaced or crowded,
swollen (bases of persistent stipules). Leaves oblong, 0.5-0.8 cm long, 0.2-0.4

1980)

DWYFR— FLORA OF PANAMA (Family 179. Ruhiareae)

39

D

Figure 8. Anytophyllum lavarum Schum. — A. Habit (x'/i). — B. Flower, opened (x5). — C.
Fruits (x5).-— D. Seeds (x5). [After Wilbur ct aL 11033.]

cm wide, obtuse at the apex, obtuse to truncate at the base, the costa and the
lateral veins evanescent, carnose, cricoid, lustrous, scarcely bicolorous, marces-
cent on drying; petioles to 1 mm long or absent; stipules connate, disposed as a
short cup, to 0.25 cm long, the free portion deltoid, longer than the cup and often
contiguous vv^ith the stipule above, the margin erose with a few ascending irregular
subulate teeth. Inflorescences terminal, to 1.5 cm beyond the uppermost leaves,
usually with 2 peduncles, to 0.3 cm long, each bearing a terminal cymule. Flowers
with the hypanthium rotund, to 0.65 mm long, petaloid, glabrous, the calycine
cup very short, with patches of flabellately disposed minute oblong glands and
squamellae intermixed within, the teeth 4, erect, oblong or ovate oblong, often
obtuse, to 1.3 mm long, often with an upright gland at the junction of 2 lobes,
glabrous, petaloid; corolla white, white tinged with purple, blue or lavender pink,
the tube campanulate oblong, ca. 2.5 mm long, the lobes not cucullate, villosulose
on the adaxial face; stamens 4, the anthers oblong, 0.8-1 mm long, dorsifixed,

40 ANNALS OF THF MISSOURI BOTANICAL GAROFN [Vol. 67

the filaments slender, attached above the middle of the tube, the style slender,
ca. 4 mm long, slightly dilated apically, the stigmas 2, clavate when connate, ca.
0.4 mm long; ovary thin walled, the septum thin, the ovules several, arising on
a distinct stipe attached almost at the base of the septum, the locule ample. Fruits
short pedicellate, capsular, rotund or compressed rotund, to 1.5 mm in diam.,
truncate, the calycine lobes persistent, a thin annular ring marking their point of
origin, the valves splitting from the apex.

Arcytophylliitn lavanini is restricted to high mountainous regions of Costa
Rica and Chiriqui. A photo of /l. lavarum at MO is labeled as a Pittier collection
(unnumbered) from Costa Rica, deposited in Berlin, but this is not the type.

chiriqui: Volcan de Chiriquf, Davidson 1293 (F, MO). Between Barii and Respinga, 3000 m,
D'Arcx 10095 (MO). Chiriqui Volcano, 3374 m, Maxon 5349 (US). W side of Volcan Barii, ca. 3000
m. Mori & Boltcn 7342 (MO). Trail to summit, Volcan de Chiriqui 2500-3400 m, Mori & Kallunki
5748 (MO). Volcan de Chiriqui, W slope, Wilbur ct al. 11033 (F, MO). Potrero Muleto to summit of
Volcan de Chiriqui, Woodson A Sclwrr 437 (MO). Loma Larga to summit of Volcan de Chiriqui',
Woodson ct id, 1087 (F. MO).

9. BATHYSA
Bathysa Presl., Bot. Bemerk. 84. 1844. type: B. stipuJata Presl.

Trees or shrubs. Leaves opposite, the stipules interpetiolar, entire, occasion-
ally persistent. Inflorescences terminal, paniculate. Flankers abundant^ the ca-
lycine cup truncate, the teeth 4-5; corolla subrotate to infundibuliform, the lobes
4-5, imbricate; stamens 4-5, inserted on the mouth, the anthers dorsifixed,
exserted; ovary 2-celled, the ovules numerous. Fruits capsular, septicidally de-
hiscent, splitting into 2 valves from the apex; seeds minute, horizontal, com-
pressed or angular, winged or not, the testa reticulate.

Bathysa ranges from Panama through Venezuela, Brazil, and Peru. It is a
genus of about 10 species.

1. Bathysa veraguensis Dwyer." type: Panama, Lao & Gentry 53! (MO, holo-
type). — Fig. 9.

Trees to 5 m tall, the branchlets terete, minutely velutinous, the nodes well
spaced. Leaves large, widely oblong, 10.5-31.0 cm long, 4.5-18.0 cm wide,
widely deltoid toward the apex, occasionally rounded, acuminate, the acumen
short, widely deltoid or narrowly deltoid, to 1.8 cm long, 0.3-0.5 cm wide in the
middle, basally widely cuneate, obtuse or truncate, the costa prominulous above,
prominent beneath, the lateral veins 15-19, to 2 cm apart in the middle of the

"^Bathysa vcrui^nensis Dwyer. spec. nov. Arhores ad 5 m altae. Folia magna, 10.5-31.0 cm
longa, 4.5-18.0 cm lata, venis lateralibus 15-19 papyracea supra in costa venisque aureo-villosula

cetera minute scabridula subtus dense aurcovelutina. Inflorcscentiac terminales amplae, ad 30 cm
longae, cymulis plerumque 3-fioribus; bracteis ovato-lanceolatis, ad 1.3 cm longis. Flores cupula
calycis ampla ca. 2.5 mm alta, apice ca. 0.6 cm lata, lobis 4-5, ad 5 mm longis: corolla alba tubo 3.3-
5.0 mm longo, lobis ca. 6 mm longis; antheris ca. 3.2 mm longis, filamentis crasso-subulatis. Fructus
oblongi vel turbinati, ad 1.5 cm long], ca. 0.5 cm lati, extus puberuli calyce persistenti magno ad I
cm longo, fortasse 2-plo longitudine pericarpium superanti scminibus polymorphis plano-compressis
reticulatis ad 0.8 mm longis.

1980]

DWYHR— FLORA OF PANAMA (Family 179. Ruhiaceae)

41

Figure 9. Bathysa veraguensis Dwyer. — A. Inflorescence and leaf (x. 45). — B. Leaf underside
(X.45).— C. Flower, opened (x3.15). [After Lao & Gentry 53L]

42 ANNALS OF THE MISSOURI BOTANICAL GARDKN [Vol. 67

blade, the venules irregular perpendicular, papyraceous, discolorous, dark red to
brown when dry, golden villosulose above on the costa and the veins, otherwise
scabridulous, beneath densely golden velutinous; petioles and stipules not seen.
Inflorescences terminal, solitary, pyramidal paniculate, to 30 cm long, to 30 cm
wide, velutinous; peduncle to 8 cm long, to 0.35 cm wide, thick, lignose, the
branches often opposite, to 15 cm long, the pairs well spaced, often 6-7 cm apart,
the cy mules often 3-flowered, disposed on long or short secondary branches or
on tertiary branches, each branch provided with 3 flowers, rarely with one flower;
bracts ovate lanceolate, to 1.3 cm long, acute, appressed villose outside, inside
with a few hairs; bracteoles ovate subulate. Flowers pedicellate, the pedicels 0.3-
1.0 cm long, hypanthium turbinate, 0.3 cm long, puberulent, the calycine cup
expanded, 2.5 mm long, 0.6 mm wide at the apex, eglandular within, the lobes
4-5, oblong triangular, 5 mm long, often ascending, 0.4-0.5 cm wide at the base,
obtuse, coriaceous, puberulent or appressed villose within; corolla white, the
tube compressed cylindrical, 3.3-5.0 mm long, often to 0.5 cm wide, glabrous
outside, within densely golden villose near the mouth, the lobes imbricate in bud,
oblong rotund, 6 mm long; stamens 5, the anthers narrowly oblong, 3.2 mm long,
slightly curved at anthesis, obtuse, exserted, the filaments attached near the
mouth, thickly subulate, 3 mm long; ovarian disc ringlike, 3.0-4.5 mm long, 1.3-
1.6 mm wide, 0.5-0.65 mm deep, the style thick, 7.5 mm long, the stigmas 2,
thick, narrowly oblong, 2.5 mm long. Fruits immature? oblong or turbinate, to
1 .5 cm long (including calyx), ca. 0.5 cm wide, the persistent calyx as in flowering
stage, perhaps 2 times the length of the pericarp, densely puberulent outside,
densely appressed villose within; seeds polymorphic, mostly oblong, to 0.8 mm
long, compressed, reticulate.

Bathysa verai^uensis is known only from Panama. The genus is not otherwise
known in Central America.

cocle: Cerro PikSn, ca. 3100 ft, Duw-r & UiUatlun 8641 (MO). Cerro Pilon, hill below summit,
ca. 2500 ft, DuycT ct al. 4544 (MO), veraguas: Cerro Tute, 1200 m, Lao & Gentry 531 (MO). NW
of Santa Fe, 2.1 km from Escuela Agn'cola Alto de Piedra, M<m & Kallunki 4768 (MO).

10. BERTIERA

Bertiera Aubl., Hist. PI. Guiane 180. 1775. ''Berthiera'' typh: B, j^uianensls
Aublet.

Shrubs or small trees. Leaves petiolate or sessile; stipules connate, the sheath
bifid. Inflorescences terminal, cymose paniculate, pedunculate, the bracts nar-
row. Flowers with the hypanthium subglobose or turbinate, the calycine cup
truncate and edentate or with 5 teeth; corolla funnelshaped, the tube usually
sericeous outside, the lobes 5, ovate, contorted; stamens 5, the anthers ovate or
subulate, dorsifixed, the connective acuminate or produced at the apex, the fil-
aments short, attached to the tube; ovarian disc cupular or annular, the style
slender, glabrous, the stigmas entire or disposed as 2 narrow lobes, the ovary
2-celled, the ovules numerous, the placentas entire, affixed to the septum. Fruits
baccate, fleshy; seeds numerous, small, angulate, the testa foveolate or granulate.

Bertiera occurs in tropical America and in Africa. It is a genus of about 30
species.

1980] DWYHR— FLORA OF PANAMA (Family 179. Ruhiaceae) 43

Literature:

Standley, P. S. 1934. 65. Bcrtiera Aubl., N. Am, Fl. 32(3): 180-181.

a. Leaves with 8-9 lateral veins; stipules 1.5(-2.0) cm long 1. B. angusfifoUa

aa. Leaves with ca. 7 lateral veins; stipules 0.9-1.2 cm long 2. B. guianensis

1. Bertiera angustifolia Benth., Bot. Voy. Sulphur 103. 1844. type: Cocos Island,
Costa Rica, not seen.

Shrubs or trees to 5 m tall, the branchlets terete, smooth, densely pilose, the
nodes well spaced. Leaves arising as juveniles from a cylindrical bract or pair of
bracts, to 2 cm long, the bract drying brown, puberulent; leaf blades ovate oblong,
9-17 cm long, 2.7-7.5 cm wide, acute toward the apex, acuminate, the acumen
often inconspicuous, obtuse or rounded at the base, the costa prominulous, the
lateral veins 8-9, strongly ascending, sometimes immersed above, papery when
dry, rubescent above, glabrous except for the costa, glabrous beneath except for
the costa and veins, the hairs appressed, elongate; petioles to 0.5 cm long, pu-
bescent; stipules triangular subulate, to 1.5(-2.0) cm long, ca. 0.5 cm wide at the
base, ultimately appressed to the stem. Inflorescences terminal, solitary, panic-
ulate, puberulent, to 17 cm long, to 7(-10) cm wide; peduncle slender, 3.5-7.0
cm long, to 0.18 cm wide, the rachis somewhat flexuous, the branches alternate
or opposite, rarely 4-verticillate; bracts and bracteoles linear subulate, equal to
the nonfloriferous parts of the branchlets. Flowers small (in bud); hypanthium
oblong, to 1.5 mm long, pubescent, the calycine cup small, the teeth 5, small,
triangular; corolla tube 1.5-2.0 mm long, carnose, glabrous or sparsely puberulent
outside, densely villose inside, the lobes deltoid, to 1.3 mm long, acute; stamens
5, the anthers oblong, to 1.5 mm long, short acuminate at the apex, the acumen
ca. 0.3 mm long. Fruits sessile, rotund, 3 mm in diam., drying black, glabrate to
sparsely pubescent, the hairs short, strigose, the ribs obvious, obtusely margin-
ate, the calycine cup vestigial.

Bertiera angustifolia is known only from Costa Rica (Cocos Island), Panama
and Peru. Standley erroneously annotated Mexia 6358 from Pongo de Manse-
riche, Dept. Loreto, Peru as Bertiera parviflora Spruce. This collection matches
well the three Panama collections cited below. The photo (MO) of the type of
Bertiera parviflora {Spruce 2774) does not match the three Panama collections;
the lateral veins are only 5-6 in the type but 8 or 9 in B. angustifolia.

Bentham described the leaves of B. angustifolia as ''pellucid-punctate,'' but
this is not so in the Panamanian collections. The only genus of Rubiaceae in
Panama with pellucid-punctate leaves is Rustia,

BOCAS DEL TORo: Between Quebrada Menaco and Buena Vista, Chiriqui Trail, Kirkbride & Duke
666 (MO), darien: Cana, 1750 ft. Stern et al. 403 (MO), veraguas: Rio Primero Brazo, 2.5 km
beyond Agricultural School, Alto Piedra. 700-750 m, Croat 25477 (MO).

2. Bertiera guianensis Aubl., Hist. PI. Guiane 1: 180. 1775. type: French Guiana,

Auhlet (P, holotype, not seen). — Fig. 10.

B. paliistris A. Rich,, D.C.. Prodr. 4: 392. 1830.

B. aeciuiiiinunosa Steyermark, Mem. New York Bot. Card. 17(1): 319. 1967. svntype: Panama,

Cooper 586 (NY, holotype).

44 ANNALS OF TFIE MISSOURI BOTANICAL GARDEN (Vol. 67

B. iiidanensis var. leiophxUa Steyermark, Mem. N.Y. Bot. Garden 17: 319. 1967. type: Panama,

Pitticr5l78 (NY, holotype).
B. tennis Lundell, Wrightia 4: 49. 1968. type: Guatemala, Contreras 4318 (LL, holotype).

Shrubs or small trees to 4 m tall, the branchlets terete, slender, appressed
villose, the nodes well spaced. Leaves oblong, 8-17 cm long, 2-6 cm wide, acute
and often vaguely acuminate at the apex, acute or cuneate at the base, the costa
prominulous above and beneath, ca. 0.1 cm wide proximally, the lateral veins ca.
7, arcuate, occasionally strict and sharply ascending, discolorous or not, glabrous
above and beneath except villose along the costa and veins; petioles slender, to
0.2 cm wide, 0.2-2.0 cm long; stipules deltoid, narrowly or roundly so, 0.9-1,2
cm long, ca. 0.3 cm wide, pubescent. Inflorescences terminal, solitary, cymose
thyrsoid, to 16 cm long, to 6.5 cm wide, the peduncle to 7 cm long, stiff but thin,
1.0-1.5 mm wide, densely villose, the branches well spaced, the lowermost op-
posite or whorled, subperpendicular; bracts persistent, subulate, to 6 mm long,
the flowers secund. Flowers with the hypanthium subrotund, ca. 1 mm long,
pubescent, the calycine cup short, the teeth 5-6, vague or as distinct short points;
corolla white, the tube cylindrical, to 6 mm long, 0.8 cm wide in the middle,
glabrate outside and within, the lobes 5, oblong, acute, villose within; stamens
5-6, the anthers 0.8-1.5 mm long, acuminate at the apex, the acumen to 0.35 mm
long, acute, sagittate at the base, attached near the mouth; stigmas 2, oblong
lanceolate, ca. 2 mm long, the style slender, ca. 2.5 mm long, the ovary thick
walled, replete with resinous dots. Fruits sessile, rotund, 3.5-4.0 mm in diam.,
the pyrenes 6-costate, drying black; seeds obtusely 3-angled, ca. 1.5 mm in diam.,
muricate.

Bertiera guianensis ranges from Mexico to Panama, the Guianas, and south
to Bolivia. It also occurs in the West Indies.

BOCAS DEL TORo: Bucna Vista, Almirante, Cooper 586 (NY, US), barro Colorado island:
Avlles 9 (MO); Croat 12503 (MO); Foster 774 (MO), chiriqui: San Felix, Fittier 5178 (US), darien:
Airstrip. Cana, 480 m, Croat 37997 (MO). Cativo Swamp near Tusijuanda, 1 day up Rio Balsa, Duke
13533 (MO). San Jose River, Duke ct- Bristan 397 (MO, NY). Between Paya and Palo de Los Letras,
Duke & Kirkhricle 14030 (MO). Rio Tuquesa, middle Tuquesa Mining Co. Camp called Charco Peje,

Mori 7057 (MO). From Paya to Pucro, Stern et al. 225 (MO). Cana, Stern et at. 693 (MO). Panama:
Chiman, Lewis et al. 3243 (MO, UC, VEN).

11. BOROJOA

Borojoa Cuatrecasas, Serie Bot. Aplicada, Serv. Divulg., Seer. Agric. y Ganad.
(Call, Colombia) 2(5): 474. Aug. 1949. (Preprint of Rev. Acad. Colombia Ci.
7: 474. 1950.) type: B, patinoi Cuatrecasas.

Small dioecious trees. Leaves large, usually somewhat coriaceous, petiolate;
stipules partly connate basally, interstipular, often persistent. Inflorescences ter-
minal, pistillate flowers 6-8-merous; male flowers sessile, crowded, disposed in
a terminal head; pistillate flowers usually terminal, solitary. Staminatey/ouY^rv 4-
5-merous; corolla hippocrateriform or funnelform, pubescent outside and within,
the lobes contorted in bud; anthers 5, linear, dorsifixed; ovary 6-8-celled, the
carpels united along their margins, the ovules numerous, the stigmas 6-8. Fruits
baccate, large, rotund, the pericarp wall very thick; seeds embedded in a slimy
pulp, disposed in rows, horizontally attached.

1980]

DWYER— FLORA OF PANAMA (Family 179. Riihiacaic)

45

Figure 10. Bertlcra giiianensis Aubl.— A. Habit (x'/z).— B. Fruit (x3.5). [After Stern et al.

693.]

46 ANNALS OF THE MISSOURI BOTANICAL GARDEN |VoL 67

Borojoa is known from Panama, Colombia, Venezuela, and Brazil. It is a
genus of about 8 species.

Literature:

Cuatrecasas, J. 1950. Borojoa. Nuevo Gen. Rubiaceae. Aug. 1949; Revista Acad.

Colomb. Ci. Exact. 7: 474-477.
Patino, U. M. 1950. Noticia sobre el Borojo, una nueva especie frutal de la costa

Colombiana del Pacifico, Revista Acad. Colomb. Ci. Exact. 7: 478-489.

a. Lateral veins of leaves 6-12.

b. Lateral veins of leaves 10-12; fruits to 6.5 cm in diam 1. B. atlantica

bb. Lateral veins of leaves ca. 7; fruits to 4.5 cm in diam. 2. B. panamensis

aa. Lateral veins of leaves 13-20 3. B. patinoi

1. Borojoa atlantica Dwyer.^ type: Panama, Kennedy & Foster 2197 (MO, ho-
lotype).

Trees to 10 m tall, the branchlets terete, glabrous, the nodes well spaced.
Leaves elliptic or obovate elliptic, 18-27 cm long, 6-13.5 cm wide, widely
deltoid toward the apex, acuminate, the acumen triangular, 1-2 cm long, basally
obtuse, cuneate or truncate, the costa lightly porcate above, prominent beneath,
the lateral veins 10-12, arcuate, slender, the intervenal areas spreading reticulate,
concolorous, drying green, thin coriaceous, glabrous above, minutely puberulent
beneath; petioles 0.3-1.2 cm long, rigid or curved, puberulent; stipules (only one
seen) connate?, ovate, to 1.3 cm long, ca. 0.5 cm wide, acute, subcoriaceous, pu-
berulent. Inflorescences and flowers not seen. Fruits apparently sessile, rotund,
to 6.5 cm in diam., lignose, lightly wrinkled on the surface, lightly puberulent,
with many slender longitudinal grooves when dry, the wall thick, to 2 cm wide,
the persistent calycine cup umbonate, to 1 cm long; seeds many, embedded in a
fleshy pulp, subplane, often angular, to 1 cm long.

Borojoa atlantica is known only from Panama. It is readily distinguished from
B. panamensis by the more numerous lateral veins of its leaves and it is from B,
patinoi whose leaf blades have still more lateral veins (up to 20). The fruits of the
new species and those of B. patinoi are similar.

Borojoa atlantica seems closely related to B. universitatis Steyermark, the
type of which {Aristequieta 6466\ US, isotype) is from the Parque Nacional de
Guatopo, 650 m, Venezuela. The lateral veins of B. universitatis number about
18 and the wall of the large fruit (in the dried state) is only about 1 cm thick.

colon: 2-3 miles up from the Rio Guanche, Kennedy c& Foster 2197 (MO). Panama: El Llano-
Carti Road, Folsom 2620 (MO); Folsom & Collins 1517 (MO). Gorgas Memorial Labs, yellow fever
research camp, Campamento Quatro, 5-10 km NE, Altos de Pacora, 600 m, Mori A Kallunki 3336
(MO).

** Borojoa atlantica Dwyer, spec. nov. Arbores ad 10 m altae. Folia elliplica vel obovato-elliptica.
18-27 (vel plus) cm longa, 6-13.5 cm lata, apicem versus lato-deltoidea acumine triangulari, 1-2 cm
longo, basi obtusa cuneata vel truncata venis lateralibus 10-12, tenui-coriacea infra minuto-puberula
supra glabra; petiolis ad 1.2 cm longis; stipulis fortasse ovatis, ad 1.3 cm longis. Fructits fortasse
sessiles rotundi, ad 6.5 cm diam., pariete crasso, ad 2 cm lato, lignoso; seminibus multis in pulpa
carnosa inclusis subplanis oblongis saepe angularibus, ad 1 cm longis.

1980) DWYER— FLORA OF PANAMA (Family 179. Ruhiaceue) 47

2. Borojoa panamensis Dwyer, Phytologia 17: 446. 1968. type: Panama, Duke &
Lallathin 15014 (MO, holotype).

Small trees, the branchlets subterete, somewhat rimose, ultimately angular,
presumably glabrous, the nodes to 9 cm apart, the scars of the petiole prominent,
subrotund, ca. 0.4 cm in diam. Leaves elliptic rotund, to 19 cm long, to 12.5 cm
wide, widely deltoid at apex, short acuminate, cuneate and subequilateral at base,
the lateral veins ca. 7, arcuate, stiffly chartaceous, the lamina glabrescent, the
costa with many minute golden hairs and the axils of the lateral veins golden
barbate; petioles to 2 cm long, to 0.2 cm wide in middle; superior stipules per-
sistent, connate to the middle, the sheath cylindrical, to I cm long, the triangular
basal prominence to 6 mm long, ca. 0.5 cm wide, at maturity torn and refiexed,
the free portions obovate rotund to elliptic, to 2 cm long, to 1 cm wide, acuminate,
often wider above middle, thin coriaceous, dark brown, with many small veins
and with a median keel. Flowers not seen. Fruits terminal, sessile, solitary,
rotund, to 4.5 cm in diam., glabrous, minutely rugulose, the persistent calycine
cup like a volcano's crater, to 0.5 cm long, the orifice ca. 0.4 cm wide.

The species is known only from the type locality in Panama.
Duke & Lallathin note on their collection that the fruits are "green, probably
edible when ripe. Fruit, when sawed into, blackening as if possessed of genipine."

cocle: Cerro Pilon near El Valle, ca. 2500 ft, Duke & Lallathin 15014 (MO); Lallathin 5014
(MO).

3. Borojoa patinoi Cuatrecasas, Nuevo Gen. Rubiaceae. Aug. 1949; Revista
Acad. Colomb. Ci. Exact. 7: 474. 1950. type: Cuatrecasas 21137 (F, holo-
type). — Fig. 1 1.

Trees to 7 m tall, the branchlets smooth, plane, glabrous, the bark rimose, the
nodes well spaced, to 6 cm apart, the pedicel scars often prominent. Leaves
elliptic, to 36 cm long, to 17 cm wide, cuneate at the apex, basally cuneate,
truncate or obtuse, the costa prominulous above, prominent beneath, to 1.8 mm
wide, obviously porcate distally, the lateral veins 13-15, widely arcuate, to 3 cm
apart, usually 1-1.5 cm apart, the venules pinnatiform, spreading, discolorous,
subcoriaceous, glabrous; petioles to 4 cm long, glabrous; stipules tending to per-
sist, often reflexed at maturity, connate, appressed to stem, the sheath cylindrical,
the parts ovate elliptic to ovate, 2.5-4 cm long, to 1.2 cm wide, acute at the apex,
stiffly chartaceous with a slender, median keel, venose, the veins crowded, prom-
inulous. Inflorescences terminal, the male flowers numerous, disposed in a ter-
minal head about equal to the petiole; bracts disposed as 2, connate, decussate
pairs, soon spreading or reflexed; bracts subtending the solitary female flower,
the inner pair suborbicular, the outer pair similar in size and texture to the sti-
pules. Maie flowers 4-5-merous, the hypanthium and the calycine tube tubular,
0.7-0.8 cm long, glabrous, stiffly membranous, truncate, the teeth absent or
vague; corolla white, to 25 mm long, the tube hippocrateriform, to 18 mm long,
thickly petaloid, densely sericeous outside, the lobes 5, 6-7 mm long, oblong,
obtuse, somewhat shorter than the tube; stamens 5, the anthers linear, ca. 12 mm
long, the filaments slender, short; female flowers 6-7-merous; style to 7 mm long,
the stigmatic lobes 6, linear lanceolate, slightly longer than the style, acute, pap-

48

ANNALS OFTML: MISSOURI BOTANICAL GARDEN

[VoL 67

Figure IL Borojoa patinoi Cuatrecasas — A. Branch with leaves and fruit (x.35). [After Duke
MI8L\—B. Fruit cross section (x.35). [After Kirkhridc & Duke 407,]

illose. Fruits sessile, rotund, applelike, 7-8 cm in diam., subtended by the per-
sistent bracts, glabrous, the remains of the calyx represented by a delicate ring
bordering a terminal cavity, ca. 1 mm deep, the wall very thick, 1.2 cm in diam.;
seeds subplane, subovoid, 6-7 mm long, 7-12 mm wide, embedded in a pulp.

The above description has drawn freely on Cuatrecasas (1949) as the Panama
collections are sterile with one exception and this is in bud.

Borojoa patinoi is known only from Panama and Colombia. ''Borojo,'' ''Bo-
rojo Hembra,'' "Borojo Macho,'' or ''Burijo."

Duke in his Darien Ethnobotanical Dictionary states: 'This favorite fruit tree,
one fruit making a bowl of chicha, has followed the Choco (Indians) into Panama
(from Colombia), in whose flora it is unreported." Duke adds, speaking of him-
self: I am called Borojo among the Darien Choco, because for a while, I was

distributing borojo seedlings like Johnny Appleseed."* He adds that the fruits are
edible, are aromatic, and take more than one year to ripen.

DARir.N: Cerro Pirre, Bristan 495 (MO). Santa Fe, Duke & Bristan 310, 311 (both MO). Rio
Tucuti between Rio Tucutf and Rio Urugantf, Duke 5280 (MO). Without specific locahty, Duke 8332
(MO). Finca Othon near Yape, Duke 11820 (MO). Rio Morti, Drill Site 7, ca. 250 m, Duke 14181
(MO). Ri'o Punusa and Rio Pucro, Duke 14637 (MO). Rio Paya, 20 minutes from mouth of Rio Paya,
Kirkhride & Duke 407 (MO), san blas: Eslogandi, 200 m, Duke 10196 (MO).

12. BORRERIA

Borreria G. F. W. Meyer, Prim. FK Esseq. 79: 1818. nomen conserv. contra
Borreria Acharius (1810, Lichen), type: B, suaveolens G. F. W. Meyer, type

conserv.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaccae) 49

Tanlavel Adanson, Fam. PI. 2: 145. 1763. type: Rheede, 168. Hortus Indicus Malabaricus 9, (ah. 76.

(design. Standley 1916) = Borreria artUitlahs (L.f.) F. N. Williams.
Borrera G. F. W. Meyer, Prim. Fl. Esseq. 79. 1818. orth. mut. Sprengel, Gen. 1: 93. 1830.

Herbs, annual or perennial, rarely suffrutescent. Leaves with the petioles
united to the stipules forming a setiferous sheath. Inflorescences capitate, usually
sessile, axillary or terminating the stem. Flowers with the calycine lobes 2 or 4,
equal or unequal, often with interdental teeth, the corolla reduced, funnelshaped,
the lobes 4, the stamens usually attached near the mouth; ovary 2 celled with an
ascending ovule in each cell. Fruits capsular, coriaceous or membranaceous,
2-valved, the valves septicidally dehiscent; seeds 1 per cell.

Borreria occurs throughout the warmer parts of the world. It is a genus of
about 150 species.

Literature:
Steyermark, J.

W

831.

Highland— Part IX. Mem. New York Bot. Card. 23: 805-

a. Plants more than 1 dm tall.

b. Inflorescences with heads 1.1-2.0 cm in diam.

c. Leaves 0.1-0.25 cm wide or if wider, corolla lobes pilulose within

6. B, siiaveolens

cc. Leaves 0.6-3.2 cm wide.

d. Leaves 2 at a node, corolla lobes pilulose within; stipular sheaths pubes-
cent 2. B. laevis

dd. Leaves 5-6(-8) at some of the nodes, corolla lobes not pilulose; stipular

sheaths glabrous - 1. B- densiflora

bb. Inflorescences with heads (0.3-)0. 5-1.0 cm wide.

e. Inflorescences with heads axillary, or if terminal heads present, these much

reduced in diam.; stems villose; leaf blades usually drying yellow green or

flushed with red 3. B. latifoUa

ee. Inflorescences with heads both terminal and axillary, similar in diam.; stems
glabrate; leaf blades not drying yellow green or flushed with red.

f. Leaves 2 at a node.

g. Leaves 0.3-1 .0 cm wide; corolla ca. 1 mm long; fruits with one carpel

with 2 calycine lobes 4. B. odmoidcs

gg. Leaves (0. 6-) 1.0-3.0 cm wide: corolla ca. 2.5 mm long; fruits with

one carpel with 3 of the 4 calycine lobes — 2. B. laevis

ff. Leaves 5-6 at some of the nodes 1. B. densiflora

aa. Plants less than 0.6 dm tall.

h. Leaves to 6 mm long 5. B. puinilio

hh. Leaves 10-25 mm long 4. B. oeimoides

1. Borreria densiflora DC, Prodr. 4: 542. 1830. type: Jamaica, Swartz. not seen.

Spermacoce spinosa L. Sp. PI., ed 2. 148. 1762, nomen illegit. (cites S. suffrutieosuni Loefl.; see

Steyermark 1972).
S. spinosa Swartz., Obs. Bot. 45. 1791. not Jacq. 1760, nomcn nudum.

S. spinosa Cham. & Schl., Linnaea 3: 340. 1828.

Tardavel spinosa (L.) Standley, Contr. U.S. Natl. Herb. 18: 122. 1916.

5. radicosa Brandegee, Univ. Calif. Publ. Bot. 10: 416. 1924. type: Mexico, Brande^ee 9223 (CAL,

not seen).

Herbs to 0,5 m tall, the stems arising from a taproot, erect, decumbent or
prostrate, often the branches restricted basally and strict, occasionally with short
branches above, angular, smooth, glabrous or puberulent, the nodes well spaced.

50 ANNALS OF THE; MISSOURI BOTANICAL GARDEN [Vol. 67

Leaves 5-8 at some of the nodes, of variable length, some 3 times longer than
others, linear lanceolate to linear, to 6 cm long, to 0.8 cm wide, acute at the apex,
attenuate at the base, the costa prominent beneath, the lateral veins few and
scarcely distinguishable, papyraceous, stiff, smooth, the costa often with scat-
tered erect calcareous bodies, the margin scabrid, revolute; petioles absent or
seemingly present as the blades attenuate basally, the setiferous sheath cupuli-
form, 2-4 mm long, the setae 15-20 linear, as long as or longer than the sheath.
Inflorescences subtended by 4-8 leaves, terminal or axillary in upper axils, the
heads sessile, 1.5-2.0 cm in diam. with slender, white bristles among the flowers.
Flowers with the hypanthium oblong, the calycine cup ca. 0.3 mm long, petaloid,
glabrous, the calycine teeth 2, subulate or linear oblong, 1.3-1.8 mm long: corolla
white, cupular, 1.5-2.0 mm long, strongly constricted basally, the lobes 4, lan-
ceolate or oblong, to 1 mm long; stamens 4, exserted, the anthers oblong rotund,
0.3-0.5 mm long, the filaments slender, to 0.5 mm long, the style 1.3-1.7 mm
long, the stigmatic lobes minute. Fruits pyriform or obovate oblong, the 2 carpels
to 2.5 mm long, somewhat compressed, smooth, shiny, glabrous except pubescent
to arachnoid pubescent at the apex, the 2 awns erect, shorter than the body of
the fruit, pubescent.

This species is widely distributed throughout tropical America. For an elab-
orate discussion of '^Borreria dcnsiflora DC. (as) the earliest legitimate name for
the species now passing as B. spinosa,'" see Steyermark (1972) p. 814.

BOCAS DEL TORo: JuHCtion Rfos Changuinola and Teribe, Lewis el al. 923 (GH, MO, US). Isia
Colon, Wedel2946 (MO), canal zone: Fort Lorenzo, Baker ct Oliver 1043 (GH, K, MO, UC, US).
Panamerican Highway, La Chorrera, IT Any 9452 (MO). Toro Point, Fort Sherman, Duke 4324 (MO).
Fort Sherman, Dwxer 7191 (MO). Between Rodman Marine Base and Chorrera, Nowieke et al. 3606

(MO, SCZ, UC). Fort Amador, Txson 2017 (MO). Fort Sherman, Tyson 2248 (MO), barro Colorado
island: Croat 4407, 6984, 7394, 12609, 12884 (all MO); Shattuck 421 (MO), chiriqui: Methodist
Camp near Nueva Swissa, Croat 13525 (MO). Burica Peninsula, 12 mi W of Puerto Armuelles, 400-
500 m, Croat 22181 (MO). Llanos Francia, 4 mi from Boquete, Dwyer & HayJen 7612 (MO). Cerro
Punta and Bambito, Wilbur et al. 10902 (MO). Cerro Punta and Las Nubes, Wilbur et al. 13233 (MO).
colon: Mouth of Rio Piedras, Lewis et al. 3176 (MO), herrlra: Salinas de Chitre on Bahia Parita,
Croat 9683 (MO). Between Rfo Bayano Dam and Canitas, DArey 9400 (MO). Panama: Isla Taboga,
Allen 1272 (MO). Punta Chame, D'Arcy 10234 (MO). San Jose Island, Johnston 838 (MO). 2 km W
of El Llano, Nee 7958 (MO). Nuevo Emperador, Troetseh 8 (MO), san blas: Mandingo Airport,
Duke 14824 (MO), veraguas: Mouth of Rio Concepcion, Lewis et aL 2766 (COL, MO, UC, VEN).

2 km NW of Atalaya, Nee 8234. 8243 (both MO).

2. Borreria laevis (Lam.) Griseb.,^ Goett. Abh. 7: 231. 1857.— Fig. 12.

Spermaeoce laevis Lam., Tabl. Encycl. Meth. 1: 273. 179L rvPE: Santo Domingo, W.L, not seen.
Spermacoce evhioides H.B.K., Nov. Gen. Sp. PI. 3: 269. 1820. type: not seen.
Tanlavel laevis (Lam.) Standley, Contr. U.S. Natl. Herb. 18: 122. 1916.

Spermacoce guianensis Brem., Recueil Trav. Bot. Neerl. 33: 714. 1936. type: Surinam. Rombouts
75 (Utrecht?).

Herbs, upright or scandent, to V?, m in length, the stems smooth or rimose,
glabrous to glabrate, often retrorse pilose, the nodes well spaced. Leaves ovate
elliptic to elliptic, 1.5-7.0 cm long, 0.6-3.0 cm wide, acute at the apex, rounded
at the base, occasionally attenuate acute, often inequilateral, the costa promi-
nulous beneath, the lateral veins 5-6, strongly arcuate, chartaceous, concolorous.

** For more complete synonymy see Steyermark (1972) p. 819.

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

51

t.vt

Figure 12. Borrerla laevis (Lam.) Griseb.— A. Habit (x'/3).—B. Fruit(x8). [After Duke 4902.]

52 ANNALS OF THL MISSOURI BOTANICAL GARDEN [Vol. 67

scabrous above and beneath, glabrous above, glabrate beneath; petioles to 1 cm

long; stipules connate, the base of the petiole adnate to the sheath, the latter
cylindrical, appearing to be subnodal, to 4 mm long, puberulent to villosulose,

the awns several, narrowly subulate, 2-7 mm long. Inflorescences axillary and
terminal, globose capitate, 0.6-1.0 cm in diam. Flowers with the hypanthium
glabrous or puberulent, oblong, ca. 1.5 mm long, the calycine teeth 4, unequal,
subulate, to 0.5 mm long, ciliolate; corolla cylindrical, 2.5 mm long, the lobes 4,
longer than the tube, pilose within; stamens 4-5, the anthers subrotund, ca. 0.3
mm long, the filaments 1.3 mm long; style to 2 mm long, the stigmas rotund
capitate, ca. 0.25 mm in diam. Fruits oblong, 1-2 mm long, glabrous to minutely
pubescent, dehiscent, one carpel bearing 3 of the 4 calycine lobes; seeds oblong,
1.0-1.5 mm long, 0.75 mm wide, the transverse grooves prominent.

Borreria laevis is a common weed in Central America and in most of South
America. It also occurs in the West Indies. ''Sab Tutu" (Cuna, Duke 14439),

BOCAS DEL TORo: Isia Bocas del Toro. Durkee 71-155 (MO). Without other locality, Wedel 243
(MO). Water Valley, Wedel 595, 991 (both MO). Isla Colon, Wedel 2833 (MO), canal zone: Fort
Amador, DArcy 9328 (MO). Between Rodman Marine Base and Chorrera, Nowicke et al. 3585 (MO,
SCZ, UC). Ancon Hill, Seihert 107 (MO). Fort San Lorenzo, Tyson & Blum 3701 (MO). Ancon Hill,
Woodson et al. 1325 (MO). Salamanca Hydrographic Station, Woodson et (d. 1617 (MO), barro
COLORADO island: Croat 4045, 4188, 4644, 5796, 6078, 6946, 7047. 7060, 7064, 7322, 9199 (all MO);
Ehin^er 48 (MO); Havden 81 (MO); McDaniel 5025 (MO); Netting 49 (F); Shattuck 478 (MO); Starry
26. 74 (both MO), chiriqui: N of Cerro Punta, Croat 10448. 10471, 10508 (all MO). I mi N of El
Hato del Volcan, Croat 10703 (MO). Road to Rio Serano, Folsom et al. 2093 (MO). Boquete to 3 mi
N, 3300-4200 ft, Lenis et al. 630 (GH, K, MO, NY, US). 11 mi W of Puerto Armuelles, Liesner 82
(MO). Puerto Armuelles, Liesner 140 (MO). Upper Rio Chiriquf Viejo, Monte Lino, Seihert 247
(MO), cocle: Behind Club Campestre, Di4ke 13273 (MO). NE of El Valle, Lewis et al. 1698 (MO,
UC). Between Las Margaritas and El Valle, Woodson et al. 1288 (MO), colon: Ridge behind Garotte,

D'Arcy 9332 (MO). Achiote, Zimmerman, Folsom et al. 3322 (MO), darien: Boca de Cupe, Allen
984 (MO). El Real, Bnrch et al. 1076 (COL, MO, UC). S of El Real, Duke 4902 (MO). From Pucro

to Rio Pucro, Duke 5375 (MO). 10 mi S of El Real on Rio Pirre, Duke 5444 (MO). 1-4 mi N of Pucro,
Duke 13019 (MO). Rio Pucura between Cerro Mali and Cerro Tacarcuna, near Rio Pucro base camp,
640 m. Gentry & Mori 13951 (MO). Paya, Stern et al, 259 (MO), los Santos: Loma Prieta, Duke
11863, 11895 (both MO). 25 mi SW of Tonosi, Lewis et al. 2983 (MO, UC). Panama: Near Jenene,
Duke 3831, 3847 (both MO). Piria, ca. 150 m, Duke 14439 (MO). 3 mi E of El Llano, Tyson 1723
(MO). Road to Cerro Campana, 7.6 km from Panamerican Highway, Witherspoon & Witherspoon
8366 (MO). SAN BLAs: SE of Puerto Obaldia, Croat 16736 (MO). Puerto Obaldia, Gentry 1555 (MO).

3. Borreria latifolia (Aubl.) Schum. in Mart., Fl. Bras. 6(6): 61. 1888.

Spennacoee latifolia Aubl., Hist. PI. Guiana 55. 1775. type: Cayenne, French Guiana, Auhlet, not

seen.
Borreria perrottetii DC, Prodr. 4: 548. 1830. typl: not indicated.
Tardavel latifolia (Aubl.) Standley, Contr. U.S. Natl. Herb. 18: 122. 1916.

Herbs, often prostrate, often erect and more than 1 m tall, the stems villose,
porcate, the nodes well spaced. Leaves with the blades often spreading, disposed
at right angles to the stem or deflexed, trapeziform, ovate oblong, oblong rotund
or oblong, 1,5-6.0 cm long, 0.7-3.2 cm wide, acute at the apex, the costa prom-
inulous beneath, the lateral veins ca. 5, immersed above, prominulous beneath,
chartaceous, scabrous, often with white bulbous-based hairs above, drying yellow
green or flushed with red; petioles absent or to 0.5 cm long; stipules with the
sheath subtriangular in outline, often split as the inflorescence expands, the setae
to 5 mm long. Inflorescences with the flowers disposed in rotund heads in the

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 53

axils, these usually to I cm wide. Flowers with the hypanthium short, ca. 1 mm
long, densely appressed white villose to glabrous (fide Standley), the calycine
cup much shorter than the lobes, the lobes 4, lanceolate to ovate, to 1.5 mm long,

acute, villose; corolla white, the tube funnelshaped, 2.5-3.0 mm long, glabrous,
the lobes 4; stamens 4, the anthers oblong or ovate, 0.35-0.50 mm long, the
filaments at least twice as long as anthers, attached at the mouth; style minute,
knoblike or conspicuously bifid, the ovarian disc scarcely developed. Fruits ob-
long or turbinate, to 3 mm long, densely villose, opening apically and often ex-
panding into 2 valves, each valve circular in top view, the smooth light-colored
interior of the valve revealed; seeds oblong, ca. 2 mm long, smooth, drying
orange, shiny, reticulate, the calyx tending to wither at dehiscence.

Borreria latifolia is widely distributed in tropical America. It also occurs in
the West Indies. ''Cansa-mozo'' (Standley, 1928).

Steyermark (1972) has 2 varieties and several forms which do not hold up as
far as the Panamanian material is concerned. In the synonomy of B, latifolia I
have included only a minimum of synonomy as compared with Steyermark's
many synonyms.

CANAL zone: Panamerican Highway near La Chorrera, DArcy 9432, 9453 (both MO). Ft. Kobbe,
Duke 3905 (MO). E of Gatiin Locks, Duke 4306 (MO). 5-9 mi N of Gamboa, Gentry 2408 (MO).
Chiva-Chiva Trail, Tyson 1370 (MO). Curundu, Tyson 6288 (MO). Miraflores Lake, White 253 (MO).
BARRO COLORADO ISLAND: Croat 5982. 6461 , I28I6, 12824 (all MO), chiriqui': Palo Alto, E of Bo-
quete. Stern et al. 1042 (MO), colon: Sardinilla, 7-8 mi E of cement plant, Blum d Tyson 497 (MO).
0.7 mi NE of Rio Piedras, Nee & Mori 3641 (MO). Panama: Arraijan, Duke 4681 (MO). Cerro Jefe,
Gentry 2138 (MO). San Jose Island, Johnston 434, 1085 (both MO). Goofy Lake, Lewis et al. 229
(GH, MO, UC, US). Chiman, Lewis et aL 3306 (MO, PAN). El Llano-Carti Road, 5 km N of
Panamerican Highway, Nee 7879 (MO), vlraguas: 1-2 mi above Santa Fe, Gentry 3063 (MO).

4, Borreria ocimoides (Burm. f.)^*' DC, Prodr. 4: 544. 1830.

Spermacoce ocimoides Burm. f., Fl. Ind. 34. 1768. type: Tropical Asia.

Borreria parviflora G. F. W. Mey., Prim. Fl. Esseq. 83. 1818. type: not seen.

B. tampicana DC, Prodr. 4: 544. 1830, in part, type: Mexico, Berlandier 37 his (?B, not seen; photo,

MO).
B, pringlei Wats., Proc. Amer. Acad. Arts 25: 152. 1890. type: Watson 2464 not seen.
Tardavel ocymoides (Burm.) Heirn, Cat. PI. Welw. 2: 503. 1898.

Terrestrial herbs, often procumbent, rarely epiphytic on tree roots, to V3 m
tall, the stems simple or branched, angular, delicately alate, glabrous, the nodes
well spaced. Leaves lance ovate or ovate trapeziform, 1.0-2.5 cm long, to 1 cm
wide, acute or occasionally obtuse at the apex, basally acute to cuneate, the costa
immersed, the lateral veins 4-5, strongly arcuate ascending, concolorous, sca-
brous above and beneath; petioles absent or to 0.15 cm long; stipules connate
and adnate to the base of the petiole, the sheath cupshaped, scarcely measurable,
to 1.5 mm long, the setae 6-10, to 2.5 mm long, usually curled. Inflorescences
terminal and axillary, sessile, capitate, 0.2-0.8 cm wide. Flowers minute, the
calycine lobes 2-3(-4), erect, subulate, ca. 1 mm long; corolla white, to 1 mm
long, the lobes 4, acute; stamens 4, the anthers minute. Fruits capsular, the

^^ For more extensive synonymy see Steyermark (1972). Only these names have been used for
Panamanian material.

54 ANNALS OF THH MISSOURI BOTANICAL GARDEN [Vol. 67

valves 2, splitting and separating, the persistent calycine lobes glabrous or pu-
berulent; seeds oblong, ca. 1 mm long, brown, reticulate.

Borrena oclmoides is a widespread weed in the tropics.

CANAL zone: Military Road, K-9, Ehingcr 513 (MO). Gaillard Highway, V2 km NW of Summit
Garden, Nee 6929 (MO). Salamanca Hydrographic Station, Woodson et al. 1619 (MO), barro Col-
orado island: Croat 8268, 8303, 8826. 10396, 11339, 12277, 12582, 12938. 13152, 17037 (all MO);
Shattuck 47, 497 (both MO), chiriqui': Districto Guanabano, along Quebrado Quanabano, Croat
22510 (MO). Boquete, 3300-4200 ft, Davidson 597 (MO); Lewis et al. 334 (GH, K, MO, NY, UC,
US); Partch 69-76 (MO); Woodson & Schery 729 (MO). Cerro Punta, 6000 ft, Tyson 7128 (MO).
cocle: EI Valle, Allen 2825 (MO). La Mesa, Gentry 6826 (MO), darien: El Real, Ri'o Tuira, Stern
ct al. 774 (MO). Panama: Between Panama and Chepo, Dod^e et al 16698 (MO). Hacienda la Joya,
Dodge et al. 16891 (MO). Cerro Campana, road near summit, Duke 6000 (MO). Confluence Rio
Pacora and Rio Corso, ca. 450 m, Duke 11957 (MO). Goofy Lake, Dwyer 7137 (MO). San Jose Island,
Johnston 902 (MO). Cerro Campana, Lewis et al. 3103 (COL, DUKE, F, K, MO, NY, UC). W slope
of Cerro Campana 2500 ft, Tyson et al. 2349 (MO). 2 mi S of Tocumen Airport, Tyson & Clewell

5929 (MO).

5. Borreria pumilio Standley, Ann. Missouri Bot. Garden 27: 342. 1940. type:
Panama, Allen 822 (F, holotype, isotypes MO, NY, US).

Herbs to 6 cm tall, unbranched or with 1-2 branches, the stem with appressed
white hairs. Leaves sessile, linear oblong, 4-6 mm long, the costa and veins not
visible or the lateral veins 4-5, scarcely visible beneath, arcuate, granulose above

r

and beneath; stipular sheath ca. 1 mm long, the setae several, to 1 mm long.
Inflorescences capitate, terminal or in the uppermost axils, the heads ca. 0.5 cm
in diam., the outermost bracts linear, to 9 mm long. Flowers with the corolla
white, the tube somewhat longer than the calycine lobes. Fruits subrotund, each
mericarp oblong, ca. 0.8 mm long, the persistent calycine lobes 1.0-1.5 mm long,
glabrous to pubescent, usually slightly longer than the body of the fruit.

Borreria pumilio is known only from Panama. Because of the scant material,
the description of flowers is taken from Standley's diagnosis.

CANAL zone: Fort Kobbe, Curatella Savanna, Duke & Massell 6573 (MO), cocle: Nata, Allen
822 (F. MO, NY. US).

6. Borreria suaveolens G. F. W. Meyer, Prim. Fl. Esseq. 81. 1818. type: Surinam,
in Arenosis Continentis Circa Arouabischkreck, Meyer, not seen.

B. tenella Cham, & Schlecht., Linnaea 3: 317. 1828. type: Brazil, Sello, not seen.

B. tenella var. suaveolens (Meyer) Schum.. Mart., Fl. Bras. 6: 56. 1888.

B. capitata var. suaveolens (Meyer) Steyermark, Mem. N.Y. Botanical Garden 23: 825. 1972.

Perennial herbs, smaller plants somewhat cricoid in habit, the stems often
arising from a taproot, frequently unbranched to the apex, densely pubescent to
glabrate, the nodes well spaced. Leaves sessile or subsessile, 2(-6) at a node, the
shorter leaves linear or narrowly ovate lanceolate, 2-6(-8) cm long, to 0.25(-0.5)

cm wide, occasionally falcate, acute at the apex, rounded or obtuse at the base, the
costa prominent beneath, the lateral veins ca. 4, strongly ascending, usually ob-
scure, stiffly papyraceous, usually scabrid, the margins revolute; stipules with or
without a distinct sheath, the latter to 2 mm long, with 4-8 setae on each side,
these 3-5 mm long, rubescent. Inflorescences globose capitate, the heads ter-

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 55

minal, axillary in the 2 superior nodes, to 1 cm in diam., the leaflike bracts
reflexed in older heads, the bracts densely aggregated in the head, linear subulate,

3.5 mm long. Flowers with the corolla funnelshaped or subcylindrical, 3.5 mm
long, glabrous outside; stamens 4, the anthers oblong, ca. 1 mm long, the fila-
ments 1.2-1.5 mm long, attached above the middle of the tube; style slender, 2-

3 mm long, the stigmatic area minute, capitate, ca. 0.2 mm long. Fruits oblong,
1.5-2.0 mm long, the calycine lobes often persistent, as long as the body of the

fruit, the latter splitting at the pubescent apex; seeds not sulcate.

Borrcrla suaveolens is well distributed throughout tropical America; it is a

common species in savannas from low to relatively high elevations.

CANAL zone: Rio Indio, Allen 17395 (MO). Drowned Rio Azote Caballo, Dodge et al. 16833
(MO). Ri'o Indio, Dod^t^c & Allen 17377 (MO). Ancon Hill, Standley 26206 (MO), chiriqui: Volcan
de Chiriqui, Allen 989 (MO). S of Boquete, Allen 4691 (MO). 1 mi N of EI Hato de Volcan, Croat
10700 (MO). Cerro Colorado, 20-28 mi from San Felix, 1200-1500 m, Croat 33385 (MO). Boquete,
D'Arey 9773B (MO). El Barii above Boquete, 1200-1800 m, D'Arey 9928 (MO). Lava flow between

El Hato and Bambito, D'Arey 10025 (MO). Boquete, Davidson 682 (MO). 3 km NE of El Hato del
Volcan, 1500-1800 m, Davidse & D'Arey 10383 (MO). Volcan, Duke 9153. 9184 (both MO). Llanos
Francia, near Boquete, Dwyer s.n. (MO). !4 mi S of Bambito, Lazor & Correa A. 2700 (MO). Volcan

Baru, ca. 2000 m, Mori & Bolten 7385 (MO). Boquete, Partch 69-37 (MO), Llanos Francia, Stern et
al. 1155 (MO). Cerro Punta, ca. 6000 ft, Tyson 7128 (MO). Rio Chiriqui Viejo Valley, El Volcan,
White 209 (MO). Panama: Rio Pacora and Chepo Highway, Duke 5925. 5926 (both MO). Cerro
Campana, Ehinger 364 (MO); Sullivan 436 (MO); Tyson et al. 2383 (MO). Rio Tartare, Woodson &
Schery 1009 (MO), veraguas: Sona, Allen 1050 (MO).

13. BOUVARDIA

Bouvardia Salisbury, Parad. Lond. pL 88, 1807. type: 5. triphylla Salisbury,

Aeginetia Cav., Anales Ci. Nat. 3: 129. 1801; Ic. 6: 51. 1801, not Aeginetia L, (1753).

Shrubs or herbs. Leaves opposite or 3-4(-6) per node, membranous; stipules
interpetiolar with a basal sheath. Inflorescence terminal, cymose. Flowers di-
morphic, calycine lobes 4, persistent; corolla tubular or salverform, the lobes 4,
valvate; stamens 4, the anthers linear or oblong, in pin flowers included and

subsessile, in thrum flowers included and with filaments distinct and exserted;

styles of 2 lengths, usually slender, the ovary 2-celled, the ovules numerous,
attached peltately to a complete septum. Fruits capsular, globose or oblate, de-
hiscing at first loculicidally, then septicidally; seeds numerous, vertically imbri-
cate, winged, the wing entire.

Bouvardia. a genus of about 50 species, ranges from Mexico to Panama.

Literature:

Blackwell, W. H. 1968. Revision of Bouvardia. Ann. Missouri Bot. Gard. 55
1-30.

1. Bouvardia glabra Polak., Linnaea 41: 565. 1877. type: Polakowsky 337 (F,
frag, holotype, photo F, US). — Fig. 13.

B. i^lahra var. ^raedis Polak., Linnaea 41: 566. 1877. type: Polakowsky 172 (not seen; F, photo).
B. glabra var. obtusa Loesner, Verh. Bot. Ver. Brandenberg 65. 106. 1923. type: Seler 2920 (not
seen).

56

ANNALS OF THE MISSOURI BOTANICAL GARDEN

I Vol. 67

Figure 13. Bouvardla glabra Polak. Flowering branch (x.94). [After Mori cS Bolton 7356.]

Shrub to 1!^ m tall, the branchlets smooth, glabrous, drying tan. Leaves
tending to persist at the apex of the twiglets, ovate lanceolate to widely elliptic,
3.5-5.0 cm long, 1-2 cm wide, acute at the apex, acute at the base, the costa
subplane above and beneath, the lateral veins 3-6, strongly ascending and ar-
cuate, glabrate, chartaceous, discolorous, brown above, greenish beneath; peti-
oles slender, to 3 mm long; stipules triangular with 3-5 subulate processes, the
median process more elongate, often longer than the body. Inflorescences 3-7-
flowered. Flowers pedicellate, the pedicels to 1.5 cm long; hypanthium glabrous,

1980) DWYER— FLORA OF PANAMA (Family 179. Ruhuucae) 51

drying dark, the teeth 4, falcate subulate, unequal, 3-7 mm long; corolla tube
cylindrical, to 2 cm long, to 0.2 cm wide in the middle, glabrous outside, sparsely
pilose above the middle, the lobes ovate oblong, to 5 mm long, to 4 mm wide,
obtuse; anthers (? all) subsessile, 3 mm long, ca. 1 mm wide, obtuse; style (only
longer styles seen) to 2.3 cm long, the stigmas to 4.5 mm long. Fruits on pedicels
to 1.7 cm long, to 0.7 mm wide, glabrous, compressed rotund, to 0.8 cm long,
glabrous.

BouvanUa glabra ranges from southern Chiapas, Mexico, to Guatemala, Costa
Rica and Panama.

CHiRiQUi: W side of Volcan Baru, oak forest, 2000-2500 fl, Mori & Bolten 7356 (MO).

14. CALYCOPHYLLUM

Calycophyllum DC, Prodr. 4: 367. 1830. lectotype: C candidissimum (Vahl)
DC. {Macrocnemum candidissimum Vahl).

EnkyHsta Benth. Hooker's J. Bot. Kew Card. Misc. 5: 230. 1850. type: E. spruceana Benth.

Trees, Leaves petiolate, the stipules small, deciduous. Inflorescences pa-
niculate, terminal, corymbose, ebracteolate or bibracteolate. Flowers in bud en-
closed by membranous bracts, sessile or subsessile, the hypanthium oblong, the
calycine cup truncate, sometimes partly expanded into a large foliaceous stalked
blade, the corolla with the tube short, funnelform, the mouth villous, the lobes
6-8, imbricate or contorted, 1 lobe exterior, the stamens exserted, the anthers
versatile, the filaments subulate, the style slender, the stigmas 2, narrowly oblong,
the ovules disposed in 2 cells, numerous, imbricate, the placentas adnate to the
septum. Fruits capsular, oblong cylindrical, 2-celled, septicidally bivalvate, the
valves coriaceous; seeds numerous, imbricate, usually minute, the testa mem-
branous, produced at each end as an elongate wing.

Calycophyllum has approximately 6 species in the West Indies and 1 in Central
America and South America.

1. Calycophyllum candidissimum (Vahl) DC, Prodr. 4: 367. 1830.— Fig. 14.

MacrocncmuDi cancHdissiniuni Vahl, Symb. Bot. 2: 3. 1791. type: not seen.

Mussaenda Candida Poir. in Lam. Encycl. Melh. Bot, Suppl. 4: 395. 1797. typf: not seen.

Mussacnda candissima (Vahl) R. & S. Syst. Veg. 5: 251. 1819.

Trees to 15 (-40) m tail, the bark reddish brown, often exfoliate, the twiglets
smooth, glabrous, rubescent. Leaves oblong rotund, obovate oblong and occa-
sionally falcate, to 15 cm long, to 7 cm wide, obtuse at the apex, cuspidate, the
cusp usually triangular, to 2.5 cm long, ca. 0.3-0.5 cm wide in the middle, basally
attenuate cuneate or widely cuneate, often slightly inequilateral, the costa plane
or immersed above, prominulous beneath, the lateral veins 8-10, arcuate, the
intermediate veins parallel and branching near the margin, the intervenal areas
spreading reticulate, papyraceous, often drying red brown, glabrescent or villos-
ulose above and beneath, rarely with minute white raphides; petioles 3 (-4) cm
long, slender, to 1 .3 mm wide: stipules deciduous, oblong, ca. 2.5 cm long, obtuse,
scarious, rubescent, villose. Inflorescences terminal (sometimes appearing axil-

58

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol, 67

Figure 14. CalycophyUum candissimum (Vahl) DC. — A. Habit (X-/5). — B. Opened flower
and leaf (x2). — C. Ovary cross section (x4). [After MUlcr 1962,]

lary), paniculate, rarely shorter than the uppermost leaves, often flat topped or
convex at the apex, to 20 cm long, usually with 3-4 peduncles arising from the
tip of the twiglet, these strongly ascending, each with several strongly ascending
branches, the flowers disposed in compound dichasia, the median flower of each
cluster sessile. Flowers with the hypanthium turbinate, ca. 3 mm long, ca. 1.5
mm wide, truncate, glabrescent or villosulose with farinose glandular dots inter-
spersed, the calycine cup less than 1 mm long, the teeth absent or often one
expanded into a petiolate lamina, obovate spatulate, the blade to 3.3 cm long, to
2.5 cm wide, petaloid, glabrous, venose, the claw slender, ca. 0.2 cm wide; corolla
white, the tube compressed cylindrical, ca. 3.3 mm long, densely villose within

I980J DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 59

above the middle, the lobes 5, strongly reflexed, oblong, 3-4 mm long, obtuse,
glabrous, venose; stamens 5, the anthers oblong, ca. 1.5 mm long, subversatile,
the filaments slender, elongate, to 3 mm long, exserted, attached at the mouth;
ovarian disc pulviniform, pubescent, 0.75-1 mm long, 0.4-0.6 mm wide, the style
3.3-5.3 mm long, ca. 0.15 mm wide, the stigmas 2, slender, ca. 2.2 mm long,
curved or erect, puberulent, the ovary rotund in cross section, 2 loculate, the
septum moderately thick, the ovules numerous. Fruits sessile or subsessile, ob-
long, 0,5-1 cm long, truncate or rounded apically, usually obtuse or rounded at
the base, lignose, villosulose, often with minute white raphides, drying tan, with

ca. 8 delicate ribs, splitting from the apex, the margins of the valves wide; seeds
to 5 mm long, the wing acuminate at each pole.

Calycophyllum candidissimum is widely distributed in Mexico and Central
America, ranging south to Colombia and Venezuela. It also occurs in the West
Indies. The hard, fine-grained wood has a variety of uses. In Panama it is used

for charcoal. Standley (1928, p. 355) lists 5 common names: ''Alazano,'' ''Gua-
yabo,'^ ^^Harino,'' ^'Madrono," and ^^Salamo.''

CANAL zone: Albrook, Blum 2086 (MO). Chagres River, Boyd-Roosevelt Highway, Blum &
Tyson 1981 (MO). Madden Dam, DoJ^n^ et al. 16500 (MO); Duyer 8380 (MO); Lewis ct al. 35 (F,
GH, K, MO, NY, UC, US). Gatiin, Johnston 1736 (MO). Boy" Seoul Camp Road, Madden Dam,
Mori 4047 (MO). 1 km S of Madden Dam, Nee 8905 (MO). Rodman Naval Ammunition Depot, 1 mi
SW of Cocoli, Wilbur et al, 12891 (MO), barro Colorado [SLand: Avilcs 60 (F); Croat 8661, 12905
(both MO), cocle: El Valle, A^uilar (MO). Rio Mata Ahogado, Allen 126 (MO). Cerro Pilon, Lal-
lathin 113 (MO), darien: Santa Fe, Rio Sabanas, Duke 10226 (MO). Punta Garachine, Duke 10472
(MO). Hydro Pico, Pico Pendejo, Rio Sabanas, Duke 15434 (MO). Rio Chucunaque between Yaviza
and junction with Tuira, Gentry & Mori 13483 (MO). Campamento Buena Vista, Rio Chucunaque,
Stern et al. 966 (MO). Piriaque Camp, Tvson et aL 4761 (MO), los santos: Tonosf, Duke 12486
(MO); Oliver et al. 3544 (MO); Stern et aL 1816 (MO). Panama: Pacora, Allen 1122 (MO). Isia El
Rey, Correa 113 (MO). Cerro Azul, Croat 13007 (MO). Old Panama, Cooper 339 (MO); Greennian
& Greenman 5142 (MO). Isia Espiritu Santo, Duke 10437 (MO). Bejuco, Duke 4581 (MO). Saboga
Island, Miller 1962 (MO, US); Tyson & Loftin 5125 (MO). Chiva-Chiva Road, Smith & Smith 3340
(MO). W of Chepo, Tyson 5369 (MO).

15. CEPHAELIS

Cephaelis Swartz, Prodr. 3: 45. 1788. Nomen conserv. type: C. mascosa (Jacq.)
Swartz type cons.

Carapichea Aubl., Hist. PI. Guiane 1: 167. 1775. Nomen rejic. type: C. guianensis Aubl.
Tapo^otuea Aubl., Hist. PI. Guiane 1: 367. 1775. Nomen rejic. type: T, violacea Aubl.
Even Aubl., Hist. PI. Guiane 1: 103. 1775. Nomen rejic. type: E. i^uianensis Aubl.
Ipecacuanha Arruda, Diss. PI. Brazil 44. 1810. type: /. officinalis Arruda.

Shrubs, trees, or occasionally woody herbs. Leaves often glabrous and with
many lateral veins; stipules free or connate, frequently persistent. Inflorescences

axillary or terminal, capitate, the head often branched, subtended by an involucre
of bracts, the bracts often large, often highly colored, free or connate, becoming
smaller and more concave adaxially. Flowers usually sessile; hypanthium short,

the calycine cup short, truncate or dentate, often irregularly so, the teeth (4-)5
(-7); corolla usually white, the indumentum variable, the tube funnelform or
salverform, usually expanded basally, the lobes usually 5, valvate in bud; ovary
2-celled, the septum thick, a single ovule in each locule arising from the base of

60 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

the ovary. Fruits fleshy, the 2 pyrenes smooth or costate, the endocarp usually

sulcate on the inner face.

World

about 200 species. I have elected to separate Cephaelis from Psychotria although
there are reasons for reducing the former to the synonomy of the latter, as J.
Steyermark maintains. In Panama most of the species have capituliform inflores-
cences with large colored bracts which allow for easy allocation to Cephaelis,
Standley & Williams in the Flora of Guatemala (1975) sum up the traditional
stand: '\ . . for practical purposes ... it constitutes a most convenient segre-
gate.^^

a. Inflorescences spiciform cylindrical, the rachis 5-12 cm long 7. C. dimorphandhoidcs

aa. Inflorescences mostly rounded capitate, not spiciform cylindrical, the rachis or flowering
capitulum less than 5 cm long,
b. Inflorescence terminal.

c. Heads of flowers subtended by 2 large flaring or mottled bracts obviously distinct
in color and texture from the innermost bracts; outer bracts 2-5 cm long.
d. Heads with the outermost bracts not mottled, uniformly red, occasionally

orange or yellow, rarely white.

e. Plants densely hairy; heads on erect peduncles 20. C tomentosa

ee. Plants glabrale; heads on nodding or erect peduncles.

f. Peduncles nodding; leaves to 29 cm long, papyraceous

4. C. corrcae

ff. Peduncles erect, the largest leaves to 15 cm long, coriaceous

10. C. data

dd. Heads with outermost bracts obviously mottled, of various colors

21. C vultusmimi

cc. Heads of flowers with outermost bracts more or less similar in texture and color
to those within, less than 2 cm long,
g. Heads of flowers disposed as a rotund capitate or subglobose mass.

h. Subshrubs less than 1 m tall; mass of flowers 3-5 cm wide; petioles

4-7 cm long.

i. Peduncle ca. 4 cm long 17. C. nana

ii. Peduncle 0.7-2.0 cm long 8. C, discolor

hh. Subshrubs ca. 2 m tall; mass of flowers to 2.5 cm wide; peti-
oles 0.3-1.5 cm long 12. C. glomendata

gg. Heads of flowers oblong or elliptical, not disposed as a rotund capitate or

rotund mass.

j. Heads ca. 0.5 cm long - 15. C. kennedyi

ij. Heads more than 0.5 cm long.

k. Leaves with 2 submarginal veins paralleling the margin; stem

rhizophorous 13. C. insucta

kk. Leaves with 1 vein or no vein paralleling the margin; stem not

rhizophorous.

I. Leaves drying red or brown beneath, the lateral veins

(20-)30-38.

m. Petioles absent.

n. Anthers to 2.2 mm long 9. C. dresslcri

nn. Anthers 2.5-3.0 mm long

19. C. panchoco'cnsis

mm. Petioles of largest leaves ca. 4 cm long

2. C. bristanii

II. Leaves not drying red or brown beneath (except C. dres-
slcri), the lateral veins 12-25.

o. Peduncle present.

p. Peduncle terminated by 2 large bracts and 3

branches, each branch terminated by a ca-
pitulum of flowers; lateral veins ca. 12

6. C dichroa

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaccac) 61

pp. Peduncle and general inflorescence not as

above; lateral veins 13-25.
q. Leaves sessile or subsessile, auriculate

at the base; stipules 2.0-2.8 cm long.

r. Bracts glanduliferous within, the
glands diffuse or disposed in se-
ries; corolla 10-14 mm long

9. C. dressleri

rr. Bracts not glanduliferous within;
corolla ca. 30 mm long

19. C. panchococnsis

qq. Leaves with distinct petioles 0.3-1.5 cm

long; stipules to 0.3 cm long,
s. Inflorescence with heads 1.5-3.0
cm wide; calycine teeth to 1.2 mm

long, equal; corolla tube ca. 10

mm long ^_ 2. C. ^lonicrulata

ss. Inflorescence with heads to 0.5 cm
long; calycine teeth absent or dis-
posed as mere points or obviously
unequal, to 0.3 mm long; corolla
tube to 3.2 mm long

15. C. kennedxi

DO. Peduncle absent 18. C. panamensis

bb. Inflorescence axillary.

t. Leaves with lateral veins 6-12.

u. Stipules membranous with conspicuous marginal subulate fimbriae; plants

rhizomatous 14. C. ipecacuanha

uu. Stipules subcoriaceous or chartaceous without marginal fimbriae; plants

not rhizomatous.

V. Leaves succulent, the lateral veins evanescent; stipules to 1 cm long;

inflorescences with peduncle to 1 cm 11. C iiaii^eri

vv. Leaves not succulent, the lateral veins conspicuous; stipules to 7 mm

long; inflorescences with peduncle absent.

w. Leaves 1.6-5.0 cm wide; inflorescences 1.0-1.5 cm wide

1 . C axillaris

WW. Leaves 7-9 cm wide; inflorescences ca. 2 cm wide

16. C. latistipulata

tt. Leaves with lateral veins 15-38.

X. Stipules more than 1 cm long; inflorescences with heads 1.5-2.2 cm wide.

y. Leaves with lateral veins evanescent; blades succulent

3. C. caniponutans

yy. Leaves with lateral veins obvious; blades not succulent __ 5. C. croatii
XX. Stipules less than 1 cm long; inflorescences with heads ca. I cm wide

2. C. bristanii

1. Cephaelis axillaris Swartz, Prodr. Veg. Ind. Occ. 45. 1788. type: not seen.
Not Psychotria axillaris Willd. 1798.

Tapoi^omea axillaris (Swartz) Poir., Encycl. Meth. Bot Suppl. 7: 585. 1806.
Evea axillaris (Swartz) Standley, Contr. U.S. Natl. Herb. 18: 123. 1916.

Psychotria auhlctiana Steyermark. Mem. New York Bot. Card. 23: 694. 1972. New name for
Cephaelis axillaris Swartz.

Shrubs to IVi m tall; branches few, usually arising near the middle of the
stem, terete, smooth, glabrous, often attenuate toward the nodes, dilated at the
node above the base of the stipule. Leaves usually salicoid, occasionally oblong
to obovate oblong, occasionally falcate, 2-13 cm long, 1.6-5.0 cm wide, at least
2 times longer than wide, occasionally wider below the middle, the costa plane
or grooved above, prominulous to prominent beneath, golden villosulose or gla-
brous, the lateral veins ca. 8, arcuate, prominulous, the intervenal areas spreading
reticulate; petioles 1.5-2.0 cm long, slender, to 0.8 mm wide or stout and to 1.8

62 ANNALS OF THE MISSOURI BOTANICAL GARDLN [Vol. 67

mm wide, glabrous or minutely pubescent; stipules briefly connate, oblong, to 7
mm long, to 5 mm wide at the base, retuse at the apex or split almost to the base,
glabrous. Inflorescences axillary or terminal with short twiglets subtending, ses-
sile, the flowers conglomerate in a rotund mass, 1.0-1.5 cm in diam.; bracts
purplish, 4-6, ovate, shorter than the flowers, obtuse. Flowers with the hypan-
thium short, the calycine cup ca. 0,3 mm long, the teeth 5, unequal, obtuse, to
0.6 mm long, petaloid, glabrous except marginally, with a few oblong glands on
the inside of the cup, to 0.2 mm long; corolla white, the tube cylindrical, some-
what inflated, ca. 5 mm long, to 1,8 mm wide, petaloid, glabrous outside, villose
within on the upper %, the lobes 5, oblong, glabrous; stamens 5, the anthers
oblong, ca. 2.2 mm long, 0.5 mm wide, obtuse, the filaments ca. 0.15 mm wide;
style ca, 2.2 mm long, shorter than the tube, the stigmas upright, ca. 0.3 mm
long. Fruits sessile, oblong rotund, 3.5-4.5 mm in diam., each pyrene compressed
hemispherical in cross section, smooth, glabrate, usually with minute white ra-
phides, the grooves about 4.

Cephaelis axillaris is widespread in the tropics, extending from Mexico,
through Central America, to Colombia and Venezuela. It also occurs in the West

Indies.

Steyermark (1972), who has incorporated Cephaelis into Psychotria, anno-
tated many specimens of '"Cephaelis axillaris Sw.'' as Psychotria aubletiana
Steyermark. In the synonomy under P, aubletiana he (1974) listed: ""Cephaelis
axillaris Sw., Prodr. Veg. Ind. Occ. 45: 1788, no Psychotria axillaris Willd., Sp.
PI. 1: 962. 1798, basada sobre Ronabea erecta Aubl., ni Psychotria axillaris
Veil., Fl. Flum. 67: 2. pL 32. 1827." He recognized 7 varieties, 5 of which occur

in Venezuela.

CHiRiQUi': New Switzerland, valley of Ri'o Chiriqui Viejo, Allen 1391 (F, MO). Cerro Horqueta,
Blum & Dnyer 2654 (MO). Las Nubes, 5.5 km NW of Rio Chiriqui Viejo, Busey 674 (MO). Las
Cumbres, near Cerro Punta, Croat 13712 (MO). Above Rio San Felix, 13 mi N of Rio San Felix,
800-1200 m, Croat 33424 (MO). Cerro Piinta, Davidson 211 (F). Cerro Horqueta, Dwycr et al. 547
(DUKE, F, GH, K, MO, NY, PA, UC, US). Road to Bajo Grande. Cerro Punta, Folsom 4013 (MO).
Cerro Colorado, 1500 m, Folsom iSc Collins 1767 (MO). Between Quebrada Hondo and divide on
Caldera Chiriquicito Trail, Kirkbridc & Duke 921 (MO). Divide, Chiriquicito Trail, Kirkhride & Duke
963 (MO). Cerro Pando, 16 km NW of El Hato del Volcan, 2000-2482 m, Mori & Bolten 7266 (MO).
Cerro Hornitos, 40 km NW of Gualaca, Mori d Bolten 7474 (MO). N of San Felix at Chiriqui'-Bocas
del Toro border, Cerro Colorado Copper Mine Road, 5000-5500 ft, Mori & Kallunki 5792 (MO).
Cerro de Garagara, Pittier 5643 (US). 6 mi above Cerro Punta, Boquete Trail, ca. 7000 ft, Tyson 7071
(MO). Cerro Punta, White 28 (F, MO). Upper Chiriqui Viejo, White & White 25 (F, MO). Cerro
Horqueta, Wilbur et al. 13464 (MO). Bajo Chorro, Woods<m & Sehery 699 (F, MO). Casita Alta,
Volcan de Chiriqui, Woodson et al. 853 (F, MO), darien: Serrania de Pirre, 9-10 km N of Alto de
Nique, 1520-1560 m, Croat 37856 (MO). Cerro Mali base camp, lower montane wet forest, 1400 m,
Gentry S: Mori 13824 (MO).

2. Cephaelis bristanii Dwyer & Hayden, Ann. Missouri Bot. Gard. 55: 34. 1968.
type: Panama, Bristan 575 (MO).

Suhshnihs to 30 cm tall, the stems smooth, glabrous, black, scarcely nodose,
often prostrate and bearing long narrow roots. Leaves oblong, 9-16 cm long, 3.5-

6.0 cm wide, apically deltoid, ultimately obtuse, attenuate cuneate at base, the
costa slender, prominulous or plano-convex beneath, the lateral veins 15-30, 0.5-
0.8 cm apart, arcuate, the intermediate veins evanescent, rigidly papyraceous,
concolorous, glabrous; petioles to 4 cm long, to 1.5 mm wide; stipules connate,

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 63

the sheath compressed annular, 1-2 mm long, minutely and obtusely 2-apiculate.
Inflorescences axillary, the peduncles solitary, to 2 cm long, black, smooth, the
flowers aggregated into 1-2 rotund capitula, ca, 1 cm in diam,, the bracts many,
clustered, the interior bracts becoming narrower adaxially, oblong, acute, pu-
berulent. Flowers with the hypanthium funnel shaped, scarcely 2 mm long, the
calycine cup petaloid in texture, ca. 1 mm long, the lobes 4(?), short, truncate or
obtuse, often much reduced, the margin then erosulose, often with a few straight
cilia; corolla with the tube narrow cylindrical, to 7 mm long, ca. 1 mm wide in
the middle, petaloid, glabrous outside, densely white pilose within above the
middle of the tube, the lobes 4, to 2 mm long, obtuse; stamens 5, the anthers ca.

w

0.8 mm long, the filaments linear, ca. 3 mm long; stigmas 2, erect, ca. 1 mm long.
Fruits red (fide Bristan), not seen.

Cephaelis hristanii is known from Costa Rica and Panama. ''Hinupichica''
(Cuna, fide Duke 1243),

BOCAS DEL TORo: Quebrada Lugron and Cerro Bonyik near Ri'o Teribe, Kirkhhdc & Duke 641
(MO), darien: Cerro Pirre, Bristan 575 (MO). Cana Cuasi' Trail, Cerro Campanimento near Tres
Bocas, Rio Cuasi headwaters, Kirkhrkle & Duke 1243 (MO).

3. Cephaelis camponutans Dwyer & Hayden, Ann. Missouri Bot. Garden 55: 35.
1968. TYPE Panama, Dwyer & Gauger 7346 (MO, holotype).

Subshrubs to 0.7 m tall, the branchlets smooth, glabrous, the nodes somewhat
prominent; roots slender. Leaves strongly ascending, to 18 cm long, to 5 cm
wide, attenuate acuminate at the apex, the acumen to 3 cm long, acute at the
base, scarcely inequilateral, the costa scarcely prominulous to plane beneath, ca.
2 mm wide proximally, the lateral veins 15-20, evanescent when dry, strongly
ascending, the margin somewhat thickened, subcoriaceous, smooth, glabrous;
petioles slender, to 3 cm long; stipules obovate rotund, to L5 cm long, ca. 1 cm
wide, obtuse or somewhat bifid(?) at the apex, thin, with minute raphides, the
margins erosulose. Inflorescences axillary, 1-3 per axil, the peduncles to 1 cm
long, often nodding, the flowers conglomerate in compressed-rotund heads to 2
cm wide, blackening on drying, the exterior bracts solitary, subovate, to 1.5 cm
long, coriaceous, with a median keel, the internal bracts smaller and narrower,
a pair of bracteoles enclosing each flower. Flowers with the calyx tubular, the
lobes 5-6, erect, subulate, longer than the tube, the margin scarious; corolla
purple or white; stigmas 2, erect, digitiform, ca. 1 mm long. Fruits (immature)
crowned by a persistent calyx, the pyrenes 2, elliptic; seeds with the adaxial
surface plane, the endosperm smooth, the testa somewhat pigmented, perhaps
bilamellate, the parenchyma cells somewhat isodiametric, 23-55 m^ in diam.

Cephaelis camponutans Dwyer & Hayden is known only from Panama.

BOCAS DEL TORo: Punta Pena near Chiriquicito, Lewis et al. 2171 (GH, MO), colon: Santa Rita
Ridge, Croat 15345 (MO); Dwyer & Lallatliin 9042, 9043 (both MO); Mori & Kallunki 2090 (MO).
PANAMA: 1 mi past La Eneida, Cerro Jefe, Correa & Dressier 970 (MO). Goofy Lake to Cerro Jefe,
Croat 15204 (MO). El Llano-Cartf Road, 10 mi from Intcramerican Highway, Croat 33766 (MO).
Cerro Jefe, ca. 600 m, Croat 35H7! (MO). El Llano-Carti Road, 12 km from Panamerican Highway,
D'Arcv 10609 (MO). Cerro Jefe, Dwyer A Gaui^er 7346 (MO); Cjentry 2882, 3523 (both MO). El
Llano-Carti Road, Folsom 2589 (MO); Liesner 677, 1190 (both MO): Mori et al. 4630 (MO), vera-
GUAS: 5 mi NW of Santa Fe, Liesner 974 (MO).

64 ANNALS OF THR MISSOURI BOTANICAL GARDHN [Vol. 67

4. Cephaelis correae Dwyer & Hayden, Ann. Missouri Bot. Garden 55: 36. 1968.
type; Panama, Lewis et at, 1759 (MO, holotype).

Small trees, to 5 m tall, the branchlets terete, smooth. Leaves oblong, to 29
cm long, to 12 cm wide, obtuse to deltoid at the apex, cuneate at the base, the
costa somewhat prominulous above, ca. 2.5 mm wide, prominulous beneath, the
lateral veins ca. 20, 1.0-1.5 cm apart, at first strict, becoming arcuate near the
margin, the intermediate veins pinnatiform, membranaceous, smooth, glabrous;
petioles 0.5-0.8 cm long, slender, ca. 0.25 cm wide, smooth, glabrate; stipules
briefly connate, the sheath short, 2-5 mm long, interpetiolar, truncate, the lobes
large, subrotund, longer than the sheath, obtuse, somewhat spreading, the margin
scarious. Inflorescences terminal, solitary, glabrous, the peduncle to 20 cm long,
at maturity nodding, ca. 0.3 cm wide in the middle, the exterior bracts 2, to 5 cm
long, to 5.5 cm wide, conspicuously glistening, red or somewhat purple, venose,
the interior bracts to 1.5 cm long, red to purplish, the larger exterior bracts

compressed obovate rotund, much wider than long, undulate marginate, often
with several shallow lobes, the smaller bracts obovate rotund and clawed, the
claw short or to about the same length as the blade, glabrous, petaloid. Flowers
ca. 1.5 cm long, the hypanthium and the calyx 2-3 cm long, glabrous, the cup
eglandular, the calycine teeth 5, triangular, unequal, 1-1.2 mm long, each tooth
often erosulose; corolla white, the tube cylindrical, to 10 mm long, inflated above,
stiffly petaloid, glabrous outside, puberulent within near the filament attachment,
the lobes 5, triangular oblong, ca. 3 mm long, glabrous; stamens 5, the anthers
linear, ca. 3 mm long, the filaments attached near the middle of the tube; style
linear, to 12 mm long, the stigmas 2, oblong, ca. 1.2 mm long. Fruits glistening
blue, obovate oblong, 1.5 cm long, the calycine cup persistent; seeds subtrian-
gular, flat on the ventral face, somewhat sulcate in the middle.

Cephaelis correae is found only in Panama.

con h: El Vallc, Allen 1648 (MO). Cerro Pilon and La Mesa, Corren 52 (MO); Croat 14335
(MO); Duke 12347 (MO); Duke & Lalhithin 15010 (MO); Dnver 10534 (MO); D\v\er & Correa 7961
(MO); Dwyer & Lalhithin 8690 A (MO); Gentry 5695 (MO); Lewis et al. 1759 (GH, MO, UC, US);
Luteyn S: Kennedy 1707 (MO); Porter et al. 4636 (COL, MO, SCZ, UC, YEN); Wilbur ct Lutextt
11722 (MO). La Pinlada, Hunter & Allen 541 (MO).

5. Cephaelis croatii Dwyer. ^^ type: Panama, Croat 27204 (MO, holotype; NY,
isotype).

Shrubs 1.0-1.3 m tall, the branchlets glabrous, smooth, ultimately angular,
the pith porate. Leaves elliptic oblong, 14.5-29.0 cm long, 5.5-9.0 cm wide,
rounded or obtuse and abruptly acuminate at the apex, basally cuneate or atten-

^^ Cephaelis croatii Dwyer, spec. nov. Frutices ad L3 m aiti, ramulis glabris. Folia elliptico-
oblonga. 14.5-29.0 cm longa, 5.5-9.0 cm lata, basin versus cuneata. venis lateralibus ca. 30, lato-
arcuatis, rigido-papyracea minuto-puberula; petiolis 3-5 cm longis; stipulis basi connatis ovato-ob-

longa vel oblongo-rotunda, ad 1.5 cm longa, versus apicem angusta tunc in lobos 2 subulatos vel
angusto-oblongos fissa. Inftorescentiae axillares, 4-6 cm longae, 3.0-5.5 cm latae, capitibus florum
6-8 oblongis vel rotundis, ad 1.5 cm latis, bracteis exterioribus ad 0.8 cm longis. Flores cupula calycis
cyiindrica, 0.5 mm longa, lobis 5, 0.6-1.0 mm longis; corolla alba tubo cylindrico, 6.5 mm longo,
extus glabro intus villoso lobis 5 lanceolatis, ca. 2.5 mm longis; anthcris 1 mm longis. Fruetus non
visi.

1980] DWYHR— FI.ORA OF PANAMA (Family 179. Rubiaccac) 65

uate acute, somewhat inequilateral, the costa prominulous above, prominent be-
neath, the lateral veins ca. 30, widely arcuate with an irregular vein between a
pair of lateral veins, rigidly papyraceous, slightly discolorous, minutely puberu-
lent; petioles 3-5 cm long, often twisted, to 0.25 cm wide, glabrate; stipules
connate, often turgid, ovate oblong or oblong-rotund, to 1.5 cm long, narrow
toward the apex, then each part splitting into 2 subulate or narrowly oblong
ascending lobes, these to 0.7 cm long, subcoriaceous, glabrous. Inflorescences
axillary, 5-6 cm long, 3.0-5.5 cm wide, drying black, the peduncle to 3 cm long,
to 0.3 cm wide, twisted, piano compressed, the flowers aggregated into heads,
these 6-8, oblong or rotund, to 1.5 cm wide, sessile or on branchlets, variable in
length, to 2 cm long, puberulent; bracts rotund, outermost 8-10 mm long, 8 mm
wide, concave, carnose, minutely pubescent outside, minutely ciliolate within,
purple. Flowers with the hypanthium short, the calycine cup cylindrical, ca. 0.5
mm long, turgid, petaloid, with a gland at the base of adjacent lobes, the lobes
5, triangular subulate or narrowly oblong, 0.6-1.0 mm long, obtuse, glabrous
within, ciliate; corolla white, the tube cylindrical, ca. 6.5 mm long, somewhat
dilated below the middle, glabrous outside, villose within, the lobes 5, lanceolate,
ca. 2.3 mm long, somewhat cucullate; stamens 5, the anthers oblong, 1 mm long,
obtuse, the filaments short; style slender, somewhat expanded toward the apex,
the stigmas 2, short, rotund, wider than the style. Fruits not seen.

Cephaelis croatii is known only from Panama. It is named in honor of Thomas
B. Croat, distinguished student of the flora of Panama. There are several distin-
guishing characters: relatively broad leaves with numerous lateral veins; large
bilobed stipules; and subrotund, capitate clusters of flowers, about 5 per head.

DARitN: Upper gold mining camp of Tyler Kittredge, headwaters of Ri'o Tuquesa ca. 2 air km
from continental divide, Croat 27204 (MO).

6. Cephaelis dichroa (Standley) Standley, Publ. Field Columbian Mus., Bot. Ser.
4: 296. 1929.

Evca dkhroa Standley, Contr. U.S. Natl. Herb. 18; 124. 1916. type: Panama, P'tttier 3218 (US,
holotype).

Glabrous shrubs, to 2 m tall. Leaves elliptic, 8.5-13.0 cm long, to ca. 3.2 cm
wide, acuminate at the apex, cuneate at the base, the lateral veins ca. 12, stiffly
papyraceous, concolorous, often drying yellowish, glabrous; petioles to 3 cm
long, wiry, often flexuous; stipules connate, ca. 3.5 mm long, the body com-
pressed cylindrical, each part with 2 widely spaced, erect, obtuse lobes about as
long as the sheath. Inflorescences terminal, the peduncle 5-8(-9) cm long, ter-
minated by 3 heads of flowers, the trio subtended by 2 large obovate bracts, to
2.2 cm long, ca. 1.8 cm wide. Fruits subrotund to 0.8 cm wide.

Cephaelis dichroa is known only from Panama. Standley's original description
of the inflorescences and flowers may help in the understanding of this species:

'Teduncles terminal or rarely also axillary, 3-9 cm long; bracts 15-20 mm long,
rounded ovate, acutish, slightly united at the base, green or tinged with red or
yellow, subtending 3 pedunculate heads, the peduncles 3-25 mm long; each sec-
ondary head subtended by 2 orbicular-rhombic bracts 12-15 mm long, these yel-

66 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

low or tinged with purple, each head consisting of 3 flowers, each flower sub-
tended by an oblong obovate acute bract 10-12 mm long and an oblong-linear
bracteole somewhat shorter; flowers sessile; calyx 3 mm long, twice as long as
the ovary, somewhat oblique, yellow, deeply 5-lobed, the lobes ovate, oblong,
acute; corolla yellowish white, the tube 10 mm long and 2 mm in diameter, the
5 lobes 2.5-3 mm long, oblong, obtuse; stamens included, inserted in the middle
of the tube, the filaments about as long as the anthers, these linear, 2.5 mm long;

fruits not seen/'

Another collection, Mori & Kallunki 2732 (MO), from between Fort San Lo-
renzo and Fort Sherman near Pavon Road junction with Road 82 (S 2), Canal
Zone, may be C. dichroa. It is IVi m tall with bracts magenta and fruits black,
but its sea level locality argues against such determination.

CHiRiQuf: Cerro Horqueta, Hat^en ct Ha^cn 2089 (F, US). Cerro Colorado, 50 km N of San
Felix on the continental divide, 1200-1500 m, M<ni & Dressier 7843 (MO). Cuesta de Las Palmas,
S slope of Cerro Horqueta, 1700-2100 m, Phticr 3218 (F, NY), Cerro Horqueta, Pittier 3266 (US).

7. Cephaelis dimorphandrioides Dwyer.^- type: Panama, Croat 27470 (MO, ho-
lotype).

Shrubs to 1.4 m tall. Leaves elliptic oblong, often falcate oblong, 17-30 cm
long, 6-14 cm wide, deltoid or obtuse toward the apex, acute or abruptly acute
toward the base, the costa slender above, prominent beneath, the lateral veins
13-17, arcuate, often with one smaller and shorter vein between 2 lateral veins,
the intervenal areas spreading reticulate, rigidly chartaceous, discolorous, gla-
brate above, minutely puberulent beneath; petioles lignose, to 4.5 cm long, twist-
ed, puberulent; stipules (only one pair seen at tip of twig) oblong rotund, to 0.7
mm long, deeply 2-lobed, coriaceous, the lobes rounded, farinose, glabrous with-

in. Infl'

12 cm long, to 4 cm wide,

obtuse at the apex, black when dry, the flowers more than 100. Flowers perhaps
sessile, the hypanthium short, swollen, puberulent, the calycine cup short, ca. 5
mm long, the teeth subulate, to 6.5 mm long, unequal, acute, puberulent outside,
glabrous inside, carnose, with 3 longitudinal veins, with 2-4 oblong glands, to 0.3
mm long, at the sinuses; corolla red outside, the tube narrowly cylindrical, 10-14
mm long, puberulent outside, glabrous within, subcarnose, the lobes 5, yellow
within, lance oblong, to 3 mm long, carnose, puberulent outside; stamens 5, the
anthers narrow oblong, 2.8-3.0 mm long, minutely apiculate, slender, attached
below the middle of the tube; ovarian disc 4-lobed, to 0.8 mm long, the style
linear, 9-14 mm long, the stigmas 2, reflexed, linear oblong, to 0.8 mm long.

'-' Cephaelis diniorphandrloides Dwyer, spec. nov. Frutices ad 1.4 m alti. Folia elliptico-oblonga,
saepe falcato-oblonga, 17-30 cm lotiga, 6-14 cm lata, venis lateralibus 13-17 rigido-chartacea supra
glabrescentia infra minuto-puberula: peliolis ad 4.5 cm longis; stipulis oblongo-rotundis, ad 0.7 mm
longis, bilobatis glabris. Infloreseentiae terminaies sessiles cylindricae et spicae similes, 5-12 cm
longae, ad 4 cm latae, floribus numerosis fortasse 100. Flares cupula calycis ca. 5 mm longa, dentibus
subulatis ad 6.5 mm longis; corolla extus rubra, tubo angusto-cylindrico, 10-14 mm longo, extus
puberulo intus glabro lobis 5, ca. 3 mm longis; antheris ca. 3 mm longis. Fruetus oblongi, ad 0.5 cm
longi.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 67

Fruits (immature) oblong, to 5 mm long, to 0.3 mm wide, truncate, each pyrene
smooth, black, 10-ribbed, the hairs weak, white, arachnoid.

Cephaelis dimorphandrioides is known only from Veraguas, It is distinguished
from all other Cephaelis by the cylindrical spikelike inflorescence which bears
a resemblance to the inflorescence of the legume genus Dimorphandra.

veraguas: Valley of Rfo Dos Bocas between Alto Piedra and Caiovebora, 350-400 m, Croat
27470 (MO, holotype). Road from Santa Fe to Caiovebora (Panamanian Highway 35) 16 km from
Santa Fe; on Atlantic slope near Rio Caloveborita, Mori et aL 6675 (MO).

8. Cephaelis discolor Polak., Linnaea 41: 572. 1877. type: Costa Rica, Pola-
kowsky 384 (F, MO, US, photos).

Cephaelis tonduzii Krause, Bot. Jahrb. Syst. 54. Beibl. 119: 45. 1916. typf: Costa Rica, Tonduz

11461 (F).
Evea ^uapilensis Standley, J. Wash. Acad. Sci. 15: 104. 1925. type: Costa Rica, Standley 37025 (US,

holotype).
Cephaelis nicaraguensis Standley, Trop. Woods 16: 46. 1928. type: Nicaragua, En^^lesin^ 5H (F).
C. gnapilensis (Standley) Standley, Publ. Field (Columbian Mus., Bot. Ser. 4: 295. 1929.

Siihshruhs to 1 m tall, usually unbranched, the stem terete, smooth, glabrous.
Leaves oblong, to 22 cm long, to 8.5 cm wide, subobtuse to deltoid at the apex,
acuminate, the acumen to 1.5 cm long, basally cuneate or attenuate acute, slightly
inequilateral, the costa prominulous above, prominent beneath, the lateral veins
12-14, arcuate, each pair with usually 1 subparallel irregular vein, soon evanes-
cent, the intervenal areas spreading reticulate, delicately marginate, stiffly char-
taceous, usually discolorous, glabrous; petioles to 7 cm long, slender, 0.13-0.20
cm wide, ascending, glabrous; stipules connate for 0.4-4.0 mm, each with 2
slender awns, to 8 mm long, these often reflexed, often deciduous, glabrous.
Inflorescences terminal, the peduncles 0.7-2.0 cm long, lignose, glabrate, the
flowers aggregated into a subrotund mass, 3-5 cm in diam.; bracts purple green
or purple, obovate lanceolate, 0.8-1 .0 cm long, petaloid, glabrous, venose. Flow-
ers with the hypanthium subrotund or oblong, ca. 1.5 mm long, constricted below
the calycine cup, the cup short, the lobes 4-5, subequal or equal, triangular, 0.8-
1.5 mm long, 2-3 times the length of the cup; corolla and sex organs not seen;
ovarian disc bilobed, ca. 0.5 mm long. Fruits bright blue, ca. 1 cm in diam., the
fleshy exterior often persisting, the pyrenes when united rotund, buttockslike,
the central groove conspicuous, ca. 5 mm in diam., glabrate, smooth, the raphides
minute, numerous, the calycine cup persistent, 0.5-0.8 mm in diam.

Cephaelis discolor occurs in Costa Rica, Panama and Colombia.

BOCAS DLL TORo: Water Valley, Wedel 1508 (MO), canal zone: Barro Colorado Island, Croat
11870 (MO). Pipeline Road, Gentry S: Dressier 1978 (MO), colon: Santa Rita Ridge Road, 8.5 km
from Boyd-Thatcher Highway, Mori & Kallunki 2089 (MO), darien: La Boca de Pirre, Bristan 1273
(MO). Cerro Pirre, Duke 5307 (U0)\ Duke & Elias 13767 (MO). Cocolito, Dwyer5l36 (MO). Panama:
Cerro Campana, Allen 1886 (MO). Cerro Trinidad, Allen 3774 (MO). Cerro Campana, Croat 14752
(MO). Cerro Jefe, Foster & Kennedy 1818 (F). Cerro Campana, Gentry 5783 (MO); Kirkbride &
Hayden 239 (MO); Lewis et al. 3061 (MO), veraguas: N of Santa Fe, N of Escuela Agricola Altos
de Piedras, Mori A Kallunki 2533, 2587, 2619 (all MO). Colombia: choco: Serrania del Darien, W
of Unguia on Panama border, 1 150 m, Gentry et al. 17038 (MO).

68 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol.67

9. Cephaelis dressleri Dwyer.''' type: Panama, Croat 16801 (MO, holotype).—

Fig. 15.

Subshruhs to 1.5 m tall, the stem lignose, rough, glabrous, the pith partly
porous, sometimes rooting toward the apex. Leaves occasionally sessile, obovate
oblong, 10-44 cm long, 6.5-22.0 cm wide, rounded or widely deltoid at the apex,
short acuminate, the acumen triangular, to 1 cm long, to 1.5 cm wide at the base,
cuneate toward the base finally slightly auriculate, the costa prominulous above
or prominent, to 0.2 cm wide, prominent beneath, to 0.35 cm wide, dark brown
and densely minutely puberulent, the lateral veins 21-25, slightly arcuate, forming
an undulate marginal vein to 1 cm from the margin, the venules irregular pinnati-
form and spreading, rigidly papyraceous to subcoriaceous, 1 of the 2 leaves drying
slightly dark purple red beneath, minutely ferrugineous puberulent on the veins
and venules; petioles, if present, to 4 cm long, stout; stipules (only 1 seen at the
tip of the stem) ovate lanceolate, to 2.8 cm long, to 1 cm wide, subcoriaceous,
glabrous, the veins somewhat prominulous, fanshaped, the awns narrowly ovate
subulate, to 0.9 cm long, acute. Inflorescences terminal, 4.5-8.0 cm long, the
peduncles 1-4 cm long, 0.3-0.5 cm wide, glabrous, bearing several short branch-
es, the flowers disposed in a few subsessile oblong or subrotund heads, ca. 2 cm
in diam., the cluster of heads compressed rotund, 3.5-5.5 cm wide; outermost
bracts rotund or spathulate, to 12 mm long, brown, ultimately acute, concave,
within with an irregular series of glands on each side of a median vein or with a
few diffuse glands, ciliate; interior bracts several, narrower and more spathulate.
Flowers subsessile; hypanthium scarcely measurable, the calycine cup to 2 mm
long, petaloid, with ca. 10 unequal, narrowly oblong or widely subulate glands
attached toward the base of the cup, the calycine lobes 5, subulate or triangular,
to 3 mm long, slightly unequal, acute, densely ciliate; corolla white, the tube
narrow cylindrical, 10-14 mm long, petaloid, glabrous or minutely puberulent
outside, glabrous inside except white villose near or above the middle, the lobes
5, narrowly lanceolate, to 4.5 mm long, 1 mm wide, conspicuously cucullate near
the middle, ultimately obtuse, carnose; stamens 5, the anthers narrowly ovate
oblong or oblong, 2.5-3.0 mm long, sagittate at the base, the filaments slender,
ca. 2.5 mm long, attached ca. 2 mm below the mouth; ovarian disc oblong rotund,
ca. 1 mm long, the style slender, 6.5-9.0 mm long, the stigmas slender, ca. 2 mm
long, densely villosulose. Fruits sessile, elliptic oblong, to 1.2 cm long, 0.4 cm
wide, smooth, glabrous, each pyrene 4-sulcate, the ribs obtuse angular, the often
persistent calycine cup and teeth to 3.5 mm long.

Cephaelis dressleri is known from Panama and Colombia. It is named in honor
of Dr. Robert Dressier, student of the flora of Panama and a specialist in the

" Ci'pluiclis dressleri Dwyer, spec. nov. Suffrutires ad L5 m alti, caule lignoso aspero glabro
interdum apice rhizophoro. Folia sessilia vel subsessilia obovato-oblonga, 10-44 cm longa, 6.5-22.0
cm lata, rotunda apice vel vix latodeltoidea, versus basim cuneata ultime vix auriculata venis late-
ralibus ad 25, rigido-papyracea vel subcoriacea puberula; stipulis ovato-lanceolatis, ad 2.8 cm longis,
ad 1 cm latis, setis solitariis ad 0.9 cm longis, acutis. Inflorescentiae terminales, 4.5-8.0 cm longae,
peduncuio 1-4 cm longo, caespite florum compressorotundo, 3.5-5.5 cm lato: bracteis exterioribus
rotundis vel spathulatis, ad 12 mm longis. Flares cupula calycis dentibus ad 3 mm longis; corolla tubo
10-14 mm longo, extus glabro, lobis ad 4.5 mm longis; antheris 2.5-3.0 mm longis. Fructus elliptico-
oblongi, ad 1.2 cm longi, glabri.

19801

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

69

Figure 15. Cephaelis dressleri Dwyer. — A. Leaf and inflorescence (xVi). — B. Flower (x3).
C. Fruit (x3). [After Dressier 4677.]

70 ANNALS OF THE MISSOURI BOTANICAL GARDLN [Vol. 67

Orchidaceae. It is distinguished by its large sessile or subsessile leaves, round at
the apex, with many lateral veins, and subauriculate at the base. The glands
within the larger bracts and within the calycine cup are diagnostic.

colon: Near Agua Clara at rain gauge, Santa Rita lumber road. Dressier 4677 (MO). darii':n:
Trail from Tacarcuna village, Rio Tacarcuna to Cerro Mali, 800-1300 m. Gentry 13598 (MO), san

BLAs: Puerto Obaldia, Croat 16801 (MO). Colombia: choco: Ri'oTigre W of Unguia, base of Serrania
del Darien, 300 m, A. Gentry & L. Agiiirre 15191 (MO).

10. Cephaelis elata Swartz, Prodr. 45. 1788. type: Jamaica, Swartz. not seen

Cephaelis punicea Vahl, Eclog. Amer. 1: 19. 1796. type: Jamaica, von Rohr, not seen.
Tapogomea elata Aubl. ex Lam., Encycl, Melh. 7: 585. 1806. type: not *
Cephaelis costarlcensis Schlecht., Linnaea 28: 546. 1856. type: not seen.

Uragoga punicea (Vahl) Schum. in Engl. & Prantl, Nat. Pflanzenfam. IV: 120. 1891.
Evea elata (Sw.) Standley, Contr. U.S. Natl. Herb. 18: 123. 1916.

Cephaelis rhiricpiiensis Standley, Ann. Missouri Bot. Card. 28: 469. 1941. type: Panama, Woodson
iSi Schery 610 (MO, holotype).

Shrubs or trees to 8 m tall, the branches few to many, smooth, terete or
angular, glabrate, occasionally constricted below the nodes, the nodes often swol-
len (persistent stipules). Leaves narrowly oblong or obovate oblong, 10-25 cm
long, 1.5-7.0 cm wide, obtuse, widely deltoid or rounded at the apex, acuminate,
the acumen short, acute, basally cuneate to obtuse, the costa prominent, prom-
inulous to subplane above, prominent beneath, the lateral veins (12-) 16-22,
widely arcuate or at first widely divergent and then arcuate marginally, the margin
often slightly revolute, stiffly papyraceous to subcoriaceous, not herbaceous
(Panama), lustrous and glabrous above, glabrous or pubescent on the costa and
the veins beneath; stipules persistent, connate, turgid, cupshaped, to 1.2 cm long,
each part of the sheath bilobed, the lobes usually hemispherical, as long as or
longer than the sheath, obtuse, coriaceous, glabrous, the margins scarious, often
widely so. Inflorescences terminal, the peduncle solitary or rarely 2-4 at the tip

of the twiglet, 0.5-3.5 cm long; heads including 2 outermost bracts to 6.5 cm
wide; outermost bracts ovate rotund, to 7 mm long, deep red or rusty orange.

CANAL zonh: Camp Pina, Duke 4400 (MO). Pena Road, Tyson ct Chu 1700 (MO). Fort Sherman,
Tyson & Dwyer 1215 (MO), chiriqui': Bajo Chorro, Boquete District, Davidson 103 (MO). Cerro
Horqueta, Kirkhride 131 , 132 (both MO). Between Pinola and divide on Chiriquicito-Caldera Trail,
Kirkhride & Duke 845 (MO). Bajo Chorro, Woodson & Schery 610 (MO). cocLt: Cerro Pajila, N of

EI Valle de Anton, Allen & Allen 4168 (MO). Cerro Caracoral, Duke & Dwyer 15103 (MO). El Valle,
[:>\vyer & Correa 7998 (GH, MO, US); Lewis et al. 1713 (GH, MO. US), colon: Santa Rita Ridge,
Croat 14/46 (MO), dariln: Cerro Pirre, Bristan 462 (MO); Duke 6570 (MO). Summit camp between
Sasardi and Mortf, Duke 10034 (MO). Cerro Pirre, Duke & Ellas 13752 (GH, MO). Top of Cerro
Mah', old helipad, Gentry & Mori 1366 (MO). Summit camp, Darien-San Bias border, Oliver et al.
3692 {COL. MO, UC, VEN). Cana-Cuasi Trail, Terry & Terry 1480 {MO). Panama: Cerro Jefe, Blum
& Duke 2197 (MO). Cerro Campana, Blum et al. 2371 (MO). El Llano-Carti Road, 330 m, Croat
33742 (MO). Cerro Jefe, Duke 8018 (MO); Duver 444 (MO). Cerro Azul, Dwxer 1369 (MO). Cerro
Jefe, Elias & Ilayden 1797 (MO); Folsom 2012 (MO). El Llano-Carti road, 6 km N of Panamerican
Highway, Folsom 2551 (MO). El Llano-Cartf road, 13.7 km N of Panamerican Highway, Eolsom
2579. 3505 (both MO). Top of Cerro Jefe, Folsom 3614 (MO). Cerro Campana. Harden 118 (MO).
Cerro Jefe, Kirkhride J- Crehhs 33 (MO). Altos del Rio Pacora, Lewis et al. 2262 (COL, DUKE, K,
MO), 2356 (COL, MO, UC, VEN). Cerro Jefe, Tyson 3189, 3205, 3231, 3254, 3285, 3288, 3373, 3389,
3425, 3546, 3577, 3588 (all MO); Witherspoon et al. 8282, 8503 (both MO), san blas: SE of Puerto
Obaldia, Croat 16750 (MO). E of Cangandi-Mandinga airport road, 2-5 mi S of Mandinga airport,
Duke 14771 (MO). Between Rfo Diabolo and Ri'o Acuati near Nargana, Duke 14865 (MO), vlraguas:
3.9-5.0 mi N of Santa Fe, Gentry 2951 (MO). Mouth of Rfo Concepcion, Lewis et al, 2806 (COL,

19801 DWYHR— FLORA OF PANAMA (Family 179. Ruhiaceae) 71

DUKE, MO, UC, VEN). Cerro Hornitos, 2238 m, Mori & BoUcn 7496 (MO). 10 km NW of Santa
Fe, road to Calovebora, Mori & Bolten 7639 (MO). N of Santa Fe, Escuela Agricola Alto de Piedra,
Mori & Kallunki 2507 (MO).

11. Cephalis gaugeri Dwyer.^* type: Panama, Mori et aL 3779 (MO, holotype).

Subshrubs to V3 m tall, the stem rhizomatous at the base, the roots fibrous,
the stem unbranched, ca. 0.3 cm wide, glabrous but with minute black puncta-
tions, the nodes well spaced. Leaves succulent, oblong, 5.0-7.5 cm long, 1.8-3.0
cm wide, widely deltoid toward the apex, acuminate, the acumen to 0.3 cm long,
ultimately obtuse, toward the base cuneate, obtuse or rounded, slightly inequi-
lateral, the costa prominulous above, subplane beneath, red on drying, to 0.2 cm
wide, the lateral veins ca. 7, arcuate, somewhat inconspicuous, the fine veinlets
obscure, the margin lightly revolute, coriaceous, concolorous, smooth, glabrous;

petioles to 1 cm long, rigid, glabrous; stipules free, often persistent, compressed-
rotund, to 1 cm long, to 1.3 cm wide, with 2 triangular or oblong lobes, ca. 0.5
cm long, the lobes erose or with subulate teeth, the body of the stipule rigidly
scarious, glabrous, the raphides dense. Inflorescences axillary, the peduncle ca.
1 cm long, ca. 0.3 cm wide, the fruits disposed in a globose head, to 2.5 cm in
diam. Fruits sessile, oblong, to 1.5 cm long, obtuse or truncate toward the apex,
carnose, smooth, glabrous, red, purple toward the apex, the bracts few, large,
imbricate, compressed rotund, to 0.8 cm long, perhaps much wider than long,

subcoriaceous, glabrous except ciliolate toward the margin, the margin erose, the
raphides dense.

Cepliaelis gaugeri is known only from Panama. It is named in honor of Mr.
George Gauger, a microbiologist in the Canal Zone who has assisted this author
in collecting plants in Panama.

The new species is distinguished by its stem being rhizomatous, its leaves
being succulent with few, inconspicuous lateral veins, and its stipules being large
and lobed.

Panama: Roads to Cerro Jefe and Altos de Pacora, Mori ct al. 3779 (MO).

12. Cephaelis glomerulata Donnell Smith, Bot. Gaz. 16: 12. 1891. type: Guate-
mala, ./. D. Smith 1637. not seen. — Fig. 16.

Psychotria glomerulata (Donnell Smith) Stcyermark, Mem. N.Y. Bot. Garden 23: 670. 1972.

Subshrubs to 2 m tall, the stem simple above or occasionally with 2 terminal

branches, the twiglets terete, glabrous, the nodes well spaced. Leaves lanceolate,
ovate lanceolate or oblong, occasionally inequilateral, occasionally salicoid, 5-

13 cm long, 1.6-5.5 cm wide, deltoid or acute at the apex, acuminate, the acumen

'^ Ci'phacUs gaugeri Dwyer. spec. nov. Sujfrutices ad V3 m aiti rhizomati caule simplici. Folia
succulentia, oblonga, 5.0-7.5 cm longa. 1 .8-3.0 cm lata, versus apicem lato-deltoidea venis lateralibus
ca. 7 in sicco evanescentibus, coriacea laevia glabra; petiolis ad 1 cm longis; stipulis compresso-
rotundis, ad I cm longis, ad 1.3 cm latis, prope medium lobis 2 ad 0.5 cm longis, in margine erosis
vel dentibus subulatis praeditis. Inftorescentiae axillares, pedunculo ca. 1 cm longo, ca. 0.3 cm lato,
fructibus in caespitem globosum ad 2.5 cm dispositis. Fructus sessiles oblongi, ad 1.5 cm longi,
carnosi, laeves, glabri, rubri.

72

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 67

Figure 16. Cephalis glomendala Donnell Smith. — A. Inflorescence (xVs). — B. Flower
(x2.4).— C. Fruit (x L2). [After Hammel 1819,]

to 1.5 cm long, ultimately acute or obtuse, cuneate or obtuse at the base, the
lateral veins 12-16(-20), widely arcuate, with several smaller veins between ad-
jacent laterals, these soon branching, the intervenal areas spreading reticulate,
chartaceous to papyraceous, glabrous, usually lustrous; petioles 0.3-1.5 cm long,
glabrous; stipules connate, erect, somewhat inflated, cylindrical, to 3 mm long,

truncate, glabrous. Inflorescences terminal, sessile or subsessile (the peduncle to
0.5 cm long), the bracts and flowers aggregated into an oblong or compressed

rotund head, 1.5-2.5 cm long, often somewhat wider than long, the outermost

bracts ovate rotund, to 1.5 cm long, to 1 cm wide, obtuse, rounded or bilobed at

the apex, subcoriaceous, glabrous, venose, eglandular within, white or green,

turning purple in fruit, with progressively smaller, narrower, and more concave

bracts within, each flower enclosed by 2 tightly enveloping bracteoles. Flowers

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceac) 73

sessile; hypanthium ca. 1 mm long, glabrous, the calycine cup about equal to the
hypanthium, delicately petaloid, glabrous, eglandular within, the teeth 5, irregular
but mostly subulate or triangular, to 1.2 mm long; corolla white, the tube cylin-
drical, ca. 1 cm long, petaloid, glabrous outside, puberulent within in the middle
of the tube for ca. 4 mm; stamens 5, the anthers subsessile, oblong, 2.2 mm long,
obtuse, minutely apiculate, the filaments attached near the middle of the tube;
style slender, about as long as the tube, the stigmas 2, dilated, oblong, 1.3 mm
long, subplane. Fruits fleshy, oblong or subrotund, to 1 cm long, china blue at
maturity, glabrous.

Cephaclis glomcridata extends from Guatemala to Panama. '^Uasika'' {Duke
14378).

BOCAS nr.L TORo: Punta Pena, near Chiriquicito, ca. 1000 ft, Lems et al. 2175 (COL, MO, UC,
VEN). Old Bank Island, Weclel 1930 (MO, US), 1941 (US). Fish Creek, Chiriqui Lagoon, Wedel2l99
(MO, US). Fish Creek Mountains, Wedcl2309 (MO, US). Bastimentos, Chiriqui Lagoon, Wedel 2907
(MO). Isia Colon, Wedel 2968 (MO, US); Woodson et al, 1936 (F, MO), canal zone: Telephone
cable trail between splice S16 and S49, Rio Indio, Steyermark & Allen 17428 (MO). Fort Sherman
near road to Gatiin along Road S-5, Tyson & Dn-yer 1215, 1243 (both MO), cocle: 16.7 km N of
turnoff to Coclesito from Llano Grande, Hammel 1819 (MO), colon: Santa Rita Ridge, Croat 14143
(MO); D\v\er8404, 8413 (both MO); Dnver et al. 8995 (MO). Highway to 8 mi E of Santa Rita Ridge,
800 ft, D\xyer et al, 8986 (MO). Santa Rita Ridge road. Gentry 1887 (MO). Santa Rita Ridge, Gentry
8822 (MO); Lewis et al, 5272 (MO). Santa Rita Ridge, 2.7 mi by gravel road NE of Transisthmian
Highway, Nee & Mori 3675 (MO). Panama: Road to Carti, 13 km N of El Llano, Bnsey 384 (MO).
Cerro Jefe, 5 mi S of Summit, Basey & Croat 284 (MO). El Llano-Cartf Road, 10 mi from Panamerican
Highway, 330 m, Croat 33756 (MO). Cerro Jefe, Croat 35869 (MO). Rio Sancantf, ca. 2 mi upstream
from Piria, ca. 120 m, Dake 14378 (MO). Goofy Lake, to 8 mi S of Goofy Lake toward Cerro Jefe,
Dwyer 7103 (MO). Between Cerro Jefe and La Eneida, 2100-2900 ft, Dnyer 8241 (MO). El Llano-
Carti Road, Folsom 2616 (MO). El Llano-Carti Road, 12 km N of Panamerican Highway, Folsom
3597 (MO). Cerro Jefe, ca. 1000 m, Gentry 6147 (MO). Cerro Jefe, ca. 3100 ft, Gentry et al. 3459
(MO). Cerro Jefe to La Eneida, Hayden 1016 (MO, UC), El Llano to Carti-Tupile, 16 km above
Panamerican Highway, Kennedy et al. 2429 (MO). Altos del Rio Pacora, 2500 ft, Lewis et al. 2262
(COL, DUKE, K, MO, UC, VEN). El Llano to Carti'-Tupile, 300-500 m, Liesner 1276 (MO). Gorgas
Memorial Labs, 5-10 km NE of Altos de Pacora, ca. 600 m, Mori & Kallunki 3325 (MO). Cerro Jefe,
2700 ft, Tyson 3373 (MO), veraguas: Mouth of Rio Concepcion, Hayden 2806 (COL, DUKE, MO,
UC, VEN). 3-4 km W of Santa Fe, 2500 ft, Nee 11308 (MO), san blas: Inland from airport at

Mandinga, Duke 8917 (MO).

13. Cephaelis insueta Dwyer.**^ type: Panama, Mori & Kallunki 3108 (MO, ho-
lotype).

Shrubs (?), the stems rhizophorous to the bases of the uppermost leaves,
square, dark red when dry, glabrous, the nodes crowded, the petiole scars sub-
rotund, to 0.8 cm long, the pith perhaps porate. Leaves sessile, oblong, 17-43
cm long, 8-16 cm wide, widely deltoid at the apex, acuminate, the acumen tri-
angular, ca. 1 cm long, the blade spathulate toward the base, finally subauriculate,
the costa subplane above or prominulous, prominent beneath, to 0.3 cm wide,
the lateral veins ca. 42, scarcely arcuate, the submarginal veins 2, these somewhat

^^ Cephaelis insueta Dwycr, spec. nov. suffrutices (?) caule rhizophoro etiam basibus foliorum
superiorium. Folia sessilia, oblonga, 17-43 cm longa, 8-16 cm lata, lamina basim versus spathulata
ultime subauriculata venis lateralibus ca. 42 vix arcuatis venis submarginalibus 2 margini subparal-
lelibus tenui-chartacea discoloria supra glabra subtus puberula. Inflorescentiae terminales ad 16 cm
longae, ad 6 cm latae, pedunculo recto, ad 10 cm longo; bracteolis minutis ovato-subulatis, ca. 1 mm
longis. Flores non visi. Fruetus elliptico-oblongi, ad 0.9 cm longi, ad 0.4 cm lati, laeves glabri opaco-
rubri calyce ca. 1 mm longo.

74 ANNALS OF THE MISSOURI BOTANICAL GARDRN [Vol. 67

parallel to the margin, the one nearer the margin inconspicuous, slender, 0.1 mm
from the margin, the other prominulous, 1 mm from the margin, the intervenal
areas spreading reticulate, thin chartaceous, discolorous, glabrous above, mi-
nutely puberulent beneath on the costa and the veins. Inflorescences terminal
(only one seen in fruit) to 16 cm long, to 0.6 cm wide, puberulent, the peduncle
erect, to 10 cm long, to 0.5 cm wide, flattened, the branches few, ascending, the
lowermost perhaps opposite, to 4 cm long; bracteoles minute, ovate subulate, ca.
1 mm long. Flowers not seen. Fruits on short pedicels; elliptic oblong, to 0.9 cm
long, to 0.4 cm wide, smooth, glabrous, dark red, the calyx persistent, ca. 1 mm
long, each pyrene 4-sulcate.

Ccphaelis insueta is known only from Panama. The Latin adjective means
''unusual'' which seems appropriate as the stem of the new species is rhizopho-
rous even to the apex; the basally spatulate leaves are striking. The 2 very reduced
submarginal veins which parallel the margin suggest the situation in e.g. Faramca
talamancariim Standley. The collectors of the new species note: ''detritus col-
lected in the leaf axils penetrated by adventitious roots from the stem."

VERAGUAS: 1 1 km from Escuela Agricola Alto de Piedra forest along Rio Dos Boca, tropical wet
forest, Atlantic slope, Mori & Kallunki 3108 (MO).

14. Cephaelis ipecacuanha (Brot.) A. Rich., Bull. Fac. Med. 4: 92. 1818.

Callicocca ipecacitcinha Brot., Trans. Linn. Soc. London 6: 137. 1802. type: Brazil, Gomez, not

seen.
Ipecacuanha officinalis Arruda, Diss. PI. Brazil 44. 1810. Not seen, reprinted in part as Appendix to

Koster, Travels in Brazil 2: 374-375. 1817. typh: Described from cultivation.
Evea ipecacuanha (Brot.) Standley. Contr. U.S. Natl. Herb. 18: 123. 1916.
Ouragoga ipecacuanha (Brot.) Farw., Drugg. Circ. 61: 175. 1917.

Suhshriihs to 0.25 m tall, the stems simple, arising from a rhizome, to 0.8 cm
in diam., the nodes often crowded toward the apex of the stem, and often rough
with stipular scars. Leaves oblong, obovate oblong, elliptic oblong, oblong sub-
rotund, 8-14 cm long, 3.2-9.0 cm wide, obtuse or rounded at the apex, occa-
sionally widely cuneate, acuminate, the acumen often vague, usually widely tri-
angular, to 1 cm long, basally obtuse, rounded, attenuate acute or cuneate (and

the lamina abruptly contracted), occasionally vaguely auriculate, the costa slen-
der, prominulous above and beneath, the lateral veins ca. 6, arcuate, uniting well

below the margin to form a submarginal undulate vein, with 1-2 smaller lateral
veins between a pair of lateral veins, stiffly papyraceous or chartaceous, glabrous
or occasionally sparsely puberulent above and beneath; petioles either absent or
to 0.5 cm long, glabrous; stipules persistent, appressed to the stem, thin, oblong
or ovate rotund, to 1 1 mm long, the body 3-5 mm long, with ca, 10 linear subulate
awns to 7 mm long, these spreading fanlike or curled, glabrous. Inflorescences
axillary and/or terminal, glabrous, the peduncles erect or deflexed, 1.2-4.0 cm
long, ca. 0.15 cm wide, terminated by a bracteate cluster of flowers, the heads
compressed rotund, 1.5 cm in diam., to 3.5 cm wide when bracts spread, the
bracts elliptic oblong to subrotund, 8-10 mm long, 3.5-8.0 mm wide, glabrous,
stiffly petaloid, with a few marginal, weak, subulate projections at the base, to
1.5 mm long, to 0.2 mm wide basally. Flowers sessile, distylous; hypanthium

19801 DWYKR^FLORA OF PANAMA (Family 179. Rnhiaccae) 75

0.1-0.2 cm long, glabrous, the calycine cup 0.1-0.2 mm long, eglandular within,
glabrous, the teeth 5, unequal, irregular, scarious, tending to be triangular, to 0.5
mm long; corolla tube cylindrical, to 4 mm long, petaloid, glabrous outside, pu-
berulent within from filament attachment to the mouth, the lobes 5, oblong or
ovate oblong, ca. 1.5 mm long, acute; stamens 5, the anthers exserted or included,
narrowly oblong, ca. 1.7 mm long, the filaments short or elongate, attached near
the middle of the tube; ovarian disc rotund, ca. 0.3 mm long, the style filamentous,
0.2-0.6 cm long, the stigmas minute. Fruits fleshy, red, oblong or rotund, to 1
cm long, the pyrenes smooth, glabrous or with a small ringlike scar.

Cephaclis ipecacuanha is well distributed throughout tropical America. The
roots and rhizome contain an alkaloid emetin, an emetic and expectorant. This
drug is known in English as ''ipecac'' or in Spanish as ''Raicilla.'' The United
States Pharmacopeia lists Rio or Brazilian ipecac as Cephaclis acuminata Kar-
sten, ''known in commerce as 'Cartagena, Nicaragua, or Panama ipecac' " which
is supposed to have different pharmacological properties from Rio or Brazil ipe-
cac. The name Cephaclis acuminata is an unpublished name which may refer to
plants which do not occur in Panama. Panamanian material seen to date is quite
uniform taxonomically and agrees with other material from South America which
has been going under the name Cephaclis (or Psychotria) ipecacuanha,

BARRO COLORADO ISLAND: Burtlctt & Lusscr 16723 (MO); Busex & Croat 237 (MO); Croat 4096,
4104, 4269, 6609, 12427, 151 17 (all MO); Harden 122, 127, 155, 1037 (all MO); Starry 55 (MO).
darien: Punta Alegre, Amhulo (MO). Ri'o Tuira and Rio Paca» Duke 5027 (MO). Line CC, Duke
5237 (MO). Ri'o Chucunaque, 2-10 mi above Cuna-Darien border, Duke 8566 (MO). Without other
locality, Sexton I46A, I46B (both MO), san blas: Headwaters of Rio Mulatupo, Elias 1770 (MO).

15. Cephaelis kennedyi Dwyer.^*' type: Panama, Kennedy 1779 (MO, holotype).

Suhshruhs to 2 m tall, the branchlets terete, smooth, glabrous, the nodes well
spaced. Leaves narrowly oblong or obovate oblong, 4-9 cm long, 0.8-3.0 cm
wide, obtuse toward the apex, often abruptly acuminate, the acumen to 1 cm
long, often falcate, ultimately obtuse or acute, basally acute or cuneate, slightly
inequilateral, the costa prominulous above, prominulous beneath, ca. 0.35 mm
wide, the lateral veins ca. 20, widely arcuate, subprominent, chartaceous, gla-
brous; petioles to 1 cm long, 0.8 mm wide, glabrous; stipules connate, the sheath
cupshaped, to 2 mm long, truncate, with narrowly subulate bodies, to 1 mm long,
yellow when dry, with 2 terminal subulate bristles arising immediately below the
margin of the sheath, these bifid above the middle, often longer than the sheath.
Inflorescences terminal, sessile or subsessile, capitate, oblong, to 0.6 cm long,

"* Cephaclis kennedyi Dvvyer, spec. nov. Snffrutices ad 2 m alti. glabri. Folia angusto-oblonga, 4-

9 cm longa, 0.8-3.0 cm lata, abrupto-acuminata acumine ad 1 cm longo, venls lateralibus ca. 20,
chartacea glabra; petiolis ad 1 cm longis; stipulis connatis vagina cupuliformi, ad 2 mm longa, truncata
setis 2 subulatis statim infra marginem vaginae affixis saepe quam vagina longioribus eis supra medium
bifidis. Inflorescentiae sessiles vel subsessiles terminales capitato-oblongae, ad 0.6 cm longae, ad 0.5
cm latae, bracteis exterioribus ad 6 mm longis, ca. 4 mm latis. Flores cupula calycis ca. 0.5 mm
longa, truncata dentibus nullis vel 5 punctis minutis praedita vel dentibus 2 longitudine inaequalibus,

ad 0.3 mm vel dente uno; corolla tubo ad 3.2 mm longo; anlheris oblongis, ca, 1 mm longis, Fructus
non visi.

76 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vol. 67

to 0.5 cm wide, outer bracts 2, oblong, 5.5-6.0 mm long, ca. 4 mm wide, obtuse,
subcoriaceous, glabrous, subplane, with numerous fimbriate glands within at the
base; interior bracts 2, somewhat stiff, concave, with a few basal bristles; flowers
8-10, each flower enclosed in 3 bracteoles. Flowers with the hypanthium to 0.5
mm long, the calycine cup 0.5 mm long, somewhat swollen, stiffly petaloid, gla-
brous, rigidly petaloid, the teeth variable, sometimes absent, sometimes disposed
as 5 minute points, sometimes with 2 unequal teeth, to 0.3 mm long, sometimes
with one tooth and sometimes the margin with one oblong petaloid obtuse lobe
to 2.5 mm long, and to 1.5 mm wide; corolla with the tube cylindrical and inflated,
to 3.2 mm long, rigidly petaloid, glabrous without, barbate within near the fila-
ment attachment region, the lobes 5, the anthers oblong, ca. 1 mm long, the
filaments linear, attached above the middle of the tube; style to 0.1 mm long, the
stigmas 2, 0.2 mm long, the ovarian disc compressed rotund, 0.6 mm long, con-
spicuously 2-lobed. Fruits not seen.

Cephaelis kennedyi is named in honor of Dr. Helen Kennedy, a student of the
flora of Panama. The new species is known only from Panama. It is clearly
distinguished from its closest relative, C. mucosa (Jacq.) Sw., by its much re-
duced heads and its calycine cup either lacking teeth or exhibiting a great range
of variability in the shape and in the size of the teeth. A bladelike lobe on the
calyx cup of one flower may be an anomaly.

Panama: 6 km from above the Panamerican Highway on the road from El Llano to Carti-Tupile,

Kenneth 1779 (MO), san blas: trail to Darien from Puerto Obaldia, 10-15 km WSW of Puerto Obaldia,
Mori et al. 6859 (MO).

16. Cephaelis latistipulata Standley, J. Wash. Acad. Sci. 18: 281. 1928. type:
Costa Rica, Standley 39695 (US, holotype; F, photo).

Suhshruhs to 1.5 m tall, the stems unbranched, somewhat angular, glabrous,
the internodes well spaced. Leaves obovate, elliptic or ovate, 14-21 cm long,
(0.5-)7-9 cm wide, rounded or obtuse at the apex, the acumen short, acute and
somewhat decurrent on the petiole at the base, the costa prominent, the lateral
veins 1 1-12, coriaceous, smooth, glabrous; petioles elongate, to 7 cm long; stip-
ules persistent, kidney shaped or round, 0.8-2.2 cm long, to 2 cm wide, briefly
bilobed at the apex, the margin undulate and serrate, the lobes to 10 mm long,

acute. Inflorescences in axillary heads, to 1.5 cm long, to 2 cm wide, the peduncle
obsolete; involucral bracts 5, ovate or oblong, to 1.5 cm long, to 1.2 cm wide.
Flowers sessile, the calycine cup with 5 teeth, these lanceolate, 2-5 mm long,
pilose, denticulate; corolla funnel shaped, 4-6 mm long, glabrous or elongate
pilose outside, white pilose within near the mouth, the lobes triangular, 1-2 mm
long, pilose marginally; stamens subsessile, the anthers oblong, ca. 1 mm long;
ovarian disc deeply bilobed, the style 3-5 mm long, glabrous or pilose, the stigmas
1 mm long. Fruits not seen.

Cephaelis latistipulata is known from Costa Rica and Panama. I have drawn
freely on Molina's (1951) description of the flower in the above diagnosis and his
description of the bracts and bracteoles reads: "bracteas involucrales 5, cori-
aceas, enteras o bipartidas, corto-pilosas en ambas caras, obtusas o acuminadas,
ovadas o anchamente oblongas, de 12-15 mm de largo, de 10-12 mm de ancho.

19801 DWYER— FLORA OF PANAMA (Family 179. Rithiaceae) 11

bracteas internas numerosas, pilosas por ambas caras, agudas o acuminadas,
obovadas, margenes sencillo-pilosos, de 10-12 mm de largo, 6-10 de ancho, brac-
teolas lineares o oblanceoladas, subcoriaceas, pilosas por ambas caras, de 6-7
mm de largo, de 1-3 cm de largo."

CHiR[QUi': Cana-Cuasi Trail, Terry 1481 (F).

17. Cephaelis nana (Standley) Standley, J. Wash. Acad. Sci. 17: 171. 1927.

Evea norm Standley, J. Wash. Acad. Sci. 15: 105. 1925. type: Panama, Standley 27550 (US, holotype;
F, isotype).

Subshmbs less than 1 m tall, unbranched, glabrous, the internodes relatively
short. Leaves elliptic to oblong, 9-22 cm long, 5-10 cm wide, obtuse to acuminate
at the apex and the base, the lateral veins prominent, ca. 15, subcoriaceous,
drying green; petioles to 4 cm long; stipules persistent, deeply bilobed, the narrow
lobes 4-6 mm long, usually obtuse. Inflorescences solitary, terminal, subsessile
or the peduncles to 4 cm, subglobose, to 1.5 cm long, to 3 cm wide, containing
3-5 heads; outermost bracts involucral, obovate or spatulate, acute or rounded
at the apex, to 1.5 cm long, to 1.5 cm wide, free, purple, membranous, the internal
bracts oblanceolate or spatulate, rarely narrowly oblong, to 1.2 cm long, to 0.6
cm wide, frequently with minute teeth at the base. Flowers not seen. Fruits
subglobose, to 1 cm long, to 1 cm wide, fleshy, blue, the pyrenes ovoid or elliptic
oblong, ca. 0.5 cm long.

Cephaelis nana is known only from Nicaragua and Panama.

CANAL zone: Hills N of Frijoles, Standley 27550 (F, US).

18. Cephaelis panamensis Dwyer.*^ type: Panama, Mori & Kallunki 3714 (MO,
holotype).

IShriibs, the branchlets woody, smooth, glabrous, the nodes swollen, well
spaced except crowded at the apex of the branchlet. Leaves usually sessile,
verticillately disposed at the apex of the branchlet, elliptic or obovate oblong, 15-
28 cm long, 7.5-15.0 cm wide, acute or deltoid at the apex, the base 1-2 cm wide,
conspicuously truncate, often somewhat auriculate, the costa prominulous above,
prominent beneath, to 0.3 cm wide, the lateral veins 15-17, strongly prominulous
beneath, the lowermost often strict, the upper arcuate, the undulate submarginal
vein inconspicuous, the intervenal areas patulous reticulate, the smaller veins
pinnatiform, chartaceous, concolorous, green when dry, minutely puberulent be-
neath; petioles absent or to 1.3 cm long, lignose, to 0.3 cm wide; stipules decid-
uous, the scars conspicuous, to 6 mm long, to 2.5 mm wide. Flowers not seen.
Inflorescences terminal, the fruits massed into a rounded head, to 3 cm in diam.,
the peduncle absent, the outermost bracts apparently few, triangular oblong, to

'^ Cephaelis panamensis Dwyer, spec. nov. IFrutices ramulis glabris. Folia apice ramuli verti-
cillatim dispositia elliptica vel obovato-elliptica, 15-28 cm longa, 7.5-15.0 cm lata, basi 1-2 cm lata
conspicue truncata saepe auriculata venis lateralibus 15-17 chartacea glabra infra minuto-puberula;
petiolis nullis vel ad 1.3 cm longis. Inflorescentiae terminales fructibus congestis caespite rotundo,
ad 3 cm diam. pedunculo nullo. Friictus sessiles oblongi, ad 1.2 cm longi, glabri.

78 ANNALS OF THR MISSOURI BOTANICAL GARDRN [Vol. 67

1.7 cm long, rigid, glabrous except ciliolate at the apex. Fruits sessile, oblong,
rotund or truncate at the apex, to 1.2 cm long, smooth, glabrous, blackish green
when dry, the calycine cup bowl shaped, ca. 1.5 mm long, to 2.5 mm wide,
truncate, not lobed, rigid, minutely ciliolate, each pyrene drying tan brown with
a slender median groove.

Ccphaclis panamensis is known only from Panama. Unfortunately flowers
were not collected and I am not certain that the species belongs in Cephaelis.
The conglomerate inflorescences appear to be subtended by several bracts. The
fact that some leaves are sessile and are basally truncate or vaguely auriculate
is diagnostic. The relatively large fruit is more suggestive of Cephaelis than of
Psychotna, The calycine cup is toothless and somewhat bowl-shaped.

colon: Rio Guanche, ca. 2.5 km upriver from bridge on road to Porlobelo, tropical wet forest
along small creek to the S of the river, 10-100 m, Mori & KaUunkl 3714 (MO).

19. Cephaelis panchocoensis Dwyer.*** type: Panama, Gentry & Mori 13895 (MO,
holotype).

Succulent suhshruhs, to 0.75 m tall, the stems somewhat smooth, glabrous,
drying red, the superior nodes well spaced. Leaves elliptic or obovate elliptic,
10-35 cm long, 3.5-14.0 cm wide, rounded or obtuse toward the apex, acuminate,
the acumen short and wide, to 1 cm long, acute or attenuate acute basally, the
costa immersed above, prominent beneath, the lateral veins 20-38, plane or prom-
inulous above, leaving the costa at almost right angles and arcuate only near the
margin or strict and sharply ascending, prominulous and conspicuous beneath,
arcuate, the undulate submarginal vein 2-8 mm from the margin, not conspicu-
ously undulate on proximal portion of the blade, the smaller veins inconspicuous,
the intervenal areas smooth, chartaceous to subpapyraceous, discolorous, drying
red to brown beneath, glabrous and minutely farinose above, minutely puberulent
especially on the veins beneath; petioles absent or to 1 cm long; stipules free,
ovate lanceolate, to 1.6 cm long, ca. 0.6 cm wide, obtuse?, drying red, subcori-
aceous, apparently glabrous. Inflorescences terminal, capitate, 2-4 cm long, 2-
6 cm wide, the head appearing solitary or of 2-6(or more?) smaller heads aggre-
gated tightly into a unit head on suppressed branches or several heads on 3
branches terminating a peduncle to 2.5 cm long, stout, to 4 mm wide, glabrous,
drying red; bracts numerous, involucral, the outermost oblong, to 2 cm long,
about V2 as wide as long, obtuse, stiff, venose, glabrous, the margin irregular,
callose, drying tan, with a few glands within. Flowers sessile, the hypanthium
ca. 1 mm long, glabrous, the calycine cup cylindrical, to 3.5 mm long, the teeth

^^ Cephaelis panchocoensis Dwyer, spec. nov. Suffrutices succulenti. Folia elliptica, 10-35 cm
longa, 3.5-14.0 cm lata, versus apicem rotundata vel obtusa venis lateralibus 20-38 chartacea ad
subpapyracea subtus rubra ad brunnea supra glabra sed farinosa subtus in venis minuto-puberula;
petiolis nullis vel 1.0-4.5 cm longis; stipulis ovato-lanceolatis, ad 1.6 cm longis, ca. 0.6 cm latis.
Inflorescentiae terminales capitatae, ad 6 cm longae, glabrae pedunculo nullo vel ad 2.5 cm longo
saepe ramis tribus ad 1.5 cm longis terminato, floribus in unum caespitem ad 2.2 cm in diam. vel in
caespites plures dispositis. Flores subsessiles; hypanthio glabro cupula calycis cylindrica, ad 3.5 mm
longa; corolla alba tubo gracili, ca. 10 mm longo extus glabro; antheris ca. 2 mm longis. Fructus
partim bracteolis inclusi, oblongi, ad 8 mm longi, leviter costati glabri, calyce ad 3 mm longo.

1980] DWYER— FLORA OF PANAMA (Family 179. /?///b,V/(frt<') 79

5, short, obscured by the ciliate margin, the cup glabrous with a few glands
within, these polymorphic, narrow oblong to ovate oblong, to 1.2 mm long; co-
rolla white, narrowly cylindrical, to 10 mm long, ca. 0.6 mm wide in the middle,
glabrous outside, villosulose within near the mouth, the lobes 5, oblong, to 3 mm
long, attenuate above the middle with a minute adaxial projection ca. 1 mm below
the apex; stamens 5, the anthers narrowly oblong, to 2 mm long, subsessile,
attached just below the mouth; ovarian disc oblong, to 0.8 mm long, drying black;
style linear, ca. 14 mm long, the stigmatic area scarcely wider than the style, to
2 mm long. Fruits oblong, to 8 mm long, ca. 4 mm wide, glabrous, red, lightly
costate, the calyx cup persistent or deciduous.

Cephaelis panchocoensis is known only from Panama and Colombia. It is
recognized by its many-veined leaves and terminal capitate inflorescences. The
leaves dry a pink red beneath, have the lateral veins either leaving the costa at
right angles or strict and sharply ascending. The undulate submarginal vein of the
blade becomes subparallel to the margin on the lower Va of the blade. Another
diagnostic character is the callose margin of the bracts. The flowers appear to be
few per head. My description of the flower is based on Gentry et al. 16756. The
calycine cup is strikingly elongate with several glands within. On dissecting the
calycine cup of the fruit of Gentry 4767, I observed several triangular glands
regularly arranged at the base of the cup; these I was unable to find in dissecting
the calycine cup of Gentry 16756. The style is noteworthy in being linear and
extending beyond the elongate corolla tube. The new species is related to Cepha-
elis dressleri Dwyer, described as new in this paper. Cephaelis panchocoensis
is an upland species, while C. dressleri is a lowland species.

PANAMA: DARrrN: La Boca de Pirre, Bristan 1277 (MO). Cerro Pirre base camp on Rio Perre-
cenega, Gentry 4767 (MO). S slope of Cerro Tacarcuna above Rio Pucro base camp, 700-1000 m,
Gentry & Mori 13895 (MO). E of Tres Bocas, shortest headwater of Rio Cuasi, Kirkhride & Duke
1198 (MO). COLOMBIA: choco: Upper Rio Tigre near base of Serrania del Darien, E of Unguia, 250-
300 m, Gentry et al. 16756 (MO).

20. Cephaelis tomentosa (Aubl.) Vahl, Eclog. Amer. 1: 19. 1796.

Tapogomea tomentosa Aubl., Hist. PI. Guiane 160. 1775. type: not seen.
Fsychotria poeppigiana Muell.-Arg. in Mart., Fl. Bras. 6(5): 370. type: not seen.
Evea tomentosa (Aubl.) Standley, Contr. U.S. Natl. Herb. 18: 123. 1916.

Shrubs to 2 m tall, the branches terete, ultimately angular, golden hirsute or
villose, often constricted below the nodes. Leaves lanceolate or ovate lanceolate,
8-23 cm long, 3-9 cm wide, acute at the apex, attenuate acuminate, the acumen
to 2 cm long, acute, rounded or obtuse at the base, the costa prominulous above
and beneath, the lateral veins 9-12, arcuate, the intervenal areas spreading retic-
ulate, membranous to chartaceous, densely golden villose above and beneath;
petioles to 2 cm long, densely hairy; stipules connate, usually persistent, cylin-
drical, the sheath 2-6 mm long, truncate or tapering toward the apex, each part
usually with 2 erect narrowly subulate awns to 11 mm long, usually twice the
length of the sheath, densely hirsute. Inflorescences terminal, 5-6 cm wide, the
flowers conglomerate, the head subtended by numerous bracts, the outer pair
large, flaring, deep red; peduncles 2.5-7.0 cm long, densely villose. Flowers with
the hypanthium short, pubescent, the calycine cup scarcely measurable, the lobes

80 ANNALS OF YUE MISSOURI BOTANICAL GARDLN [Vol. 67

5, narrowly oblong or narrowly triangular subulate, to 2 mm long, petaloid, with
a linear oblong gland at the sinuses to 0.5 mm long; corolla yellow, the tube
narrowly cylindrical to 13 mm long, somewhat expanded near the mouth, petal-
oid, densely elongate ciliate outside, barbate inside ca. 3 mm below the filament
attachment, the lobes 5, triangular, ca. 3 mm long; stamens 5, the anthers sub-
sessile, narrowly oblong, 2.5-3.5 mm long, attached at the mouth, usually slightly
exserted; style slender, included or barely exserted, the stigmas digitiform,
scarcely wider than the style, ca. 2.5 mm long. Fruits berrylike, bright blue,
rotund to broadly oblong, less than 1 cm in diam.

Ccphaclis tomcntosa is well distributed throughout tropical America. Standley
(1938) remarked that ''. . . in most herbaria (it) is represented by more specimens
than perhaps any other tropical member of the family. Oddly enough I have not
seen any herbarium material from the Province of Chiriqui, Panama; undoubtedly
it occurs there." ''Uasika'' (Cuna, fide Standley).

BOCAS DFL TORo: Santa Catalina, Blackuell et al. 2805 (COL, DUKE, MO, UC, VEN). Almi-
rante, near Nigra Creek, McDaniel 5050 (MO). Water Valley, Wcdcl 811 (MO). Chiriquf Lagoon,
WecM 1081, 1279 (both MO). Water Valley, Chiriqui Lagoon, Wedel 1744 (MO). Old Bank Island,
Chiriqui Lagoon, Wedcl 21 18 (MO). Little Bocas, Chiriqui Lagoon, Walcl 2534 (MO), barro Col-
orado island: Avilcs 31 (MO); Croat 4364 (MO); Ehinger 612 (MO); Slwtttak 53, 531. 1045 (all
MO); Starry 155 (MO); Woodson & Schery 987 (MO), canal zonf: Crossing of Cruces Trail and
Madden Dam Road. Bartlctt & Lasser 16344 (MO). Near Juan Mina, Bartlctt & Lasser 16556 (MO).
Cerro Viejo on K 16 C, Blum 1224 (MO). Chagres, 4 mi S of river, Buscy 389 (MO). Madden Forest
Reserve, Candido F. I (MO). Pipeline Road, 0.2 mi N of ATTC Gate, Croat 4697 (MO). Pipeline
Road near Gatun Lake, Croat 4704 (MO). Radar Station Road 1 mi N of Summit Garden, Croat
10276 (MO). Madden Forest SE of Rio Pedro Miguel, D'Arcy & DArcv 6099 (MO). Rio Azote
Caballo, Sabanas, Dodge ct al. 16844 (MO). Hill C-6, Ft. Sherman, Duke 4401 (MO). George Greene
Memorial, Las Cruces Trail, Dwycr 6755 (MO). Chagres, Fcndler 177 (MO). Gatiin Station, Hayes
s.n. (MO). Road between Gamboa and Summit Gardens, Kennedy 2257 (MO). Pipeline Road, Ken-
nedy & Andrews 1896 (MO). Navy Reservation N of Gamboa, Rohyns 65-54 (MO). Las Cruces,
Sefhert 577 (MO). Pipeline Road, Stern & Cluunhers 18 (MO). 6 mi N of Gamboa, Tyson 1510 (MO).
7 mi N of Gamboa, Tyson 6635 (MO). Fort Sherman, Tyson c5 Blutn 1679 (MO). 1 mi N of Summit
Garden, Tyson & Blum 1968 (MO), cocle: El Valle, Aguilar CH3 (MO). Cerro Pilon, 700-900 m,
Duke 12184 (MO). La Mesa, El Valle, Folsom & Butcher 2807 (MO). Ft. Cerro Pilon, ca. 2000 m.
Porter et al. 4584 (MO, SCZ). Cerro Valle Chiquito, 700-800 m, Seihert 505 (MO). El Valle de Anton,
650-700 m, Woodson & Schery 171 (MO). Las Margaritas and El Valle, Woodson et al. 1297 (MO).
colon: Juan Mina Plantation, Rio Chagres, Allen 4109 (MO). Santa Rita Ridge, East Ridge, Dwyer
& Correa 8405 (MO). Santa Rita, N of Agua Clara Rain Gauge, Kennedy 2764 (MO). Santa Rita
Ridge, Lewis et al. 5263 (COL, MO, SCZ, UC, VEN). 0.7 mi NE of Rfo Piedras Bridge, Pilon to
Portobelo, Nee & Mori 3654 (MO). 8 km NE of Santa Rita, ca. 650 ft, Wilbur & Weaver 10840 (MO).
darien: Headwaters of Rfo Chico, 500-750 ft, Allen 4598 (MO). Ri'o Chico, across from Boca de
Tesca, Duke 5214 (MO). Pico Piriaque, Duke 8138 (MO). Ri'o Sancanti, 2 mi upstream from Piria, ca.
120 m, Duke 14383 (MO). Puerto St. Dorothea, Dwyer 2228. 2255 (both MO). Quebrada ^^Camachi-
muricatc*' near Casa de Bartolo. Kennedy 2861 (MO). Manene to Mouth of Ri'o Cuasi, Kirkhride &
Bristan 1543 (MO). Cana, 1750 ft, Stern et al. 654 (MO). LOS Santos: Loma Prieta, Cerro Grande,
2400-2800 ft, Lewis et al. 2240 (COL, DUKE, UC, VEN). Panama: Road to Nuevo Emperador,
Austin 19 (MO). Cerro Azul, Carrasquilla 198 (MO); Castrejon 39 (MO). Cerro Campana, Croat
12139 (MO). Nuevo Emperador, De Garcia 30 (MO). Near Jenene, Duke 3882 (MO). Panama Viejo,
Duke 5730 (MO). Cerro Jefe, Duke 9459 (MO). Cerro Azul, ca. 2000 ft, Dwyer 2124, 3074 (both MO).
Cerro Campana, ca. 3000 ft, Dwyer & Kirkhride 7854 (GH, MO); Ehinger 337 (MO). Cerro Azul,
Ehinger 986 (MO). Nuevo Emperador, Fernandez 116 (MO). Cerro Jefe, Ferris 160 (MO). Cerro
Campana, Gentry 1829 (MO). Ri'o Maestro, 2 mi from coast. Gentry 2205 (MO). Cerro Azul, Gomez-
Pompa et al. 3394 (MO); Guerra 6 (MO). Nuevo Emperador (Arraijan), Guerra 9 (MO). Las Guacas,
Kant 41 (MO). 16 km above Panamerican Highway, road from El Llano to Carti-Tupile, ca. 350-400
m, Kennedy 2531 (MO). Peluca Meterological Station, Kennedy cfe Dressier 2976 (MO). 12-16 km
above Panamerican Highway, El Llano to Carti-Tupile Road, Kennedy et al. 3128 (MO). 2-3 mi S
of Goofy Lake, Lewis et al 261 (F, GH, K, MO, NY, PA, UC, US). Cerro Campana, Lewis et al.

1^80] DWYER— FI.ORA OF PANAMA (Family 179. Rithiaccae) 81

1920 (GH, MO, UC, US). Altos del Rio Pacora, 2500 ft, Lewis ct aL 2302 (MO, UC). Cerro Campana,
McDaniel 6887 (MO). Cerro Jefe, Mori 7108 (MO). Cerro Campana, 45 km SW of Panama City, Mori

& Bolten 7687 (MO). Road past Cerro Azul, Mori & Kallunki 2I82A (MO). 4.5 km N of Lago Cerro
Azul, 675 m, Nee 7037 (MO). Cerro Azul, Porter et aL 4110 (MO, SCZ, UC, VEN). Cerro Campana,
Porter et al. 4160, 4246 (both MO). Capira, Saldana 6 (MO). Cerro Jefe, Tyson 3441. 3590 (both
MO). Cerro Campana, Webster & Breckon 16505 (MO). Cerro Jefe, Wilbur A Weaver 11096 (MO).
Arraijan, Woodson et al. 777, 1400 (both MO), san blas: Between Rio Diablo and Rio Acuatf, Duke
14893 (MO). VERAGUAS: Cerro Tuti, Folsom & Edwards 3343 (MO).

21. Cephaelis vultusmimi Dwyer.^^ type: Panama, Gentry & Dwyer 4811 (MO,
holotype). — Fig. 17.

Siibshriibs to 0.5 m tall, the stem smooth, black or dark brown when dry,
pilose, the hairs golden, slender, to 2 mm long, the nodes well spaced. Leaves
oblong or oblong lanceolate, 9-15 cm long, 3-9 cm wide, deltoid or acute at the
apex, acuminate, the acumen to 2.5 cm long, obtuse or truncate at the base,

rarely subacute, often conspicuously inequilateral, the costa prominulous above,
prominent beneath, the lateral veins ca. 11, strongly arcuate, subprominent be-
neath, not forming an undulate submarginal vein, rigidly chartaceous, bullate but
not drying bullate, concolorous, green with weak curled hairs above, often with
withered stiffer and more elongate hairs beneath, those along the costa to 2 mm
long, perpendicular; petioles 0.5-1.8 cm long, to 0.3 cm wide; stipules persistent,
erect or spreading, free, the body rotund, the lobes 2, erect, widely subulate,
1.5-2.0 cm long, to 0.4 mm wide at the base, caudate tipped, scarious, densely
golden hirsute, the hairs to 2 mm long. Inflorescences terminal, solitary, the
peduncle to 8 cm long, to 0.5 cm wide including the hairs, densely golden-hirsute
or barbate, the hairs to 2 mm long, the head ca. 5 cm wide; the large exterior
bracts connate toward the base, triangular to 3.5 cm long, to 4.5 cm wide, bright
red toward the base, purple to yellow in the middle, green toward the apex,
venose, ciliate on both sides, the hairs similar to those of the leaves. Flowers
with the calycine teeth triangular, unequal, to 1 mm long, hirsute marginally with
a gland between adjacent teeth, these to V^ the length of the tooth.

Cephaelis vultusmimi is known only from Panama. It is the only new species
in this paper without either a complete floral description or a fruit description.
The inflorescence as found within the outermost bracts is very immature. I located
one bud in which only the hypanthium and the calycine teeth were obvious. The
corolla was very immature and little was discernible except the fact that the hairs
were very dense. The bracts are strikingly unique and are suggested by the spe-
cific name vidtusmimi (clown's face). The leaves when seen in the field were
bullate; this feature was lost in drying. A 2 x 2 Ektachrome of the species has

been deposited in the herbarium of the Missouri Botanical Garden.

colon: Santa Rita Ridge Road, E. ridge, Gentry <Si Dwyer 48! I (MO).

^^ Cephaelis vidtusmimi Dwyer, spec. nov. Suffrutices ad 0.5 cm alti, caule dense piloso. Folia
oblonga vel oblongo-lanceolata, 9-15 cm longa, 3-9 cm lata, apice deltoidea vel acuta, acumine ad
2.5 cm longo, venis lateralibus ca. 11, rigido-chartacea, dense hirsuta; petiolis 0.5-1.8 cm longis;
stipulis liberis lobis 2 erectis lato-subulatis, 1 .5-2.0 cm longis, apicem versus caudatis, dense hirsutis.
Infioreseentiae terminales solitariae forma Cephaelis tomentosa simili bracteis exterioribus conspicue
variegatis rubris purpureis luteis viridibusque.

82

ANNALS OF THR MISSOURI BOTANICAL GARDFN

[Vol. 67

Figure 17. Ccphaelts vidlisniimi Dwyer. — A. Habit (x!/i). — B. Vegetative node (xl). [After
Gentry & Dwyer 481 1 .]

16. CHIMARRHIS

Chimarrhis Jacq., Sel. Stirp. Amer. 61. 1763. type: C cymosa Jacq.

Pseudochimarrhis Ducke, Arch. Jard. Bot. Rio de Janerio 3: 255. 1922. type: P. turhinata (A. P.
DeCandolle) Diicke (= Chimarrhis turhinata A. P. DeCandolle).

Trees, Leaves crowded at the apex of the branchlets; stipules caducous. In-
florescences axillary, corymbose cymose, bracteate. Flowers small, odoriferous;

19801 DWYER— FLORA OF PANAMA (Family 179, Ruhiaceae) 83

calyx cup campanulate, reduced, truncate or dentate; corolla white, the tube
short and broad, villous within, the lobes 5, valvate in bud; stamens 5, inserted
in the throat, the anthers dorsifixed, the filaments often exserted, villous at the
base; style short, the stigmas 2, obtuse, ovules numerous on an axile peltate
placenta. Fruits capsular, small, oblong, dehiscent, septicidally into 2 valves,
each valve ultimately divided; seeds numerous, compressed, reticulate, the en-
dosperm fleshy.

Chimarrhis, a genus of about 14 species, is found in the West Indies, Costa
Rica, Panama, and in South America.

a. Leaves 10-20 cm long, 6-11 cm wide; stipules 20-30 mm long; fruits ca. 5 mm long

1. C lot if olio

aa. Leaves 3.5-9.5 cm long, 2-5 cm wide; stipules to 1 1 mm long; fruits ca. 2 mm long

2. C, parvifiora

1. Chimarrhis latifolia Standley, Publ. Field Columbian Mus., Bot, Ser. 4: 265.
1929. type: Panama; Cooper & Slater 260 (F, US).

Trees to 25 m tall, the wood yellow, the buttresses high, the ultimate branch-
lets angular, smooth, glabrous; nodes well spaced, the stipular scars ringlike,
often drying black, the leaf scars ±0.5 cm in diam. Leaves widely elliptic to
elliptic oblong, 10-20 cm long, 6-11 cm wide, rounded at the apex, short acu-
minate, the acumen widely triangular, ca. 0.5 cm long, obtuse, basally cuneate
to abruptly attenuate acute, the costa prominulous above, prominent beneath,
ca, 0,2 cm wide, rubescent, the lateral veins ca. 10, arcuate, conspicuous above
and beneath, uniting to form an undulate submarginal vein, to 0.3 cm from the
margin, stiffly papyraceous to thin coriaceous, discolorous, drying dark chocolate
above, liver colored beneath, somewhat lustrous, glabrescent; petioles to 4 cm
long, angular, twisted, glabrous; stipules deciduous, ovate, 2-3 cm long, acute,
thin coriaceous, rubescent. Inflorescences axillary, paniculate, to 12 cm long, to
11 cm wide, the peduncle to 6 cm long, the branches disposed in a fan, to 3 cm

long, the branchlets in a fan, the corymbs densely flowered. Flowers short ped-
icellate to subsessile; hypanthium turbinate, ca. 2 mm long, glabrous, rubescent,
the calycine cup to 0.8 mm long, somewhat expanded, stiffly petaloid, glabrous,
teeth absent; corolla white, the tube compressed cylindrical, ca. 2 mm long, stiffly
petaloid, rubescent, glabrous outside, white pubescent within above attachment
of the filaments, the lobes 5, oblong, longer than the corolla tube, obtuse, pu-
berulent within; stamens 5, the anthers ovate oblong, ca. 1 mm long, versatile,
the filaments to 4 mm long, tapering to apex, ca. 0.35 mm wide in middle, bearded
adaxially along lower %, attached near the corolla mouth; style to 3.5 mm long,
basally narrow, expanded above, the stigmatic lobes oblong rotund, ca. 1 mm in
diam., crassate. Fruits capsular, bivalvular, to 5 mm long, the outer part forming
a skirt-like structure enclosing the valves, striate, glabrescent.

Chimarrhis latifolia is known from Costa Rica and Panama, The flowers were
described from Allen 6302 (MO), Jalaca Station, Province of Puntarenas, Costa
Rica, which is close to the Province of Chiriqui in Panama. Standley did not
describe the flowers in his original diagnosis.

84 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

The wood was described by the collectors as ''yellow, heavy and hard to split,
the grain very crooked and crossed." '^Jagua amarilla."

CHiRiQui: Progreso, Cooper S: Slater 260 (F, holotype; US, isotype).

2, Chimarrhis parviflora Standley, Trop. Woods 11: 26. 1927. type: Panama,
Cooper & Slater 120 (US, holotype; F, NY, isotypes),— Fig. 18.

Trees to 18 m tall, the branches smooth, terete, glabrous, ultimately angular,
the nodes well spaced. Leaves oblong, 3.5-9.5 cm long, 2.5 cm wide, acute and
short acuminate at apex, acute or cuneate at the base, scarcely inequilateral, the
costa subplane to prominulous above, subprominent beneath, the lateral veins 9-
11, at first strict, then arcuate toward the margin, the intervenal areas smooth,
the veinlets inconspicuous, papyraceous, scarcely discolorous, drying chocolate
brown above, glabrous above, appressed villose especially on the costa and the
veins beneath; petioles 0.5-1.6 cm long, scarcely pubescent; stipules free, nar-
rowly triangular, to 11 mm long, densely appressed pubescent and farinose out-
side, glabrescent within. Inflorescences axillary or pseudoterminal, to 4 per axil,
minutely puberulent, cymose paniculate, to 15 cm long, to 9 cm wide, the pe-
duncle when present 4-9 cm long, arcuate or strict, with 3 or 4 radiate branches
terminating the peduncle 1-3 cm long, the cymules hemispherical, densely flow-
ered. Flowers on short pedicels to 2 mm long; calyx and hypanthium very short,
glabrous truncate the 4-5 teeth obtuse, ciliate; corolla funneliform, ca. 3 mm
long, the lobes 4, rounded, glabrous outside, villose within; stamens 4, exserted,
the anthers narrowly ovoid, to 0.7 mm long, the ovarian disc annular ca. 0.7 mm
long, 0.65 mm wide, 0.8 mm tall; style ca. 2.5 mm long, the stigmas broader than
long. Fruits capsular, obovoid or turbinate, ca. 2 mm long, 1 mm wide or almost
as wide as long, the valves separating almost to the base, the inner face subplane,
scarious, longitudinally grooved, the outer face weakly ribbed; seeds usually 4,
plane, subrotund. 0.5-1 mm in diam. the wing enveloping the central glandular
shiny fruit body.

The species is known only from Panama and Costa Rica. Croat 11244, taken
from a fallen tree, has the inflorescences with galls measuring 5x4 mm in di-
ameter. ''Yema de Huevo.''

BOCAS DRL TORo: Changuinolu Valley, Cooper & Slater 3, 120 (F). Daytonia Farm, Cooper 401 (F,
US). PANAMA: Barro Colorado Island, Croat 11244 (MO), veraguas: Valley of Rio Dos Bocas, road
between Escuela Agricola Alto Piedra & Calovebora, 15.6 m NW of Santa Fe, Croat 27755 (MO).

17. CHIOCOCCA

Chiococca P. Br., Civ. Nat. Hist. Jam. 164, 1756. type: Lonicera alba L. = C.
alba (L.) Hitchc.

Chiococca L., Syst. Nat., ed. 10. 917. 1789. type: C. racemosa L.

Shrubs, vines, or trees, often scandent. Leaves membranous to coriaceous,
petiolate; stipules usually cuspidate, persistent. Inflorescences axillary, racemose
or paniculate, the racemes often secund. Flowers pedicellate; hypanthium com-
pressed; calycine cup 5-lobed, persistent; corolla funnelform or campanulate,

1 9801

DWYER— FLORA OF PANAMA (Family 179. Ritbiaceae)

85

Figure 18. Chimanhis parviflora Standley. Habit (xl). [After Croat Il244.\

glabrous throughout, the lobes valvate; stamens with the filaments connate at the
base; ovary 2-celled, each cell 1-ovulate. Fruits drupaceous, orbicular, usually
laterally compressed, white at maturity,

Chiococca occurs in southern Florida, tropical America, and the West Indies.

a. Vines or sprawling shrubs; stamen filaments included in the corolla tube.

b. Corolla 3-4 mm long; leaves stiffly papyraceous to subcoriaceous, the lateral veins

4_5 1. C. alha

bb. Corolla 6-10 mm long; leaves coriaceous, the lateral veins 5-8 __. 2. C. durifoHa

aa. Upright shrubs or trees; stamen filaments exserted.

c. Leaves tapering at apex; corolla with tube 3.6-6.0 mm long 4. C. phanaestemon

cc. Leaves obtuse or subrotund at apex; corolla with tube ca. 2 mm long 3. C.jefensis

86 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

1. Chiococca alba (L.) Hitchc, Ann. Rep. Missouri Bot. Gard. 4: 94. 1893.

Lonkera aiha L., Sp. PL 175. 1753. type: not seen.

Chioccoca racemosa L., Syst. Nat. ed. 10. 917. 1759. type: not seen.

C. macrocarpa Mart. & GaL, Bull. Acad. Roy. Sci. Bruxelles 11(1): 231. 1844. type: not seen.

C. pctcnsls Lundell, Wrightia 5: 7. 1972. type: Guatemala, Contreras 2439 (LL).

C. vcstita Lundell, Wrightia 5: 8. 1972. type: Belize, Lundell 16425 (LL).

C. vcstita var. glaherrima Lundell, Wrightia 5: 9. 1972. type: not seen.

Vines or shrubs (trees?), the twiglets smooth, glabrous, usually ultimately

wiry, the branchlets perpendicular, opposite. Leaves ovate oblong, oblong, rarely
rotund, 2.2-13,5 cm long, 1.3-6.0 cm wide, tapering acutely, obtuse or somewhat
rounded toward the apex, often acuminate, the acumen often arising from a con-
tracted apex, 1.0-1.5 cm long, acute or obtuse, often falcate, basally acute, at-
tenuate acute, obtuse or rounded, the costa slender, prominulous above and
beneath, the lateral veins 4-5, arcuate, uniting to form a submarginal vein 4-5
mm from the margin in the larger blades, the intervenal areas smooth, the fine
endings of the venules regularly evanescent, stiffly papyraceous to subcoriaceous,
usually glossy above, glabrous; petioles to 0.8 cm long, glabrous; stipules often
persistent, widely triangular to oblong, 1.5-5.0 mm long, acute or contracted at
the apex into a median subulate awn, to 4 times the length of the body. Inflores-
cences axillary, solitary, or often 2 per node, 2,5-8.0 cm long, racemiform or
paniculate, the flowers occasionally solitary or in sessile cymules; peduncles slen-
der, 0.7-7.0 cm long, ca. 0.35 mm wide, glabrous, the lowermost branches to 4.5
cm long. Flowers pedicellate, the pedicels 2-5 mm long; hypanthium subrotund,
somewhat compressed, glabrous, the calycine cup ca. 1 mm long, the teeth acute,
5, about as long as the carnose, glabrous, eglandular cup; corolla white or yellow,

urn shaped, 3-4 mm long, glabrous inside and out, petaloid, the lobes 5, oblong,
slightly shorter than the tube, tapering obtusely; stamens 5, the anthers at first
linear, then narrowly oblong, ca. 4 mm long at anthesis, the filaments slender,
1.2 mm long, pubescent, at the very base connate into a ring adnate to the corolla
tube; style linear, 5-6 mm long, the stigmatic area turgid, to 1.3 mm long, the 2
lobes obtuse. Fruits white, fleshy, compressed, 4-8 mm in diam., the persistent
calyx short.

Chiococca alba ranges from southern Texas through Central America south
to Argentina. It also occurs in the West Indies. ''Lagrimas de Maria.''

BOCAS DEL TORo: Bocas, Maccaw Hills, WcJcl 553 (MO). Water Valley, Wedel 1408, 1732, 1787
(all MO). CANAL zone: Fort Sherman, Croat 14154. 15425 (both MO). Madden Forest, Croat 15266
(MO). Fort Kobbe, Duke 4207. 4716 (both MO). Albrook, Dwycr 6707 (MO). Madden Dam, Dwycr
& Elias 7492 (GH, MO, UC, US); Ehingcr 1960 (MO); Kirkhride 55 (MO). Farfan Beach Road,

Kirkhride & Elias 65 (MO). Rio Petitpie, road to Ft. Sherman from Gatun Locks (S2 or 82), Mori &
Kallunki 2693 (MO). Cocoli, Tyson 1627 (MO). Fort Sherman, Tyson Si Blum 3750 (MO). Curundii,
Tyson & Dwyer 4466 (MO). Fort Kobbe, Woodson ct al. 1415 (MO). Salamanca Hydrographic Sta-
tion, Rfo Pequeni, Woodson ct al, 1594 (MO), chiriqui: Boquete, Davidson 782 (MO), cocle: El
Valle, Woodson ct al. 74L 1253, 1760 (all MO), colon: Portobelo, D'Arcy 4011 (MO). 4 km E of
Buena Vista, Ncc 6796 (MO). Achiote, Tyson ct al. 4538 (MO), darien: Pucro to Rio Pucro, Duke
5369 (MO). Piiias, Duke 10586, 10600, 10639 (all MO). Chepigana, Duke & Bristan 8 [#278 ?1 (MO).
HERR! ra: Las Minas and Pese, Burch ct aL 1322 (GH, K, MO, NY, UC, US). 12.5 mi S of Ocii,
Lewis ct al. 1635 (GH, MO, UC, US). LOS santos: Loma Prieta, Duke 1 1886 (MO). Los Asientos,
Wendchakc 35 (MO). Panama: Nuevo Emperador, Baker 22 (MO). Cerro Campana, Bartlett &
Lasser 16908 (MO). Caldzada Larga, Croat 12392 (MO). Chame, Dodt^c 16731 (MO). Rio Canita,
Duke 3841 (MO). Rio Mamoni, Duke 5681 (MO). Cerro Campana, Duke 8668 (MO). Isia del Rey,

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhuueae) 87

Duke 10422 (MO). Coronado Beach, Duke 1 1814 (MO). San Jose Island, Duke 1 25 17 (MO). Cerro
Azul, Dwyer 1509 (MO). Cerro Campana, Dnyer & Kirkhride 7823 (GH, MO). Cerro Azul, Ebinger
1960 (MO). Cerro Campana, Ebinger I960 (MO). Chagres, Fendler 138 (MO). Panama Vieja, Maurice
797 (MO). Playa de Venado, Mejia 3 (MO). Old Fort Lorenzo, Tyson 1585 (MO). El Llano, Tyson
1742-A (MO). Cerro Azul, Tyson 2093 (MO). Playa Grande, San Jose Island, Tyson & Loftin 5088
(MO). Arraijan, Woodson et id. 1404 (MO), vkraguas: Rio Los Chorros, San Francisco, Dwyer 1354
(MO). Near Ponuga, D\yyer & Kirkbride 7439 (GH, MO, UC, US). Sona, Woodson et at. 545 (MO).
SAN BLAs: Soskatupu, Duke 8967 (MO); Elias 1693 (GH, MO, US).

2. Chiococca durifolia Dwyer. 2" type: Panama, Dwyer 11865 (MO, holotype).

Vines or climbing shrubs, the terminal branchlets opposite, perpendicular,
nigrescent, smooth, glabrous, the nodes well spaced, turgid (persistent leaf bas-
es). Leaves oblong, ovate oblong, lanceolate, 3-13 cm long, 1.3-6.0 cm wide,
deltoid obtuse or rounded at the apex, often with an acumen to 1.2 cm long,
basally rounded, obtuse or cuneate, the costa plane or prominulous, often distally
evanescent above, scarcely evident beneath, the lateral veins 4-5(-8), at first
strict, then strongly arcuate, the submarginal vein well set in from the margin,
coriaceous, smooth, glabrous, usually drying chocolate brown above; petioles to
2 cm long; stipules connate, compressed hemispherical, annular in appearance,
ca. 2.5 mm long, much wider than long, apiculate, subcoriaceous, shiny, glabrous.
Inflorescences axillary, to 8 cm long, often secund, glabrous or glabrate, racemi-
form or scarcely paniculate; peduncle absent or much reduced, to 1 cm long, the
branches opposite, few; flowers solitary or in pairs; bracteoles minute, ovate
subulate; pedicels to 0.6 cm long or obsolete. Flowers with the hypanthium rotund
to subrotund, to 1.5 mm long, glabrate, drying black, the calycine cup scarcely
developed, the teeth 5, erect, subulate, ca. 1.2 mm long, acute, puberulent; co-
rolla white, the tube subcampanulate, 6-10 mm long, thickly petaloid, glabrous
outside, puberulent ca. 2 mm above the base, the lobes 5, deltoid, scarcely shorter
than the tube, acute; stamens 5, the anthers linear, 4-5 mm long, obtuse, the
filaments slender, ca. 2 mm long, pubescent, connate at the base for ca. 0.5 mm;
style slender for almost half its length, expanding above, the stigmas 2, somewhat
unequal, oblong, ca. 1 mm long, the ovules oblong, ca. 1 mm long, obtuse. Fruits
pedicellate, the pedicels shorter than the berry, white, rotund to oblong, strongly
compressed laterally, to 0.8(-10) cm in diam., rounded or obtuse at the base,
smooth, glabrous, delicately ribbed and venose, the persistent calyx coroniform,
ca. 0.5 mm long.

'/'

Wernh

well-defined calycine teeth and the flattened fruit distinguish it from this species,
whose calycine teeth are almost obsolete and whose fruit is nearly globose in
cross section.

'^^^ Chiococca durifolia Dwyer, spec. nov. Vites vel frutices scandentes. Folia oblonga ovato-
oblonga vel lanceolata, 3-13 cm longa, 1.3-6 cm lata, venis lateralibus 5-8 supra evanescentibus,
coriacea laevia glabra; petiolis ad 2 cm longis; stipulis compresso-rotundis, ad 2.5 mm longis. Inflo-
rescentiae pedunculo nullo vel multo reducto. Flores dentibus calycis subulatis, ca. 1.2 mm longis;
corolla alba tubo subcampanulato, 6-10 mm longo. Fructus pedicellis valde compressis, rotundi, ad
0.8(-10) cm diam., plani.

88 ANNAl.S OF- THI- MISSOURI BOTANICAL GARDEN [Vol. 67

coci k: La Mesa above El Valle, Dwyer 11865 (MQ). El Petrosa, 7 km N of El Cope, Alto
Calvario, 900 m, Folsom 1330 (MO), colon: Santa Rita Ridge, Corrca ct Dressier 906 (MO): Dnyer
et al. 9008 (MO); Gentry 1422 (MO). Panama: Cerro Jefe, Correa Si Dressier 467 (MO); Croat 15235
(MO). Cerro Azul, Dwyer 1886 (MO). Cerro Jefe, Ellas S: Hay den 1799 (GH, MO). Panamerican
Highway to Cerro Jefe summit; 2L7 km from Panamerican Highway. Folsoni 3854 (MO). Cerro Jefe,
Kirkhrule ilk Crehhs 9 (MO). Cerro Trinidad, SE slope, Kirkhride & Duke 1649 (MO). Cerro Azul,
Tyson 2088 (MO). Cerro Jefe, Tyson et al. 3207 (MO). Colombia: choco: Bahia Solano, Gentry &
Fallen 17164 (MO).

3. Chiococca jefensis Dwyer.^' type: Panama, Dressier 3296 (MO, holotype). —
Fig. 19.

Trees (?), the branchlets terete, glabrous, the bark cracking, the nodes mod-
erately well spaced. Leaves widely oblong, 5.5-13.5 cm long, 2.5-7.0 cm wide,
obtuse or subrotund at the apex, acuminate, the acumen short, widely triangular,
to 0.8 cm long, acute at the apex and the base, the costa plane or prominulous
above, prominent beneath, to 0.12 cm wide, the lateral veins 7-8, arcuate, the
marginal vein undulate, 1-2 mm distant from the margin, the venules inconspic-
uous above, reticulate beneath, thin coriaceous, scarcely discolorous, farinose
punctate above and beneath; petioles rigid, to 1.5 cm long, to 2.5 mm wide;
stipules (one seen and this juvenile) connate, the sheath cylindrical, turgid, ca.
5 mm long, coriaceous, glabrous, with each part compressed rotund, ca. 1.5 mm
long. Inflorescences terminal, paniculate, to 8 cm long, to 7 cm wide, glabrous
or glabrate, shiny; peduncle absent or to 3 cm long, the branches few, spreading;
bracts and bracteoles not seen; pedicels to 1 cm long, stiff, smooth, to 0.8 mm
wide toward the apex. Flowers glabrous except for the corolla tube within; hy-
panthium short, the calycine cup cupuliform, to 2 mm long, to 3 mm wide, con-
spicuously truncate, subcoriaceous, the lobes absent; corolla white, the tube
compressed cylindrical, ca. 2 mm long, rigidly carnose, the lobes 5, oblong, to
7 mm long, obtuse, conspicuously villose at the base within, otherwise glabrous;
stamens 5, exserted, the anthers oblong, ca. 2 mm long, to 1.5 mm wide before
anthesis, contorted after anthesis, subbasifixed, the filaments thick, 2-4 mm long,
geniculate near the middle, conspicuously villose except near the bend, attached
at the mouth; style thick, to 5 mm long, ca. 0.6 mm wide, the stigmas 2, oblong,
2.2-3.0 mm long, obtuse, each stigma longitudinally bilobate. Fruits rotund, to
1 cm in diam., piano compressed, smooth, glabrous, white, the persistent calyx
to 1 mm long.

Chiococca jefensis is known only from Panama.

Panama: La Eneida, region of Cerro Jefe, Dressier 3296 (MO). Cerro Jefe, on top near Antenna,
ca. 3000 ft, Folsom ci Harp 1373 (MO).

'^ Chiococca Jefensis Dwyer, spec. nov. Arbores (?) ramulis glabris. Folia lato-oblonga, 5.5-13.5
cm longa, 2.5-7.0 cm lata, apice obtusa vel subrotunda acumine brevi lato-triangulari, ad 0.8 cm
longo, venis lateralibus 7-8 tenui-coriacea supra farinoso-punctata; petiolis rigidis, ad 1.5 cm longis,
ad 2.5 mm latis; stipulis connatis vagina turgido-cylindrica, ca. 5 mm longa, ca. 5 mm lata. Inflores-
centiae terminales paniculatae, ad 8 cm longae, ad 7 cm latae, glabrae pedunculo nullo vel ad 3 cm
longo. Flores pedicellis ad 1 cm longis; hypanthio brevi cupula calycis ad 2 mm longa, dentibus nullis;
corolla tubo compresso-cylindrico, ca. 2 mm longo, lobis 5, ad 7 mm longis; staminibus 5, exsertis
antheris oblongis, ca. 2 mm longis, filamentis crassis, 2-4 mm longis; stylo crasso, ad 5 mm longo,
stigmatibus 2, 2.2-3.0 mm longis.

1 980 1

DWYHR— FLORA Ol- PANAMA (Family 179. Ruh'uucae)

89

Figure 19. Chiococca Jefensis Dwyer.— A. Habit (x>/2). — B. Flower (x2.5). [After Dressier

3296.]

90 ANNALS OF THH MISSOURI BOTANICAL GARDLN [Vol. 67

4. Chiococca phaenostemon Schlecht., Linnaea 9: 594. 1834. type: Mexico,
Schlechtendal, not seen. — Fig. 20.

C. staminca Mart. & Gal., BulL Acad. Roy, Sci. Bruxelles II: 231. 1844. typk: not seen.

Shrubs or trees, to 30 m tall, the twigs terete, smooth, glabrous, often nodose
(persistent stipule bases). Leaves oblong or falcate lanceolate, 2.5-10.0 cm long,
0.7-3.3 cm wide, tapering to a narrow cusp, to 1.2 cm long, attenuate acute to
cuneate at the base, usually slightly inequilateral, the costa plane or prominulous
above, prominent beneath, the lateral veins ca. 8, often evanescent, delicately
prominulous, at first strict, then arcuate and forking, the intervenal areas indis-
tinct, subcoriaceous, lustrous, glabrous, delicately marginate; petioles to 2.5 cm
long, ca. 0.12 cm wide; stipules connate, compressed hemispherical or widely
deltoid, to 4 mm long, including the median subulate projection. Inflorescences
terminal or axillary, when terminal about as long as the uppermost leaves and
disposed as a spreading raceme or a narrow panicle, the peduncle 1.5-3.0 cm
long, terminated by 3 or 4 primary branches, 2 or 3 of these longer, bearing short
branches along their length or unbranched and with cymules. Flowers with the
hypanthium oblong rotund, ca. 2.2 mm long, ca. 2.3 mm wide, smooth, glabrous,
the calycine cup arising from a constricted base, ca. 0.8 mm long, glabrous, the
lobes 5, oblong, ca. 1 mm long, convex in cross section, crassate, glabrous but
puberulent marginally; corolla white or yellow, the tube campanulate, 3.6-6.0
mm long, constricted basally, thin carnose, glabrous, the lobes 5, subequal, as-
cending, deltoid, ca. 3 mm long, ca. 1.8 mm wide at the base, glabrous; stamens
5, the anthers linear, ca. 4.2 mm long, ca. 0.15 mm wide, the filaments slender,
ca. 0.5 mm long, barbate, connate for ca. 0.35 mm, not attached to the corolla
tube; ovarian disc doughnut shaped, 0.9-1.0 mm long, 0.5 mm wide, 0.6-0.75
mm high, the style linear subulate, 6.5-9.0 mm long, ca. 0.25 mm wide, narrower
toward the base, the ovary bilocular, the ovules 2, oblong, ca. 1 mm long, pendent
from the apex of the locule. Fruits on pedicels to 0.3 cm long, oblong rotund to
rotund, to 0.5 cm in diam., the calyx persistent, smooth, glabrous, occasionally
delicately ribbed or sulcate.

Chiococca phaenostemon occurs at elevations between 700 and 1800 m in
Mexico, Guatemala, Honduras, Costa Rica, and Panama.

CHiRiguf: Boqucte, Dwycr 6966 (MO). Bambito, 1300 m, Maasola 31 (MO). Boquete, Stern et
al. 2016 (MO). Volcan de Chiriquf, Terry 1359 (MO). Slopes below Cerro Horqueta, along Quebrada
Horqueta, 1600 m, Webster 16704 (MO). Valley of Upper Rio Chiriquf Viejo, White & White 37. 38
(both MO). Finca Lerida, Woodson t^ Schery 224 (MO).

18. CHIONE
Chione DC, Prodr. 4: 461. 1830. typf: C. glabra DC.

Crusea A. Rich., Mem. Mus. Nail. Hist. Nat., Ser. B, Bot. 5: 204, 1834, non Schlecht. & Cham.

1830. type: C. iihhra (DC.) A. Rich.
Saeeonia Endl., Gen. PI. 451. 1838.

Trees or shrubs. Leaves often coriaceous, petiolate; stipules often briefly
connate, small, caducous. Inflorescences terminal, cymose or corymbose, pe-
dunculate; pedicels bracteolate. Flowers with hypanthium turbinate, the calycine

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

91

Figure 20. Chiococca phacnostcnwn Schlecht. — Habit (x'/i). — B. Flower, partly dissected
(x5).— C. Ovary dissected (x5). [After White <&. White 37, \

92 ANNALS OF THL MISSOURI BOTANICAL GARDEN [Vol. 67

cup with 5 lobes, or the margin simply undulate; corolla reduced, funnel shaped,
lacking hairs within, the lobes 5, imbricate, 2 of these exterior; stamens 5, the
anthers large, exserted, dorsifixed, the filaments stout, inserted above the base
of the tube; ovarian disc swollen, the style stout, the stigmas oblong, exserted,
the ovary 2-celled, the ovules solitary, pendulous. Fruits drupaceous, 2-celled;
seeds elongate, the testa membranous.

Chione occurs in Central America and in the West Indies. There are about 15
species in the genus. It is the most poorly collected among the genera of the
Rubiaceae of Panama containing several species.

a. Leaves with the lateral veins 4-9; petioles 0.5-1.5 cm long.

b. Leaves with the lateral veins ca. 4, thickly coriaceous; fruits rotund \. C. huxifoUa

bb. Leaves with the lateral veins 5-9, papyraceous to coriaceous; fruits oblong or obovate

oblong.

c. Leaves with the lateral veins 5-6, the blades neither farinose punctate beneath

nor barbate in the axils.

d. Leaves with the largest blades to 3 cm wide; inflorescences with 2 branch-
es; fruits ca. 2 cm long 2. C. campanensis

dd. Leaves with the largest blades ca. 7 cm wide; inflorescences with 3-5

branches; fruits ca. 1.4 cm long 5. C. panamensis

cc. Leaves with the lateral veins 7-9, the blades either farinose punctate beneath

or barbate in the axils 3. C. costaricensis

aa. Leaves with the lateral veins 13-15; petioles 1.8-3.5 cm long 4, C. daricnensis

1. Chione buxifolia Dwyer & Hayden, Phytologia 15: 55. 1967. type: Panama,
Tyson et aL 3291 (MO, holotype; NY, isotype).

Shrubs 3-5 m tall, glabrous, the branchlets terete, smooth, rimose. Leaves
elliptic or oval elliptic, 2.0-6.5 cm long, 1 .5-3.5 cm wide, deltoid or round obtuse
at the apex, vaguely acuminate, the acumen obtuse, the costa plane above, prom-
inulous beneath, the lateral veins ca. 4, strongly ascending, evanescent beneath
when dry, depressed above, the marginal undulate vein well set in from the
margin, the margin revolute, coriaceous, concolorous, drying deep brown, lus-
trous. Flowers not seen but presumably solitary (the fruits are solitary at the tips
of the twigs). Fruits sessile or on pedicels to 0.5 cm long, rotund, to 1.5 cm long,
0.75 cm wide, black; the persistent lobes of the calyx minute, to 1 mm long; seeds
solitary, large, hard, the endosperm biporous in cross section.

Chione buxifolia is known only from Panama.

Panama: Cerro Jefe, 6.5 km by road N of Goofy Lake, Cerro Azul, forest remnant on slope,
Nee 9308 (MO). La Chorrera, Rodriguez 36 (MO). Cerro Jefe, 2700-3000 ft, Tyson et al. 3291 (MO,

NY).

2. Chione campanensis Dwyer. -^ type: Panama, Wilbur et al. 1020 (MO, holo-
type).

Trees with the branches crowded, smooth, glabrous, the nodes well spaced.
Leaves elliptic oblong, oblong, occasionally ovate oblong, 2.0-6.5 cm long, 1-3

'^'^ Chione campanensis Dwyer, spec. now. Arhores ?. Folia elliptico-oblonga, oblonga vel ovato-
oblonga, 2.6-5.0 cm longa, 1-3 cm lata, apicem versus lato-deltoidea, basi oblusa vel lato-cuneata
venis lateralibus ca. 6, coriacea glabra; pctiolis ad 1 cm longis; stipulis elliptico-oblongis, ad 4 mm
longis, minuto-puberulis. Inflorescentiae patulae pedunculo 1.0-1.2 cm longo, ca. 0.1 cm lato, glabro
ramis 2 angulari-ascendentibus fructibus paucis. Frnctus pedicellis ad 0.8 cm longis, angusto-obtongi,
ad 2 cm longi, ca. 0.7 cm lati, glabri.

19801 DWYER— FLORA OF PANAMA (Family 179. /?«W«rraf') 93

cm wide, widely deltoid toward the apex, acuminate, the acumen widely trian-
gular, to 3 mm long, obtuse, basally obtuse or widely cuneate, often inequilateral,
the costa plane above or scarcely prominulous, prominulous beneath, the lateral
veins ca. 6, prominulous beneath and conspicuous, strongly ascending, often
branching dichotomously, the intervenal areas spreading reticulate, thin coria-
ceous, discolorous, dark red, glabrous; petioles to 1 cm long, ca. 0.1 cm wide;
stipules elliptic oblong, to 4 mm long, acute, erect, stiff, minutely puberulent.

Infl^

1.2 cm long, ca. 0.1 cm wide, gla-

brous, the branches 2, angular ascending, the fruits few (flowers not seen). Fruits
with pedicels ca. 0.8 cm long, carnose, narrowly oblong, to 2 cm long, ca. 0.7 cm
wide, obtuse at the base, glabrous, shiny, the calycine cup persistent, to 1 mm
long, the lobes absent, the margin undulate or with 5 inconspicuous, widely tri-
angular lobes, to 0.2 mm long; disc of the ovary persistent, oblong or ovate
oblong, to 5 mm long, truncate to obtuse at the apex.

Chione campanensis is known only from Panama. It is closely related to C.
buxifolia Dwyer & Hayden whose leaves have few lateral veins and are more
obtuse at the apex. The fruits of C. campanensis are narrowly oblong, while
those of C. buxifolia are rotund.

Panama: Cerro Campana, trail to Campana Ridge, 2300 ft, Wilbur et al. 1020 (MO).

3. Chione costaricensis Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 111.
1940. type: Costa Rica, Austin Smith 1778 (F, holotype).

Chione ciUcnii L. 0. Williams, Phytologia 25: 462. 1973. typk: Costa Rica, Allen 5321 (F, holotype).

Trees 6-7 m tall, the branchlets terete, smooth, glabrous, ultimately angular,
the bark often flaking. Leaves elliptic ovate or lance oblong to oblong rotund,
6.5-14.5(-18) cm long, 3.5-7.0(-l 1.0) cm wide, acute, obtuse or rounded at the
apex, acuminate, the acumen absent or when present widely deltoid, to 0.5 cm
long, basally acute, rounded or obtuse, the costa slender, immersed or plane
above, prominent beneath, the lateral veins 7-9, arcuate, prominulous beneath,
the intervenal areas smooth, spreading reticulate, papyraceous, discolorous, gla-
brous or puberulent, barbate in the axils; petioles to 1.5 cm long, ca. 0.1 cm wide
in the middle, glabrous; stipules deciduous (one seen), connate, the sheath ca.
1.5 mm long, with each part with an obtuse lobe, to 3 mm long, glabrous. Inflo-
rescences terminal, solitary, glabrous, shorter than the terminal leaves, the pe-
duncle 2.0-5.5 cm long, the branches opposite, the pairs few, angular ascending,
the lowermost to 5 cm long; bracteoles minute, subulate. Flowers (fide Standley)
with the corolla tube 5-6 mm long, ca. 0.4 cm wide, somewhat dilated near the
mouth, the lobes 4, round elliptic, almost equal to the tube, glabrous within;
stamens exserted, the anthers ca. 3.5 mm long, narrowly oblong. Fruits pedicel-
late for 1-4 mm, usually with 2 (or more?) often persistent bracteoles at the base,
fusiform oblong, often falcately so, to 2.2 cm long, to 0.8 cm wide, delicately
longitudinally striate, glabrous, capped by a calycine cup, to 1 mm long, the
persistent annular disc protruding for 1 mm in penislike fashion.

Chione costaricensis is known from Costa Rica and Panama.

colon: Salud, Lao & Holdridge 233 (MO).

94 ANNALS OF THR MISSOURI BOTANICAL GARDEN |Vol. 67

4. Chione darienensis Dwyer.^'* type: Panama, Duke 13571 (MO, holotype).

Trees, the uppermost nodes crowded, the branchlets terete, ashen gray.
Leaves oblong lanceolate or obovate lanceolate, 13-24 cm long, 4.5-7.8 cm wide,

acute toward the apex, acuminate, the acumen to 1.3 cm long, subacute or cu-
neate toward the base, scarcely inequilateral at the base, the costa subplane
above, prominent beneath, the lateral veins 13-15, prominulous beneath, the
intervenal areas evanescent above, spreading reticulate beneath, papyraceous,
shiny, glabrous above except for the costa and the veins, the hairs small, white,
beneath sparsely appressed villose, barbate in the axils; petioles rigid, 1.8-3.5 cm
long, often twisted; stipules not seen. Inflorescences (only one seen) axillary, to
13 cm long, ca. 11 cm wide, the peduncle slender, 4.5-6.0 cm long, arcuately
disposed, terminated by 2 branches, these short and slender, to 1.5 cm long,
appressed ciliate, with the scars of the pedicels ca. 6 per branch; bracts ovate,
to 1.5 mm long, acute, concave, the margins ciliolate. Fruits falcate oblong or
obovate oblong, to 2.2 cm long, attenuate toward the base, often minutely pu-
berulent, the calycine cup to 1 mm long.

Chione darienensis is known only from Panama,

darikn: Rio Balsa, between Manenc and Tusijuanda, Duke 13571 (MO).

5. Chione panamensis Steyermark, Ceiba 3: 19. 1952. type: Panama, Hagen &
Hagen 2137 (NY, holotype; F, isotype).

Trees or shrubs, to 25 m tall, the branchlets numerous, terete, smooth, gla-
brous, the nodes well spaced, somewhat turgid. Leaves oblong or obovate-ob-
long, 3.5-9.5(-10.5) cm long, 2-7 cm wide, widely deltoid to obtuse at the apex,
cuneate, obtuse, rounded or truncate at the base, the costa prominulous and
grooved above, scarcely prominent beneath, the lateral veins 5-6, opposite,
strongly ascending, often forking well below the margin, coriaceous, lustrous,
mostly drying dark brown, the margin often revolute on drying; petioles to 1.5
cm long, rigid; stipules not seen. Inflorescences terminal, soHtary, glabrous, cy-
mose corymbose, 4.0-8.5 cm long, 3-7 cm wide, the peduncle 2.5-4.0 cm long,
angular, usually terminated by 3-5 flabellately disposed branches or the branches
opposite, few and well spaced, the lowermost to 3.5 cm long; bracts deciduous.

Flowers with the hypanthium and calycine cup turbinate, ca. 3 mm long, coria-
ceous, glabrous, the cup eglandular within, the teeth not evident; corolla white,
broadly cylindrical, ca. 4 mm long (in bud), wider than long, carnose, glabrous,
the lobes 5, oblong rotund to compressed triangular, 3.3-4.0 mm long, often wider
than long, obtuse or rounded at the apex, carnose, glabrous; stamens 5, the
anthers widely oblong, ca. 3.5 mm long, the filaments ca. 3 mm long, thick; style
subclavate, ca. 6 mm long, ca. 1 mm wide, the stigmas 2, oblong, ca. 1.5 mm

^"^ Chione darienensis Dwyer, spec, now Arhores. Folia oblongo-lanccolatae vel obovato-lanceolala,
13-24 cm longa, 4.5-7.8 cm lata, acumine ad 1.3 cm longo, venis laleralibus 13-15, papyracea nitida

supra glabra praeter costam venasque; petiolis rigidis, 1.8-3.5 cm longis. Infloresvenliae axillares, ad
13 cm longae, ca. 1 1 cm latae, pedunculo gracili, 4.5-6.0 cm longo; bracteis ovatis, ad 1.5 mm longis.
Fructus falcalo-oblongi vel obovato-oblongi, ad 2.2 cm longi.

1980] DWYER— FLORA OF- PANAMA (Family 179. Riihiciceae) 95

long, truncate. Fruits red, oblong, glabrous, to 1.4 cm long, marcescent, glandular
shiny, glabrous.

Chione panamcnsis is known only from Panama.

CHlRiQUi: Cerro Horqueta, Blum & Dwycr 2651 (MO); Hagen & Hagen 2137 (F); Kirkhride 158
(MO). PANAMA: El Llano-Cartf Road, 18 km from the Interamerican Highway, Mori & Kalliinki 5125
(MO). SAN BLAS: El Llano-Carti Road. 24.5-25 km from the Interamerican Highway, Mori c& Kallunki
5533, 5557 (both MO).

19. CHOMELIA
Chomelia Jacq., Enum. PI. Carib. 12. 1760. type: C. spinosa Jacq.

Anisoineris Presl., Symb. Bot. Upsal. 2: 5. 1833. type: A. spiiio.sa C. B. Presl.

Shrubs or trees, occasionally armed, the branchlets occasionally reduced in
length. Leaves with intervenal areas occasionally lineolate; stipules usually per-
sistent. Inflorescences cymose, axillary, the peduncle occasionally reduced in
length; bracts and bracteoles small, the latter occasionally connate. Flowers tu-
bular, the teeth persistent; corolla white or cream colored, the corolla salverform
or funnel shaped, usually sericeous outside, the lobes valvate or imbricate; sta-
mens generally not exserted, the anthers dorsifixed, sessile, attached near the
mouth; ovary 2-celled, each cell with a pendulous ovule. Fruits fleshy, the seeds
cylindrical, pendulous.

Chomelia has over 400 species, about 50 of which occur in Central and South
America. The genus has the majority of species in tropical Africa and tropical
Asia. It also occurs in the Seychelles Islands.

There are only 4 genera among the Panamanian Rubiaceae which are armed.
Apparently only one of these, Uncaria, is consistently armed. Most species of
Randia have spines. In the case of Machaonia and Chomelia only rarely are the
species with spines.

Literature:

Steyermark, J. S. 1967. Chomelia in Botany of the Guayana Highland. Mem
New York Bot. Card. 17: 333-334.

a. Plants armed.

b. Leaves with intervenal areas not lineolate 9. C spinosa

bb. Leaves with intervenal areas lineolate L C. atkintica

aa. Plants unarmed.

c. Leaves with intervenal areas lineolate L C. atlantica

cc. Leaves with intervenal areas not lineolate.

d. Sprawling shrubs: leaves with lateral veins ca. 6 7. C. psilocarpa

dd. Trees or upright shrubs; leaves with lateral veins 6-13(-18).

e. Leaves ashen white beneath; blades 7.0-18.5 cm long 4. C, leucophylla

ee. Leaves not ashen white beneath; blades 4.5-12.0 cm long,
f. Flowers and fruits not secundly disposed.

g. Inflorescence with peduncle scarcely measurable 8. C. recordii

gg. Inflorescence with peduncle 1-3 cm long,
h. Leaves coriaceous; lateral veins 7-9.

i. Leaves with petioles 0.5 cm long; stipules to 0.35 mm

long 3, r. grandkarpa

96 ANNALS OF THR MISSOURI BOTANICAL GARDLN [Vol. 67

ii. Leaves with petioles to L5 cm long; stipules to 1 cm

long

2. C. coclcnsis

hh. Leaves chartaceous; lateral veins 6 6. C. pcninsulatis

ff. Flowers and fruits secundly disposed 5. C. pananicnsis

1. Chomelia atlantica Dwyer.-^ type: Panama, Kennedy & Gra 2242 (MO, ho-

lotype).

Trees or shrubs to 6 m tall, the branchlets short or long, to 13 cm long, the
bark gray, densely appressed pubescent or glabrescent, the nodes well spaced,
at times armed, the spines solitary, to 0.4 cm long, rigid. Leaves oblong, 3-14
cm long, 1.5-7.5 cm wide, deltoid toward the apex, acute, basally attenuate acute,
rarely subcordate, sometimes inequilateral, the lateral veins 8-10, strongly ar-
cuate, the intervenal areas conspicuously lineolate, chartaceous, discolorous,
pubescent above on the costa and veins, with elongate appressed hairs on the
costa and the veins beneath, the hairs of the intervenal areas small, strigose;
petioles slender, to 2.2 cm long; stipules free, often deciduous, narrowly trian-
gular, to 10 mm long, acute with a median keel, densely pubescent especially
toward the base. Inflorescences axillary, perhaps pedunculate; flowers ca. 3;
bracteoles connate at the base of the hypanthium, each part provided with a
conspicuous bristle to 2.5 mm long. Flowers green or cream colored, the hypan-
thium oblong rotund, ca. 2 mm long, the hairs long, appressed, white, the calycine
cup to 1.5 mm long, eglandular within, truncate, the lobes 4-5, subulate, to 6 mm
long or longer, often curled; corolla with the tube 1.7-2.0 cm long, ca. 0.1 cm
wide in the middle, with appressed, white and dense hairs outside, glabrous
within, the lobes 4, narrowly ovate oblong, ca. 8 mm long, ca. 2 mm wide at the
base, with a median keel, the latter with elongate, appressed hairs; stamens 4,
somewhat included, the anthers narrowly oblong, 3.3 mm long, obtuse, the fila-
ments ca. 0.5 mm long, attached near the mouth, the style linear, 4-1 1 mm long,
the stigmas 2, ca, 1 mm long. Fruits not seen.

Chomelia atlantica is known only from Panama. It is immediately distin-
guished by the intervenal areas of the leaves being lineolate and the inflorescence
being epedunculate. It is probably related to C. tenuiflora Bentham from Surinam,
British Guiana, and Venezuela, which also has the bracteoles united below the
hypanthium and a suppressed peduncle.

colon: Swamp forest and bordering tierra firma, ca. 0.5 km from Ri'o Buena- Ventura, near
Portobelo, Kennedy & Gra 2242 (MO), dariln: Cerro Pirre, Btistan 567 (MO). Cina-Cuasf trail
(Camp 2), Chepigana, Terry & Terry 1448 (F, MO).

-^ Chomelia atlantica Dwyer, spec. nov. Arhores vel frutices ad 6 m alti, interdum armati. Folia
oblonga, 3-14 cm longa, 1.5-7.5 cm lata, ad apicem deltoidea venis lateralibus 8-10 interveniis con-
spicue lineolatis subtus ciliis elongatis appressis in costa venisque praedita; petiolis ad 2.2 cm longis;
stipulis ad 1 cm longis. Inftoresccntiae fortasse epedunculatae floribus ca. 3 bracteolis connatis cu-
puliformibus basi hypanthii ad 1 mm longis, seta ad 2.5 mm longa. Flores cupula calycis ad 1.5 mm
longa, truncata, dentibus 4-5 subulatis, ad 6 mm longis; corolla viridi vel cremea tubo ad 20 mm
longo, lobis ca. 8 mm longis; antheris ca. 3.3 mm longis. Fruetus non visi.

1980) DWYER— FLORA OF PANAMA (Family 179. Ruhiaccac) 97

2. Chomelia codensis Dwyer.-^* type: Panama, Croat 25396 (MO, holotype).

Shrubs to 3 m tall, the branchlets terete, somewhat smooth, lightly puberulent,
ascending at a sharp angle, the nodes well spaced, the lenticels many but incon-
spicuous. Leaves ovate-oblong or oblong, 4.5-10.5 cm long, 2-5 cm wide, deltoid
or subobtuse at the apex, acuminate, the acumen triangular to 0.8 cm long, ul-
timately obtuse, obtuse or rounded at the base, the costa prominulous above,
somewhat prominent beneath, the lateral veins ca. 7, arcuate, conspicuous be-
neath, the smaller veins not conspicuous, coriaceous, discolorous, dark brown
and glabrous above, beneath with a few white hairs especially on the costa and
the veins; petioles rigid, woody, to 1.5 cm long, to 1.5 mm wide; stipules decid-
uous, only seen at the apex of the twigs, rhombic oblong, to 1 cm long, 0.6 cm
wide, acute, subcoriaceous, puberulent, the veins crowded, fanlike. Inflores-
cences subterminal, axillary, cymose paniculate, to 7 cm long, to 3 cm wide, the
peduncle to 3 cm long. Flowers with the hypanthium oblong, glabrous, ca. 1 mm
long, the calycine cup expanded, 0.1 mm long, the lobes 4, triangular, 0.1 mm
long; corolla white, the tube cylindric funneliform, ca. 1.5 mm long, petaloid,
glabrous without, densely white villose toward the mouth, the lobes 4, ovate, to
2.5 mm long, to 1.8 mm wide, ultimately obtuse; stamens 4, the anthers oblong,
to 0.7 mm long, dorsifixed, the filaments scarcely longer than the anthers, at-
tached near the mouth; stigmas 2, oblong, to 0.3 mm long, obtuse, the style ca.
2 mm long, the ovules one per locule, pendent. Fruits not seen.

Chomelia codensis is known only from Panama. It is recognized by its cori-
aceous leaves and rigid, elongate peduncles. The flowers are much reduced in
length.

cocle: La Mesa above El Valle, 800 m, Croat 25396 (MO).

3. Chomelia grandicarpa Dwyer.'-** type: Panama, Kennedy & Foster 2174 (MO,

holotype).

Trees to 6 m tall, the branchlets terete, glabrescent, smooth, moss covered.
Leaves oblong, ovate-oblong or ovate, 4.5-1 1.0 cm long, 2.5-5.5 cm wide, deltoid
to subacute at the apex, acuminate, the acumen to 1 cm long, ultimately acute,

=^^ Chomelia codensis Dwyer, spec. nov. Frutices ad 3 m alti. FoUa ovato-oblonga vel oblonga,
4.5-10.5 cm longa, 2-5 cm lata, versus apicem deltoidea vel subobtusa basi obtusa venis lateralibus
ca. 7 coriacea; petiolis ad 1.5 cm longis; stipulis rhombico-oblongis, ad 1 cm longis. Infloresceutiae
ad 7 cm longae, ad 3 cm latae, pedunculo rigido, ad 3 cm longo. Flores hypanthio glabro cupula
calycis ca. 0.1 mm longa; corolla alba lubo ca. 1.5 mm longo, lobis 4 ad 2.5 mm longis; antheris ad
0.7 mm longis. Fructus non visi.

^** Chomelia grafulicarpa Dwyer, spec. nov. Arhores ad 6 m altae. FoUa oblonga, ovato-oblonga
vel ovata, 4.5-11.0 cm longa, 2.5-5.5 cm lata, venis lateralibus 6-8 venis venulisque minoribus non
visis, rigido-papyracea ad tenui-coriacea, in costa subtus villosa; stipulis minutis triangularibus; pe-
tiolis ad 0.5 cm longis. Inflorescentiae axillares pedunculo forte gracile, ad 4.5 cm longo, floribus 2-
5 in ramis 2 forte reductis dispositis. Flores sessiles cupula calycis carnosa extus appresso-villosa
lobis 4 inconspicuis. Fructus oblongi, ad 3.2 cm longi, glabri rubri carnei. seminibus 2 oblongis. ad
2 cm longis, proxime apicem Iruncalis ad basin attenuatis conspicue rostratis in sicco purpureo-nigris
laevibus inconspicue sulcatis.

98 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

rounded or obtuse at the base, the costa slender, prominulous above and beneath,
drying red, the lateral veins 6-8, arcuate, forming an undulate, submarginal vein
well in from the margin, the intervenal areas smooth, the smaller veins not visible,
rigidly papyraceous to thin coriaceous, drying deep red brown above, lighter
beneath, glabrous above and beneath but moderately villose on the costa below;
petioles stiff, terete, to 0.5 cm long, glabrate; stipules persistent only at the tips
of the twiglets, triangular, to 3.5 mm long. Inflorescences axillary, the peduncle

slender, 1.2-4.5 cm long, ca. 0.15 mm wide, often curved, pubescent, terminated

by 2 flowers or by 2 short, spreading branches, to 8 mm long, each branch with
2-5 flowers; bracts and bracteoles not seen. Flowers (in young bud) sessile, the

hypanthium scarcely measurable, the calyx urn shaped, ca. 1 mm long, carnose,
appressed pilose outside, the margin subtruncate with 4 compressed-rotund lobes,
the latter scarcely measurable; corolla tube cream colored or pink and streaked,
carnose, pubescent outside, the lobes valvate; ovules pendent, one in each of the
2 cells, the style cylindrical. Fruits presumably rotund when fresh, fleshy, red,
when dried oblong, to 3.2 cm long, to 2.5 cm wide, the seeds 2, oblong, to 2 cm
long, wider at the apex than at the base, truncate toward the apex, conspicuously
rostrate, the beak triangular, to 0.4 cm long, obviously attenuate toward the base,
drying purple black, smooth, glabrous, vaguely sulcate.

Chomelia grandicarpa is known only from Panama. The fruit is of extraor-
dinary size for a Chomelia. and in the pressed condition appears like a young
fruit of Pterocarpus (Leguminosae). This character, coupled with the few flowers
per inflorescence and the subcoriaceous texture of the leaves, readily segregates
the species from the remaining species in Panama. The fact that Folsom 3696 has
pink flowers that are streaked is noteworthy.

colon: Santa Rita Lumber Road, 8.7 km from Transislhmian Highway, Folsom 3696 (MO). 2-3
mi up Rfo Guanche, lowland rain forest, 10-20 m, Kennedy iSi Fosicr 2174 (MO).

4. Chomelia leucophylla Dwyer.'-^ type: Panama, Gentry 4262 (MO, holotype). —
Fig. 21.

Small trees, 4-10 m tall with the trunk branching above, the twigs terete,
rigid, pubescent, the nodes well spaced. Leaves oblong elliptic, oblong or sub-
rotund, sometimes falcate, 7.0-18.5 cm long, 2.2-1 1.0 cm wide, acute or deltoid
at the apex, the acumen widely triangular, to 1 cm long, often falcate, at the base
rotund or widely cuneate, the costa slender, prominulous above and beneath, the
lateral veins 7-9, arcuate, thin chartaceous, discolorous, brown above, ashen
white beneath, glabrous above except for the costa and veins, with the hairs
beneath crowded, minute, white; petioles 0.5-2.3 cm long, to 0.13 cm wide;
stipules free, narrowly lanceolate, 5-7 mm long, acute, the hairs curled. Inflo-

-' Chomelia leueophylUi Dwyer, spec. nov. Arhores 4-10 m altae. Folia oblongo-elliptica, ob-
longa vel subrolunda, 7-18.5 cm longa, 2.2-11 cm lata, venis lateralibus 7-9 tenui-chartacea supra
brunnea infra albo-cinerea; petiolis ad 2.3 cm longis; stipulis 5-7 mm longis, acutis. Inflorescentiae
ad 3 cm longae, pedunculis 1-1.5 cm longis. Flores sessiles vel pedicellis ad 1.5 cm longis; cupula
calycis brevi lobis 4 triangularibus subulatis vel foliaceis, 0.7-3 mm longis; corolla tubo ad 13 mm
longo, lobis 2.5-3 mm longo; antheris ca. 2.2 mm longis. Fructits oblongi, ad 7 mm longi, puberuli.

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

99

■.■z>.

FiGURH 21. Chomclia Icucophylla Dwyer. — A. Habit (x4.5). — B. Flower, opened to show in-
terior (x2.8). — C. Ovary cross section (x7.5). [After Holdndge 6452.]

resccnces axillary, to 3 cm long, the peduncles 1.0-1.5 cm long, slender, white
puberulent, the flowers congested on short branches, the bracts and bracteoles
small. Flowers sessile or subsessile, to 1.5 cm long, the hypanthium rotund, ca.
1.5 mm long, densely pubescent, the cup of the calyx short, provided with a few
minute red glands within, the lobes 4, triangular, subulate or foliaceous, 0.7-3.0
mm long, acute, glabrous within, often reflexed; corolla with the tube narrowly

100 ANNALS OF THE MISSOURI BOTANICAL GARDLN [Vol. 67

cylindrical, 10-13 mm long, 0.8 mm wide in the middle, puberulent outside, pu-
bescent within near the base, the lobes 4, subrotund, 2.5-3.0 mm long, with the
margins crisp, glabrous within; stamens 4, included within the tube, oblong, 2.2
mm long, obtuse, the filaments short, ca. 0.5 mm long, attached slightly below
the mouth; style to 5.5 mm long, ca. 0.25 mm wide, the stigmas narrowly oblong,
ca. 3 mm long. Fruits immature, sessile, oblong, to 7 mm long, to 3.5 mm wide,
truncate above, tapering at the base, puberulent, drying brown, with 6 ribs.

Chomclia Iciicophylla is known only from Panama. It is readily recognized by
the white undersurface of the leaf blades as suggested by the specific epithet.

colon: Lower Rio Guanche, Dressier 4346 (MO). Rio Trapiche. Holdrid^e 6452 (MO). Rio
Guanche, ca. 3 km upriver from bridge, road to Portobelo, Mori & Kallunki 5181 (MO), daru-n:
Cerro Chucula, Gentrx 4262 (MO).

5. Chomelia panamensis (Standley) Dwyer, comb. nov.

Antirhca panamensis Standley, N. Amer. Fl. 32: 264. 1934. typr: Panama, Cooper 238 (F. holotype;

NY, isotype).
Chione chamhersii Dwyer & Hayden, Ann. Missouri Bot. Gard. 54: 138. 1967. typf: Panama, Stern

iSc Chambers 144 (MO, holotype).

Trees of medium height, to 12 m tall, the bole slender, the crown small, the
twigs smooth, the branchlets smooth, rimose, glabrescent, occasionally lenticel-
late, the leaves often persistent and verticillate at tips. Leaves ovate, oblong,
elliptic oblong, occasionally oblong rotund, 6-13(-18) cm long, 3.0-6.5 cm wide,
apically acute and acuminate or obtuse and abruptly acuminate, basally cuneate
or obtuse, the lateral veins 7-8, usually immersed above, prominulous beneath,
the intervenal areas often striate lineolate, stiffly membranous to rarely stiffly
papyraceous, concolorous or not, glabrate or short pilose on the costa and the
veins and often with short strigose appressed hairs in the intervenal areas; petioles
slender, 1.0-2.5(-5.0) cm long; stipules deciduous, ovate elliptic, 4.0-5.5 mm
long, attenuate at apex, glabrous to puberulent, ciliate marginally. Inflorescences
terminal and/or axillary, cymose, shorter than or slightly longer than the upper-
most leaves, the peduncles to 6 cm long, 0.4-0.8 mm wide, strigillose, twisted,
the first 2 branches from apex of peduncle short and soon bifurcating to form
branches to 3.5 cm long, often conspicuously spreading, the flowers several,
secundly disposed, ca. 1 mm apart. Flowers sessile; hypanthium oblong, 1 mm
long, glabrous to puberulent, the calycine cup and the 4-5 oblong-rotund lobes
ca. 0.5 mm long, glabrous within, the lobes minutely ciliolate, carnose; corolla
greenish yellow, the tube ca. 7.5 mm long, 0.6 mm wide at the middle, puberulent
outside, glabrous within except hirsute at very base of the tube, carnose, the
lobes 4, oblong rotund, 1.5 mm long, or 3 of 4 equal in width, compressed rotund
to oblong, 1 mm long and the 4th oblong, 1.8 mm long, carnose, glabrous within;
stamens 4, the anthers linear oblong, ca. 3 mm long, conspicuously sagittate at
the base, subbasifixed, the filaments ca. 0.5 mm long, attached to the tube 2-3
mm below the mouth; ovarian disc hemispherical, ca. 0.3 mm long, the style
linear, 6-8 mm long, the 2 stigmas narrowly oblong, slightly wider than the style,
0,7 mm long, the ovary thick walled, the ovules 1 per cell, pendulous, narrow
oblong, ca. 0.7 mm long. Fruits sessile, fleshy, oblong, 1.0-1.3 cm long, rounded,

1980] DWYER— FLORA OF PANAMA (Family 179. /?///)(V;rffl(') 101

obtuse or truncate at the apex, smooth, glabrous, lightly costate, the ribs irregular
and inconspicuous, black, and drying black; the calyx not persistent, the scar
evanescent; seeds 2, enclosed by a very thick, hard endocarp.

Chomelki panamensis is known only from Panama and Colombia. The wood,
bright orange in color, is used in interior construction, for axe handles, paddles,
etc. The fruit, while juicy, is not eaten (fide Duke). "Tablis"; 'Tapaliso"; 'Ta-
palizo."

( HiRiQuf: Puerto Armuelles. Stern & Chambers 144 (MO), darien: Rio Ucurganti, Bristun 1167
(MO). Rio Yape, Bristan 1372 (MO). Without other locality. Bristan & Duke 179 (MO). Santa Fe,
Rio Sabana, Duke 4119 (MO). Rio Chico across from Boca de Resca, Duke 5220 (MO). Santa Fe,
Duke 8319 (MO). Rio Balsa between Quebrada Chusomocatre and Rio Areti, Duke 8709 (MO). N of
Pucro, Duke 13049 (MO). Rio Sabana. above Santa Fe, Duke 14098 (MO). Rio Congo, Holdrid^e
6290 (MO). Foothills, N slope of Cerro Pirre, Mori tS Kalluuki 5412 (MO). Campamento Buena Vista,
Rio Chucunaque above Rio Tuquesa, Stern et al. 825 (MO). 3 mi N of Santa Fe, Tyson et al. 4639 (MO).
3 mi E of Santa Fe, Tyson et al. 4704 (MO). 2 mi E of Santa Fe. Tyson et al. 4842 (MO). LOS santos:
Loma Prieta, 800-900 m, Duke 11889 (MO). Panama: Between Cafiasas and Sabalo, Duke 14472
(MO). SAN bias: Perme, G. P. Cooper 238 (F). Colombia: choco: 1 mi NE of Camp Curiche, 3 mi
E of Curiche, Duke & Idroho 11320 (MO, NY).

6.

(MO

Shrubs to 1 .3 m tall, the branchlets numerous, terete, glabrous, the tips villose,
densely lenticellate, the nodes well spaced. Leaves oblong to vaguely oblong
rotund, occasionally ovate, 5.5-15.0 cm long, 2.8-8.0 cm wide, obtuse to deltoid
at the apex, the acumen triangular, ca. 1 cm long, ultimately acute or obtuse,
acute to cuneate at the base, the costa prominulous above and beneath, the lateral
veins ca. 6, rigid and angular ascending for about ¥4 of their length then curving
arcuately to form an undulate vein to 1 cm from the margin, the smaller veins
between the laterals scarcely visible, stiffly chartaceous, somewhat discolorous,
drying dark green above, smooth, glabrate with a few hairs on the costa, often
barbate in the vein axils beneath, the hairs dense-ferrugineous; petioles stiff, 0.3-
0.8 cm long, ca. 1 mm wide in the middle, sparsely villose; stipules triangular, to
4 mm long, stiff, appressed pubescent. Inflorescences axillary or pseudo terminal,
usually 2 per node, the peduncles wiry, 1 .8-2.5 cm long, to 0.4 mm wide, glabrate,
the peduncle ending as 2 spreading branches, the flowers 15 to 25 per inflores-
cence, disposed as a few secund cymules or solitary, sessile or subsessile. Flow-
ers with the hypanthium oblong or turbinate, ca. 1.5 mm long, obtuse or rounded
basally, the hairs few, the calycine cup scarcely measurable, the teeth usually 5,
triangular or compressed rotund, to 0.3 mm long or inconspicuous; corofla white
(in bud) to 6 mm long, densely white puberulent outside, carnose, barbate basally

^8 ChomeUa peninsularis Dwyer, spec. nov. Frutices ad 1.3 m alti ramulis crebris teretibus. Folia
oblonga vel oblongo-rotunda interdum ovata, ad 15 cm longa, ad 8 cm lata, basi acuta vel cuneata,
venis lateralibus ca. 6, vena submarginali undulata a margine bene distante, rigido-chartacea laevia
solum in costa et venis sparse pubescentia axillis saepe barbatis; petiolis 0.3-0.8 cm longis, sparse
villosis; stipulis triangularibus, ad 4 mm longis, appresso-pubescentibus. Inflorescentiae axillares vel
pseudo-terminales, pedunculis gracilibus, 1.8-2.5 cm longis, glabrescentibus ramis duobus brevibus
terminatis floribus 15-25 cymulis paucis vel floribus solitariis secunde dispositis, bracteis deciduis.
Flores sessiles vel subsessiles, hypanthio ca. 1.5 mm longis, dentibus inconspicuis vel 5 et ad 0.5 mm
longis, obtusis vel acutis. Fructus non visi.

102 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

within, the lobes valvate, to 2 mm long; stamens 5, the anthers oblong, ca. 1.3
mm long, obtuse, usually exserted, the filaments shorter than the anthers; style
slender, ca. 7 mm long, the stigmas thumblike, ca. 2 mm long; ovules 1 per cell,

pendulous. Fruits not seen.

Chomelia peninsidaris is known only from lowland Chiriqui. It is readily dis-
tinguished from C. panamensis (Standley) Dwyer, a large tree which is distributed
primarily in Darien Province and in the neighboring Province of Choco, Colombia.
The new species is a shrub with thicker leaves and with fewer lateral veins.

chiriqui: Burica Peninsula near Punta de Piedra, secondary forest, Croat 22440 (MO).

7. Chomelia psilocarpa Dwyer & Hayden, Ann. Missouri Bot. Gard. 54: 139.
1967. type: Panama, Hayden 143 (MO).

Scandent shrubs, often in treetops, the branches glabrous, smooth, often
drying white. Leaves elliptic, 6.5-13.0 cm long, 2.5-6.0 cm wide, acute at the
apex, acuminate, the acumen to 1 cm long, apiculate, the apiculum ca. 1 mm
long, attenuate acute at the base, the costa prominulous beneath, the lateral veins
ca. 6, alternate, strongly arcuate, papyraceous, glabrate above, the hairs short,
white, deciduous, beneath ciliolate, the veins appressed pilose, with the shorter
hairs seriately disposed, densely barbate in the axils of the veins; petioles 0.7-
2.0 cm long, appressed subhirsute; stipules triangular or subulate, to 1 cm long,
ca. 0.35 cm wide at the base, elongate acuminate, smooth, glabrate, diffusely
pilose. Inflorescences axillary, cymose paniculate, to 9 cm long, ca. 6 cm wide,
the peduncles 4-7 cm long, ca. 0.15 cm wide, stiff, ascending, 1-7 cm long, the
cymules few flowered; bracts and bracteoles linear subulate, to 3 mm long, gla-
brate, the bracteoles of the pedicels several, aggregated at the end of the pedicel.
Flowers not seen. Fruits sessile or pedicellate, the pedicels to 2 mm long, elliptic,
ca. 1 cm long, to 0.6 cm wide, obtuse, the persistent lobes of the calyx 4, erect,
linear subulate or linear ligulate, the 2 shorter 2-3 mm long, the 2 longer 5-6 mm
long, the exocarp carnose, densely appressed pilose.

Chomelia psilocarpa is known only from the Canal Zone.

CANAL zone: barro COLORADO ISLAND! Croat 7931 , 10307 (both MO); Hayden 143 (MO).

8. Chomelia recordii Standley, Trop. Woods 7: 9. 1926. type: Guatemala, Record
8862 (Y, holotype; NY, isotype).

C. en^lcsin^ii Standley, Trop. Woods 16: 45. 1928. lype: Nicaragua, En^lcslng 49 (F, holotype).
Anisonicris recordii (Standley) Standley, N. Am. Fl. 32: 227. 1934.
A. en^lesini^ii (Standley) Standley, N. Am. Fl. 32: 227. 1934.

a. Calyx with lobes lanceolate, villose outside 8a. C. recordii var. recordii

aa. Calyx with the lobes narrowly subulate, glabrous outside

8b. C. recordii var. paucipuhescens

8a. Chomelia recordii Standley var. recordii.

Trees or shrubs, rarely armed, the spines lignose, to 1 cm long, the branchlets
distichous, often short, terete, smooth, drying grey, glabrous or ultimately pu-

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceac) 103

bescent. Leaves oblong, ovate oblong, occasionally oblong rotund or subfalcate,
usually 2 times as long as broad, 6-ll(-12) cm long, 2.5-5.5 cm broad, tapering
obtusely or rounded at the apex, occasionally deltoid, acuminate or only vaguely
so, the acumen to 0.8 cm long, obtuse, basally cuneate to obtuse, occasionally
rounded, the costa plane or prominulous above, the lateral veins 6-8, occasionally
subimmersed above, arcuate, usually forking toward the margin, chartaceous,
scattered villose above, occasionally densely white villose beneath, the hairs
elongate, spreading, occasionally confined to the costa and the veins, the inter-
venal areas glabrous or pubescent, the hairs often tufted in the axils, the margins
often sparsely hirsute; petioles to 0.5 cm long, densely villose; stipules free, ovate
lanceolate, to 8 mm long, acute, densely villose. Inflorescences axillary, of 2-3
flowers on a scarcely measurable and densely bracteate peduncle terminating a
short or rarely elongate branchlet to 8 cm long; bracts foliose, lanceolate, to 5
mm long, acute, venose, villose. Flowers sessile; hypanthium oblong, ca. 1 mm
long, densely appressed white villose, the calycine cup ca. 2.5 mm long, petaloid,
villose outside, glabrous within, the lobes 4, unequal, lanceolate, to 3.5 mm long,
acute, each with a distinct median vein; corolla white, the tube narrow cylindrical,
ca. 17 mm long, 0.5-0,8 mm wide, petaloid, glabrate or densely appressed villose
outside with white hairs, to 1.3 mm long, glabrous within, the lobes 4, oblong,
ca. 5 mm long, obtuse, petaloid, glabrous adaxially, villose to glabrate within, the
margins revolute, short fimbriate; stamens 4, the anthers sessile, narrowly oblong,
ca. 2.8 mm long, attached ca. 2 mm below the mouth; style linear, ca. 17 mm
long, the stigmas 2, narrowly oblong, ca. 1.5 mm long. Fruits oblong, ca. 8 mm
long, ca. 3.5 mm wide, truncate, obtuse at the base, black or purple red, juicy,
the hairs white, to 1.5 mm long, the calycine teeth ca. 5 mm long.

Chomelia recordii is known from Guatemala, Nicaragua, Costa Rica, Panama,
and Colombia. 'Tata de Gallina'' (fide Stern et al. 254).

darien: Between Pinogana and Yaviza, Allen 244 (MO). Rio Ucurganti, Bristan 1 177 (MO). La
Boca de Pirre, Bristan 1281 (MO). El Rio Aruza, Bristan 1365 (MO). El Rio Yape, Bristan 1407
(MO). Rio Uruceca, Bristan 1447 (MO). Without other locahty, Bristan 1509 (MO). Rio Pirre, Duke
4962A (MO). Rio Chucunaque near Quebrada Bolcillo, Duke 8580 (MO). 2 mi N of Santa Fe. Duke
10235, 10715 (both MO). 1-4 mi N of Pucro, Duke 13031 (MO). Quebrada Maskia off Rio Pucro,
above Pucro, Duke 13098 (F, MO). Between Canasas and Sabalo, Duke 14490 (MO). Trail from Paya
to Payita, Stern et al. 254 (MO). Campamento Buena Vista, Rio Chucunaque above Rio Tuquesa,
Stern et al. 865 (MO). 2-3 mi N of Santa Fe, Tyson et al. 4626, 4833, 4843 (all MO). Panama: Bayano
Cuipo forest, 28.5 km E of Bayano Bridge. Folsoni 3538 (MO). Colombia: choco: Roniero-Casta-
neda 6209 (COL).

8b. Chomelia recordii Standley var. paucipubescens Dwyer.-^ type: Panama,

Gentry & Tyson 5038 (MO, holotype).

Shrubs, the branches short, often with reduced spines at the tips of the short
branchlets. Flowers with the calycine lobes narrowly subulate, glabrescent out-
side.

Chomelia recordii var. paucipubescens Dwyer is known only from Panama.

-^Chomelia recordii Standley var. paucipubescens Dwyer, var. nov. Frutices ramulis brevibus
saepe armati spinis brevibus terminalibus. Flores lobis calycis lineari-subulatis glabrescentibus; co-
rolla tubo extus glabrescenti.

104 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

CANAL zonh: Near archeological site, edge of Madden Lake, Gentry A Tyson 5038 (MO).

9. Chomelia spinosa Jacq., Enum. PI. Carib. 12. 1760. type: Colombia, 'Mn syl-
vaticis ad pedem montis de La Popa.'' Jacquin, not seen. — Fig. 22.

Ixora spinosa (Jacq.) Lam., EncycL Meth. Bot. 3: 344. 1789.

Guettarda annatci BartL in DC, Prodr. 4: 457. 1830. type: not seen.

Chomelia filipes Benlh. in Oerst., Vidensk. MeddeL Dansk Naturhist. Foren. Kjobenhavn 1852: 41.

1852.
Caraelina fiUpes (Benth.) Kuntze, Rev. Gen. PI. 277. 1891.
C. spinosa (Jacq.) Kuntze, Rev. Gen. PL 277. 1891.
Guettarda costaricensis K. Schum. nomen 1895. type: not seen.
Anisomeris purpusii Brandegee, Univ. Calif. Publ. Bot. 6: 70. 1914. typf: Mexico, Pringie 6940 (UC,

holotype).

Chomelia hrachyloba Standley, Contr. U.S. Natl. Herb. 18: 136. 1916. type: Panama, Pittier 4673

(US, holotype).
C pnrpasii (Brandegee) Rusby, Bull. Torrey Bot. Club 52: 138. 1925.

C. longicandata Rusby, Bull. Torrey Bot. Club 52: 139. 1925. typk: Colombia, Smith 392 (US).
Anisomeris hrachyloba (Standley) Rusby, Bull. Torrey Bot. Club 52: 142. 1925.

Trees to 20 ft tall or shrubs, the branchlets terete, smooth, the twiglets often
rough with crowded nodes and withered stipular remnants, branched, the branch-
es often uniformly as short as 3 cm, usually strict, the spines axillary, divergent,
lignose, to 2 cm long, to 0.25 cm wide at base. Leaves often crowded at the end
of twigs, oblong, ovate oblong, rarely ovate rotund, 3.7-14.0 cm long, 2.5-5.6
cm wide, acute, rarely obtuse, acuminate or cuspidate, acute at base, obtuse,
often sharply contracted, occasionally truncate, often slightly inequilateral, the
costa prominulous above, prominent beneath, densely or moderately villose be-
neath, the lateral veins 8-10, prominulous above and beneath, arcuate, the in-
tervenal areas not lineolate; petioles to 2 cm long, to 0.5 mm wide, densely pubes-
cent; stipules free, narrowly triangular, 4-6 mm long, tapering into an awn, scari-
ous, pilose, often persistent. Inflorescences axillary, 1 per axil, 2-5 cm long, the pe-
duncle capillaceous, ca. 0.4 cm long, strict, arcuate or occasionally deflexed, the
cymules terminal, few flowered, the flowers erect, sessile, disposed like a can-
delabra. Flowers with the hypanthium rotund, 1.0-1.5 mm long, the calycine tube
ca. 2 times the length of the hypanthium, the teeth 4, unequal, subulate, to 0.8
mm long, sericeous outside; corolla white, the tube cylindrical, to 18 mm long at
maturity, to 0.65 mm wide at the middle, stiffly petaloid, the lobes 4, ovate
lanceolate, to 6 mm long, conspicuously hooded at the narrow apex, sericeous
outside, glabrous within, the margin scarious; stamens 4, the anthers linear ob-
long, ca. 3.5 mm long, obtuse, sagittate at the base, barely exserted, the filaments
slender, very short, attached just below the mouth; ovarian disc about 0.6 mm
long, the stigmas 2, linear oblong, ca. 1 mm long, the style ca. 17 mm long,
slightly exserted, glabrous. Fruits sessile, oblong or ovate oblong, 0.5-0.9 cm
long, obtuse or truncate at the apex, the persistent calyx ca. 1 mm long, drying
black, pubescent, delicately ribbed.

Me

bia, Venezuela and northern Brazil. Lamarck's (1789) discussion of the species
says that, 'These flowers exhale during the night a very sweet odor.'' 'Tata de
Palome.^'

1980]

DWYHR— FLORA OF PANAMA (Family 179. Ruhiineae)

105

Figure 22. Chomelia spinosa Jacq. Habit (x»/2), [After Allen 794.]

CANAL zone: 1 mi NW of Paraiso, near Gaillard Cut, Croat 12656 (MO). Summit Naval Radio
Station, Croat 12852 (MO). Old Gamboa road. Between Summit Hills Golf Course and Summit Naval
Radio, Croat 16668 (MO). Under Thatcher Bridge, D'Arcy & D'Arcy 6082 (MO). Fort Kobbe, Duke
3957 (MO). Albrook, U.S. Army Tropic Test Center, Dnycr 6731 (MO). Cocoli, Dwycr 7212 (MO),
Madden Dam, Boy Scout Road, Dnyer & Bias 7530 (GH, MO, US). Beach, Fort Kobbe, Gentry
6428 (MO). Mandingo Road, Hayden 60 (MO). Ancon Hill, Holdrid^e 6536 (MO). K-2 Road, Con-
tractors Hill, Leuis et al. 2882 (COL, MO, UC, VEN). Fort Kobbe, Luteyn 1088 (F). Gaillard
Highway, Rio Masambi Grande, Nee 7069 (MO). Madden Road, 1-2 km N of Gaillard Highway
Junction, 80-1 10 m. Nee 7614 (MO). 1 mi NE of Fort Clayton, Stimson 5192 (MO). Curundu, Survival
School, Tyson 1059 (MO). Chiva-Chiva Trail, Miraflores Lake, Tyson 1362 (MO). Farfan Beach,
Tyson 1810 (MO). Rodman Ammo Dump, Tyson 6658 (MO). Miraflores Lake, White 128 (F, MO).
Miraflores, White 131 (F, MO). Ancon Hill, Woods(m et al. 1329 (F, MO), cocle: El Valle, Harvey
5188 (F). darien: Chepigana, Duke & Bristan 265 (MO), herrera: Pese, ca. 50 m, Allen 794 (F,
MO). Las Minas and Pese, ca. 600 ft, Duke 12337 (MO). 4 mi S of Los Pozos, Tyson 2652 (MO), los
Santos: Pocri, Dwyer 1195 (MO). Panama: Madden Lake, A^uilera 16 (MO). Pacora, Carrasquilla
190 (MO). Tocumen, Dressier 3898 (MO). E of Bejuco, Duke 4556 (MO). Between El Llano and Rio
Mamoni, Duke 5530 (MO). Tocumen, Dnyer (MO); Dnyer & Hayden 16 (MO). 8 mi S of Goofy
Lake, Dwxer 7125 (MO). 4 mi W of Chepo, Gentry 5555 (MO). Taboga Island, Gentry 5719 (MO).
Archeological Site, Madden Lake, Gentry & Tyson 5038 (MO). Chepo. Pittier4673 (US), veraguas:
Roadside savanna, 2-4 mi E of Santiago, 30 m, Duke 12372 (MO). Santiago, 4 mi from Transisthmian
Highway toward Atalaya, Dnyer & Kirkhride 7411 (GH, MO, UC, US). 5 mi E of Santiago, Tyson
et al. 4274 (MO).

106 ANNALS OF I'HE MISSOURI BOTANICAL GARDEN [Vol. 67

20. CINCHONA

#

Quinquina Boehmer in Ludwig, Def. Gen. PI. 30. 1760. New name for Cinchona L.

Kinkina Adans., Fam. 2: 147. 1763. Based on Cinchona L.

Cascarilla R. ex Steud., Norn., ed. 2: 1: 302. 1840; not Adanson 1763. Nomen nudum.

Trees or shrubs. Leaves petiolate; stipules interpetiolar, free, mostly decid-
uous, glandular at the base. Inflorescences terminal, paniculate, the branches
mostly opposite. Flowers 5-merous, the hypanthium pubescent, the calycine cup
turbinate, the lobes 5, mostly reduced; corolla variable in color, hippocrateriform,
pubescent outside, glabrous within, the lobes valvate; stamens partly exserted or
included, the anthers slender, dorsifixed; ovary 2-celled, the ovules numerous,
imbricate, attached peltately to a spongy placenta, the style slender, the stigmas
2, obtuse; ovarian disc pulviniform. Fruits capsular, mostly ovoid oblong, sep-
ticidally dehiscent, opening from the base; seeds numerous, winged.

Cinchona is found in Costa Rica, Panama and in the Andes of South America.
There are about 40 species in the genus.

1. Cinchona pubescens Vahl, Skr. Naturhist. Selsk. 1: 119. 1790. type: not
seen. — Fig. 23.

C. cordifolia Mutis ex Humb., Ges. Naturfe Feeund, Berlin Mag. Neuesten Entdeck. Gesammten

May 1: Naturk. 117. 1807. XYPt: not seen.
C puhcsccns a cordata DC, Prodr. 4: 353. 1830. typk: not seen.
C tucuycnsis Karst., Fl. Columb. 1: 17. pi. 9. 1858. iype: not seen.

Trees 5-8 m tall, the branchlets angular, glabrous, drying gray or reddish gray.
Leaves oblong rotund to ovate rotund, to 30 cm long, to 20 cm wide, widely
deltoid at the apex, obtuse to subrotund at the base, the costa slender, promi-
nulous above and beneath, the lateral veins slender, 12-13, arcuate, the basal
lateral veins often leaving the costa at right angles, the smaller veins pinnatiform,
stiffly chartaceous, scarcely discolorous, drying red, glabrate above, appressed
pilose on the costa and the lateral veins beneath; petioles to 8 cm long; stipules
not seen. Inflorescences paniculate, to 35 cm long, the branches opposite, angular
ascending, the lowermost to 15 cm long. Flowers not seen. Fruits pedicellate,
the pedicels short, the capsules narrowly oblong or narrowly ovate oblong, to 3.5
cm long, to 0.7 cm wide, minutely golden puberulent, delicately longitudinally
ribbed, the persistent calyx 1-2 mm long, wider than the apex of the capsule.

Cinchona pubescens occurs in Costa Rica, Panama and in the Andes of South
America. The bark is the source of quinine. ''Quina/'

Steyermark (1974) gives an elaborate description of the flowers (from Span-
ish): "Calyx and hypanthium obconic or obconic oblong, 2.5-3.0 mm long, 2.0-
2.5 mm wide, densely pale yellow sericeous. Calyx l-2(-3) mm long, ca. 3 mm

wide, moderately strigulose on the back, the teeth widely and slightly deltoid,
acute to acuminate, much shorter than in C. officinalis, 0.2-LO mm long, strig-
ulose on back. Corolla 15-16 mm long, the tube 10-13 mm long, minutely to-
mentose on the back, glabrous within; lobes lanceolate or oblong, obtuse or
rounded at the apex, 3-5 mm long, 1.5-2.0 mm wide, tomentulose on the back.

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaccae)

107

Figure 23. Cinchona ptihescens Vahl.— A. Habit (xVi). [After Croat /5769.]— B. Dehisced
fruit (X 1).— C. Seed (x5). [After Fosher^ 20016. Colombia.]

108 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

densely villose along the margins within, otherwise glabrous. Anthers 2.5-3.0 mm
long, the filaments ca, 2 mm long, attached approximately at the upper third;
style 6-13 mm long, glabrous; disc scarcely puberulent to hispidulous."

cHiRiQuf: Disturbed cloud forest. Monte Rey above Boquete, Croat 15769 (MO). Santa Clara
region, 27 km NW of El Hato de Volcan, coffee finca Rattibor Hartmen called ''Ojo de Agua,"" 5000-
5300 ft, Mori & Boltcn 7410 (MO).

21. COCCOCYPSELUM

Coccocypselum P. Br., Civ, Nat, Hist. Jamaica. 144. 1756. "Coccocipsilum" No-
men and orthography cons, type: C. repcns Sw.

Siccliion P. Br., Civ. Nat. Hist. Jamaica. 144. 1756. Nomen Rejic. contra Coccocypselum P. Br.

Based on CoccocipsUum P. Br.
Tontanca Aubl., Hist. PI. Guiane 1: 108. 1775. typp: T. i>uianensis Aubl.

Bellardia Schreb., Gen. PI. 1: 790. 1791, not Allioni 1785. Provisional name for Tontanca Aubl.

Condalia R. & P., Prodr. 11. 1794. Nomen Rejic, not Condalia Cav. (1799). Rhamnaceae. Nomen

Cons, type: C. repcns R. & P.
Lipostoma D. Don in Edinburgh Philos. J. 1: 168. 1829. lectotype: L. capltata (Graham) D. Don

{Aeginctia capitata Graham).

Herbs, often woody, prostrate or occasionally erect. Leaves petiolate; stipules
free, persistent. Inflorescences terminal or axillary, the peduncle occasionally
absent, the flowers disposed in a head; bracts and bracteoles small. Flowers
4-merous; calyx with the lobes narrow and elongate, persistent; corolla variable
in color, infundibuliform, the mouth glabrous, the lobes valvate; stamens included
or exserted, the anthers dorsifixed; ovary 2-celled, the ovules numerous in each
locule, horizontal. Fruits baccate, ovoid or globose, shiny blue; seeds numerous,
minute, somewhat angular.

Coccocypselum ranges from Mexico through Central America, the West In-
dies, and South America. There are about 20 species in the genus.

Literature:

Steyermark, J. S. 1967. Coccocypselum in Botany of the Guayana Highland
Mem, N.Y. Bot. Garden 17: 333-334.

a. Leaves with lateral veins 8-12, ovate lanceolate to oblong, to 7.5 cm long, often drying

olive green; pubescence not reddish 4. C. lanceolatum

aa. Leaves with lateral veins 6-8, mostly ovate, rhomboid or ovate rotund, to 6.5 cm long, not
drying olive green, often with a reddish pubescence.
b. Inflorescence with peduncles LO-3.8 cm long; leaves (2-)3-6.5 cm long, often with

a reddish pubescence

c. Leaves not cordate at the base 3. C. hirsntum

cc. Leaves cordate at the base 1. C. cordifoUam

bb. Inflorescence sessile; leaves 0.8-4.2 cm long, lacking a reddish pubescence beneath

2. C. herbaceiim

1.

Mart

German Nat. Cur. 12: 14. 1824.

Geophda plcuropoda Donnell Smith, Bot. Gaz. 52: 50. 1911. typf: Guatemala, Goll s.n., not seen.
Geocardia pleuropoda (Donnell Smith) Standley, Contr. U.S. Natl. Herb. 17. 445. 1914.
Tontanea plcuropoda (Donnell Smith) Standley, N. Am. FI. 32: 148. 1921.

19801 DWYER— FLORA OF PANAMA (Family 179. /?H/7/V;r(w) 109

Cocc(H\pseJiim roihschuhii Loesner, Engler Bot. Jahrb. Syst. 60. 370. 1926. type: not seen.
Cociocypsclum pkiiropoJmn (Donnell Smith) Standley, Publ. Field Colombian Mus., Bot. Ser. 4:

281. 1929.

Herbs, stems prostrate, the hairs dense, weak, spreading, the nodes well
spaced. Leaves reniform, ovate or ovate rotund, 2.0-4.5 cm long, 1.5-3.2 cm
wide, widely deltoid to obtuse at the apex, cordate at the base, the lateral veins

Infl.

cm long.

difoUum ranges from Mexico

eastern and southern Brazil. I have seen only one collection from Panama. Stand-
ley & Williams' (1975) description reads, ''Inflorescence mostly 2-4-flowered;
bracts linear or foliaceous; calyx and hypanthium long-hirsute; calyx lobes linear
or oblong linear, acute, 2-4 mm long; corolla 12 mm long, purplish white or
sometimes blue; fruit densely hirsute, bright blue or sometimes greenish white."

2. Coccocypselum herbaceum P. Br., Civ. Nat. Hist. Jam. 144. 1756. type: Ja-
maica, not seen.

C. repens Swartz, Prodr. Veg. Ind. Occ. 31. 1788. not C. rcpens H.B.K., 1819; not Condaliu repcns

R. & P. (1798).
Tontanea hcrbacea (P. Br.) Standley, N. Am. Fl. 32: 147. 1921.

Prostrate herbs, the stems slender, glabrescent or pilose. Leaves ovate or
ovate subrotund, 0.8-4.2 cm long, 0.5-2.5 cm wide, widely deltoid at the apex,
rounded or truncate at the base, often slightly contracted at the junction with the
petiole, often inequilateral, the costa prominulous above and beneath, the lateral
veins 6-8, arcuate, membranous, glabrescent to pilose above, the hairs short or
long, the latter often scraggly, golden, bulbous at the base, often much longer
above than beneath, pilose beneath but the hairs often restricted to costa and
veins and then appressed and elongate; petioles 0.5-2.5 cm long, ca. 0.1 cm wide;
stipules small, 2.5-3.0 mm long, the awn often 5 times longer than the body.
Inflorescences sessile or subsessile, the heads few flowered. Flowers with the
calycine lobes subulate, to 4 mm long, pilose; corolla light blue, the tube to 9 mm
long, short pilose. Fruits ovoid at maturity, subrotund to ovate rotund when
immature, to 1.5 cm long, pilose, the hairs gradually deciduous, china blue.

Coccocypselum herbaceum is well distributed throughout tropical America
and the West Indies. Steyermark (1967) discusses the synonomy of C. herbaceum
at length.

BOCAS DEL TORo: Santa Catalina, Bhicknell ct al. 2730 (MO, PMA). Chiriqui Lagoon, Wedcl
1134 (MO). CANAL zone: N of El Cope, 9.4 km above El Cope, 750-900 m, Croat 44632 (MO).
Pipeline Road, D'Arcx 9273 (MO). Chagres, Fendlcr 179 (MO). Telephone cable trail between splice
816 and S49, Rio Indio. Steyermark & Allen 17433 (MO), chiriqui': Cerro Colorado, 20-28 mi from
San Felix, 1200-1500 m, Croat 33374 (MO), cocle: La Mesa above EI Valle, Dwyer 11904 (MO). La
Mesa, Folsom & Butcher 2837 (MO). El Valle, Lenis et al. 2512 (COL, DUKE, K, MO, UC, VEN).
colon: Miguel de la Borda. Croat 10028 (MO). Rio Guanche near Coastal Highway, D'Arey 10192
(MO). Santa Rita Ridge, Dwxer & Gentry 9360 (MO), herrera: 12.5 mi S of Ocu, 1200 ft. Lewis et
al. 1647 (MO). Panama: Alto de Pacora, Bnsey 804 (MO). Cerro Jefe, Croat 13059 (MO). Campo
Tres beyond Cerro Jefe, Croat 27088 (MO). Pipeline Road near Gamboa, Clewell & Tyson 3218 (MO).
Panama Viejo, Duke 5734 (MO). Cerro Azul, Duke 9378 (MO). Cerro Campana, Gentry 1853 (MO).
Cerro Jefe, summit, ca. 3000 ft, Ha\den 1004 (MO). Cerro Azul, Mori & Kallunki 2167 (MO). Cerro

1 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

Jefe, ca. 1000 m, Mori & Kallunki 3618 (MO). Cerro Jefe, Tyson ef ciL 3215 (MO), 4430 (SCZ). san
BLAs: Road between Mandinga & Cangandf, Duke 14745 (MO), vhraguas: Rfo Primero Braso, 2.5
km beyond Agriculture School near Santa Fe, 700-750 m, Croat 25458 (MO). To 7 km NW of Santa
Fe, D'Any 10279 (MO). Mouth of Rio Concepcion, Lewis et al. 2803 (MO, SCZ, UC).

3. Coccocypselum hirsutum Bartling ex DC, Prodr. 4: 396. 1830. type: probably
Trinidad, type collection not known.

Tontanea hirsuta (Bartl.) Standley, N. Am. Fl. 32: 147. 1921.
Coccocypselum hirsutum Bartling ex DC. var. hirsutum.

3a. Coccocypselum hirsutum DC. var. hirsutum.

Prostrate herbs, the nodes well spaced. Leaves ovate, ovate rhomboid, rarely
oblong or reniform ovate, (2-)3.0-6.5 cm long, 1.5-5.0 cm wide, widely deltoid
to obtuse at the apex, widely triangular to truncate at the base, usually slightly
inequilateral, densely pubescent or with a few elongate hairs above, these golden
and bulbous at the base, hirsute beneath, the hairs often elongate and stiff as well
as short and soft, the hairs sometimes restricted to the margins of the blade;
petioles to 2 cm long; stipules subulate, 3-6 mm long (Standley & Williams 1975).
Inflorescences pedunculate, the peduncles 1-4 cm long, slender, glabrescent,
often bearded. Flowers with the calycine lobes 4, linear lanceolate, 3-6 mm long,
densely hirsute; corolla to 11 mm long. Fruits 2-4 per cluster, ovate, to 2 cm
long, densely hairy or with a few hairs, often delicately lenticellate.

Coccocypselum hirsutum ranges from Mexico to Panama. It also occurs in
Trinidad, Colombia, and Bolivia.

canal zone: Cerro Pelado, 1 km N of Gamboa, 200-220 m. Nee 1 1024 (MO), chiriqui': San
Felix, Cerro Otoe, 3000 ft, Bort 69 (MO). Road from Boquete to David, KirkhriJe 85 (MO), colon:
Rio Piedras, road to Portobelo, Blum et at. 2518 (MO, SCZ). Maria Chiquita, Dnyer S: Kirkhride
7796 (MO). Santa Rita Ridge, Nee & Mori 3671 (MO). NE of Santa Rita Ridge, 650 m, Wilbur &
Weaver 10837 (MO). Panama: Goofy Lake, 200 ft, Correa & Dressier 583. 584 (both MO). Cerro
Jefe, D'Arcx 975! (MO). Cerro Azul, D'Arcy tQ D'Arcy 6227 (MO). Hills NE Hacienda La Joya,
Dod^e et al. 16896 (MO). Cerro Campana, summit, Duke 6009 (MO). Cerro Azul, Ehin^er42I (MO).
Cerro Jefe, 2900 ft, Elias & Harden 1796 (MO). Cerro Azul, Finca Melo, Kant 47 (MO). Cerro Jefe,
Kirkhride A Crehhs 28 (MO). Cerro Campana, Kirkhride 245 (MO): Le\yis et al. 1927 (GH, MO, US),
1948 (MO), 3075 (COL, MO, UC, YEN); Porter et al 4175 (MO). Cerro Jefe, 2900 ft. Porter et al.
5063 (COL, MO, UC). Cerro Campana, Witherspoon ct Witherspoon 8904 (MO).

3b. Coccocypselum hirsutum var. glabrum (DC.) L. O. Williams, Phytologia 25:
462. 1973.— FiG. 24.

C. glabrum Bartl. ex DC, Prodr. 4: 397. 1830, rvPL: not seen.
Tontanea i^lahra (DC.) Standley, J. Wash. Acad. Sci. 15: 104, 1925.

Prostrate herbs. Leaves ovate to oblong, 3-6 cm long, to 5 cm wide, widely
triangular to truncate at the base, glabrate above and below. Inflorescences with
the peduncles to 4 cm long, glabrous. Fruits ovate, to 2 cm long.

Coccocypselum hirsutum var. glabrum occurs in Mexico, Guatemala (Peten),
Belize, Nicaragua, and Panama.

CHiRK^uf: Boquete, ca. 4000 ft, Davidson 759 (MO). 4.1 mi from Boquete, road to David, Kirk-
hride 85 (MO), colon: Between Rio Piedras and Puerto Pilon, Lewis et al. 3227 (MO). Santa Rita

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaccac)

111

Figure 24. Coccocypselum hirsutum var. ghihrum (DC.) L. O. Williams. Habit (xi/2). [After
Kirkhride & Crehbs 28.\

Ridge, 2 mi E of Transisthmian Highway, Le^^•^s et al. 5242 (MO, SCZ, UC). Santa Rita Ridge Road,
ca. 1.9 km from Boyd-Roosevelt Highway, Mori & KaUunki 1808 (MO). Panama: Goofy Lake,
Correa & Dressier 584 (MO). Cerro Campana, savanna near top, Duke 6009 (MO). Goofy Lake, 2000
ft, Dwyer et al. 4137 (COL, DUKE, F, MEXU, MO, NY, MICH, PMA, VEN). Cerro Campana, ^4
way to summit, D\vyer47l9 (MO). Cerro Azul, Montenegro 65 (MO).

4. Coccocypselum lanceolatum (Ruiz & Pavon), Pers. Syn. PI. 1: 132. 1805.

ConJalia lanceohita Ruiz & Pavon, Fl. Peru. ChiL Prodr. I: 54. 1798. type: Peru, '"m memoribus

Cuchero and Chinchao, Ruiz & Pavon,'' not seen.
Coccoeypsehtm caneseens Willd. ex C. & S., Linnaea 4: 139. 1929. typf: "*Herb. Willd. n. 2844

(specimen Humboldtianum),'' not seen.
Tontanea caneseens (Cham. & Schlect.) Slandley, N. Amer. Fl. 32: 146. 1921.

Herbs, prostrate or to 25 cm tall, the stems golden green, pilose, the nodes
well spaced. Leaves ovate lanceolate to oblong, often somewhat falcate, 2.0-7.5
cm long, 1-3 cm wide, deltoid, obtuse to occasionally rounded at the apex, basally

1 12 ANNALS OF THE MISSOURI BOTANlCAl GARDEN [Vol. 67

cuneate or obtuse, occasionally truncate, acute or obtuse, usually slightly in-
equilateral, the costa plane or grooved above, plane to prominulous beneath, the
lateral veins 8-12, arcuate, chartaceous, densely pilose beneath and above, vel-
vety to the touch, often drying olive green, the veins often bearded beneath;
petioles 0.3-l.3(-3.5) cm long; stipules mostly persistent, ovate triangular, 5-6 mm
long, the awn longer than the body, densely hairy. Inflorescences subsessile or
on peduncles 2-3(-5) cm long, 0.1-0.3 cm wide, densely pilose. Flowers sessile,
numerous; hypanthium subrotund, ca. 2 mm long, densely hairy, the calycine
lobes 4, erect, ovate or oblong, 3(-6) mm long; corolla white, rarely purple, the
tube 4-5 mm long, petaloid, pilose outside, glabrous within, the lobes 4, oblong
or triangular, ca. 2 mm long; stamens 4, the anthers sessile, attached near the
middle of the tube, ovate, ca. 1 mm long; style ca. 2 mm long, the stigmas slender,
about the same length as the style. Fruits bright blue, ovoid, to 7 mm in diam.,
pilose.

Coccocypselum lanceolatum is well distributed throughout Middle America
and extends into Brazil and Bolivia. The leaves of the Panamanian collections
are smaller than those collected to the west of Panama.

CHiRiQuf: Boquete, 3800 ft, Davidson 635, 756 (both F, MO). Boquete, Llanos Francia, Dwyer
& Hayden 7583 (F, GH, MO), 7602 (MO). Boquete, Road to David, Kirkhride 84 (MO). Cerro
Colorado, road 35.6 km from Rio San Felix bridge, 1390 m, Sullivan 379 (MO). Boquete, 1200-1500
m, Woodson & Schcry 793 (MO), cocle: Between Las Margaritas and El Valle, Woodson ct al. 1275
(MO), darii'n: Gold mine, Cana, 500-600 m, Croat 37646 (MO), hkrrfra: Between Las Minas and

Pese, Biirch ct al. 1338 (MO), vhraguas: Trail between Canazas and foot of Cordillera Central, Ri'o
Canazas, Allen 183 (MO). Near San Jose on Santiago-Santa Fe Road, 400 m, Ncc 8179 (MO).

22. COFFEA

Coffea L., Sp. PI. 172. 1753; Gen. PI. ed. 5: 80. 1754. type: C. arabica L,

Shrubs, rarely pubescent. Leaves occasionally ternate, usually petiolate, the
stipules free, acuminate, persistent. Inflorescences with flowers conglomerate in
the axils, the bracteoles connate into a calyxlike cup. Flowers sessile or pedi-
cellate, the hypanthium variously shaped; calycine cup reduced, truncate, dentate
or lobulate, often bearing small glands within; corolla salverform or funnelform,
the lobes 4-8, contorted; stamens 4-8, sessile or on short filaments inserted in
the throat, the anthers linear, basally dorsifixed, exserted or included, often twist-
ed; ovarian disc swollen, the style slender or thick, the stigmas 2, narrow, the
ovary 2-celIed, the ovules solitary, subpeltately affixed to the middle of the sep-
tum. Fruits baccate, dry or fleshy, the 2 carpels often coriaceous, convex but
plane and sulcate on the inner face; seeds 2, convex, the testa membranous with
a deep groove on the ventral side.

This is a genus of about 40 species represented in the tropics of the Old World,
especially in Africa. The 2 species, Coffea arabica and C. liberica, supply most
of the coffee of commerce.

Literature:

Standley, P. C. & L. O. Williams. 1975. Coffea Linnaeus Coffee (part of Ru-
biaceae of Guatemala). Fieldiana 24, part IL 44.

1980]

DWYER— FLORA OP PANAMA (Family 179. Riihiaceae) 1 13

a.
aa.

Flowers 5-merous 1- C. arahica

Flowers 6-8-merous 2. C. liberka

1. Coffea arabica L., Sp. PI. 172. 1753. type: not seen.— Fig. 25.

Shrubs or trees, to 20 m tall, the branchlets straight or flexuous, glabrous,
scarcely or moderately swollen at the nodes, the latter well spaced. Leaves el-
liptic oblong, 8-18 cm long, 2.5-7.5 cm wide, distinctly acuminate at the apex,
the acumen to 1.5 cm long, often falcate or somewhat hooklike, usually ultimately
obtuse, attenuate or widely cuneate at the base, the costa prominulous above,
subprominent beneath, the lateral veins 9-11, uniting to form a submarginal un-
dulate vein to 0.8 cm from the margin, thin coriaceous to stiffly papyraceous,
usually drying green or grey green; petioles to 1.2 cm long; stipules linear lan-
ceolate, to 1.2 cm long, acute. Flowers not seen. Fruits plump, oblong rotund,
to 1.5 cm long, to 1 cm wide, rounded at the apex and the base, drying black
brown or dull olive, glabrous, the calycine scar scarcely elevated, ringlike, to 2
mm in diam.

Coffea arabica is native to Africa and is cultivated throughout the tropical
areas of the world. Standley & Williams (1975) discuss the cultivation of Coffea
arabica in Guatemala. The 5-parted flowers are described in their diagnosis:

flowers in clusters of 2-9 or more; sessile or nearly so, 12-18 mm long,
bractlets ovate, the inner ones connate at the base of the pedicel, shorter than
the 5-denticulate calyx; corolla lobes equalling or exceeding the tube; anthers

exserted . . . ."

CHiRiQuf: Cafetales above El Cantinero, DArcy 9846 (MO). Finca "Princesa Janca," Boquete,
D'Arcy & D'Arcy 6453 (MO). Fred Collins Finca, Boquete, Ehinger 723 (MO). Quebrada Melliza,
6 mi S of Puerto Armuelles, 0-150 m, Liesner 457 (MO), cocle: Forest behind Club Campcstre, 700
m Duke 13252 (MO). El Valle, Ehinger 948 (MO). Panama: Rio Maje along river from waterfalls
near Bayano Lake ca. 2 mi upstream, Croal 34553 (MO). Highway 79, Quebrada Sardinilla, 5 km
NNE of cement factory, 55 m, Nee 6581 (MO). Cermeiio, Ochoa 32 (MO).

2. Coffea liberica Bull ex Hiern., Trans. Linn. Soc. Ser. 2(1): 171, tab. 24. 1876.
type: Sierra Leon, Daniel! (BM, not seen; photo, MO).

Shrubs or trees to 20 ft tall, the twiglets subangular, glabrous, the nodes well
spaced. Leaves oblong or obovate elliptic, 7.5-8.5 cm wide, short acuminate at
the apex, rounded at the base, the lateral veins ca. 12, coriaceous, glabrous;
petioles to 2 cm long, stiff. Flowers not seen. Fruits oblong subrotund, to 2.0-
2.5 cm long at maturity (fide Standley & Steyermark), lightly grooved.

Coffea liberica is native to Liberia and to areas of West Africa. It is cultivated
throughout tropical areas of the world. Standley & Williams (1975) describe the
6-8-parted flowers as ". . . several in a cluster, subsessile, 2.5 cm long; bractlets
shorter than the subtruncate calyx; corolla lobes about as long as the tube; anthers
not exserted." "Liberian Coffee."

colon: Achiote, Tyson et al. 4508 (MO).

23. CONDAMINEA

Condaminea DC, Prodr. 4: 402, 1830. lectotype: C. corymbosa (R. & P.) DC

Shrubs or small trees. Leaves large, coriaceous; stipules large, bipartite. In

florescences terminal, spreading, cymose or corymbose; bracteoles absent. Flow

114

ANNALS OF THE MISSOURI BOTANICAL GARDEN

(Vol. 67

FiGURH 25. Coffca arable a L. Habit {xVi). | After Odioa 32.]

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceac) 1 15

ers pedicellate; hypanthium turbinate campanulate, the calycine cup with 3-5
lobes or teeth; corolla funnelform or campanulate, the tube cylindrical, villous in
the throat, the lobes 5, valvate, glabrous; stamens 5, the anthers exserted, narrow,
sagittate, the filaments thick, inserted below the corolla mouth; ovarian disc de-
pressed, the ovary 2-celled, the style stout, filiform, exserted, the stigma oblong,
obtuse, the ovules numerous. Fruits capsular, turbinate, truncate, thick walled,
dehiscing loculicidally into 2 valves; seeds minute, attached horizontally, the
testa reticulate.

Condaminea occurs sporadically throughout Middle America and occurs in
tropical South America. There are about 3 species in the genus,

a. Leaves oblong, to 50 cm long, the lateral veins ca, 20: petioles absent or to 1 cm long —

1. C. corymbosa

aa. Leaves lanceolate, to 15 cm long; the lateral veins ca. 10; petioles 0.7-1.2 cm long —

2. C. petiolata

1. Condaminea corymbosa (Ruiz & Pavon) DC, Prodr. 4: 402. 1830.— Fig. 26.

Macrocnemum corymhosum Ruiz & Pavon, Fl. Peruv. Chil. Prodr. 2: 48. 1799. type: not seen.

Shrubs or small trees to 7 m tall, the branches terete, stout, smooth, glabrous
or puberulent. Leaves oblong, to 50 cm long, to 21 cm wide, deltoid to vaguely
obtuse at the apex, basally truncate to subcordate, the costa plane or prominulous
above, prominent beneath, to 3 mm wide, the lateral veins usually ca. 20, usually
1.5-2.5 cm apart, coriaceous, glabrous to golden puberulent beneath, the inter-
venal areas often delicately wrinkled; petioles either absent or to 1 cm long, stout,
rimose, glabrous; stipules interpetiolar, connate, to 4 or 5 cm long, to 0.6 cm
wide at the base forming a sheath, each part with 2 subulate awns longer than
the sheath, acute, coriaceous, puberulent, rubescent. Inflorescences terminal,
spreading paniculate, to 40 cm long, the peduncle 15-20 cm long, with 3 primary
branches at the tip, 10-15(-21) cm long, angular ascending or arcuate, each ter-
minating in either 3 radially disposed secondary branches or in a few branches.
Flowers with the hypanthium oblong, to 1 cm long, to 0.6 cm wide, tapering
cuneately below, the calycine cup short, dilated, the margin with 4 or 5 broad,
strongly compressed rotund or widely triangular lobes, coriaceous, glabrous; co-
rolla white, the tube cylindrical, ca. 2x longer than wide, to 2 cm long, carnose,
glabrous inside and out, the lobes 5, triangular or oblong, to 1 cm long, glabrous;
stamens 5, the anthers narrowly oblong, ca. 7 mm long, ca. 1.2 mm wide, retuse
at the apex, the filaments short, plane, attached at the upper % of the tube; style
plane, attenuate at the base, to 1.5 cm long; stigmas 2, to 5 mm long, ca. 0.7 mm
wide, truncate, the ovary thin walled, the locules 2-3, the ovules numerous on
2-3 intrusive, C-shaped placentas as seen in cross section. Fruits^ oblong or
obovate oblong, 1-1.5 cm long, ca. 1 cm wide at the base, ligneous, glabrous,
drying black, topped by a conspicuous and shallow ringlike scar, splitting longi-
tudinally into 2 or 3 thin valves; seeds numerous.

L

Condaminea corymbosa ranges from Costa Rica to Bolivia.

CHiRiQUi: Boquete, Davidson 792 (F, MO, US). 12.4 mi N of David, Lewis et al. 692 (GH, K,
MO, NY, PMA, UC, US). 17 km NE of San Felix, road to Cerro Colorado, 1000 m, Nee 10718 (MO).
San Felix, Pitticr 5252 (F, US), darien: Piiias, Duke 10621 (MO).

116

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 67

FiGURK 26. Condaminea corymhosa (Ruiz & Pavon) DC. — A. Leaves and inflorescence
(x»/2)._B. Fruit (x3.5).— C. Seed (x21). [After Duke 10621.]

■

I980I DWVf-R— FLORA OF PANAMA (Family 179. Ruhiaceae) 117

2. Condaminea petiolata Dwyer.^^ type: Panama, Croat 34706 (MO, holotype).

Shrubs to 2 m, the twigs slender, glabrous, obtusely angled, the nodes well
spaced. Leaves lanceolate, to 15 cm long, to 4.5 cm wide, acute at the apex and
the base, the costa prominulous above and beneath, the lateral veins ca. 10,
strongly arcuate, plane above, prominent beneath, the intervenal areas open re-
ticulate, chartaceous, discolorous, the hairs appressed, white, elongate, more
dense beneath especially on the costa and the veins; petioles 0.7-1.2 cm long,
ca. 0.2 cm wide, pubescent; stipules persistent at the apex of the twigs, connate
basally, turgid, the sheath 1-2 mm long, each part with an erect triangular pro-
cess, attenuate toward the apex, to 0.6 cm long, acute, vaguely thickly carinate,
villose outside, densely villose at the base within and often with a few conspic-
uous black glands on the margin within, the glands oblong, ca. 0.5 mm long.
Inflorescences axillary, paniculate; peduncle slender, to 0.3 cm wide, the cor-
ymbs disposed toward the apex of 2-3 pairs of short, arcuate, ascending, opposite
branches, these 3-5 mm long; bracts not seen; bracteoles subulate. Flowers not
seen. Fruits sessile, oblong, coriaceous, to 1 cm long, less than 2 times longer
than wide, truncate at the apex, pubescent outside, delicately ribbed, the persis-

rpel

ocarp

finally free from mesocarp; seeds numerous, piano compressed, fusiform, ca. 2
mm long, shiny.

Condaminea petiolata is known only from Panama. It is distinguished by its
leaves having few lateral veins and by its axillary inflorescence. The fruits are
smaller than those of C corymbosa.

PANAMA: Rio Maje, near new Bayano Lake, 100 m, Croat 34706 (MO).

24, COSMIBUENA

Cosmibuena Ruiz & Pavon, Fl. Per. Chil. Prodr. 3: 2. 1802. Nomen conservan-
dum. type: C. obtusifolia Ruiz & Pavon not Cosmibuena Ruiz & Pavon
(1794). Nomen rejiciendum (Chrysobalanaceae). Buena Pohl, PL Bras. Ic. 1:

8. 1827.

Shrubs, glabrous, usually epiphytic. Leaves fleshy, petiolate; stipules large,
deciduous. Inflorescences terminal, cymose or the flowers solitary; bracts and
bracteoles present. Flowers large; hypanthium oblong or turbinate, the calycine
cup tubular or campanulate, the teeth 5-6, usually unequal; corolla salverform
or funnelform, the tube elongate, 5-6-lobed, the lobes contorted; stamens 5-6,
the anthers linear, basifixed, included, the filaments short: ovarian disc cup

^*' Condaminea petiolata Dwyer, spec. nov. Frutices ad 2 m alii, ramulis gracilibus glabris obtuse
angulatis nodis bene distantibus. Folia lanceolata, ad 15 cm longa, ad 4.5 cm lata, acuta, venis
lateralibus ca. 10, forte arcuatis, chartacea discoloria; petiolis ad 1.2 cm longis; stipulis connalis
turgidis vagina 1-2 mm longa parte utraque processu triangulari erecto ad 0.6 cm longo praedita extus
crasso-carinata villosaque. Inflorescentiae axillares corymboso-paniculatae pedunculo gracile ad 0.3
cm lato, corymbis versus apicem jugorum 2-3 ramulorum brevium arcuate ascendentium opposito-
rum, ad 3-5 mm longorum. Fructus sessiles oblongi, ad 1 cm longi, coriacei apice truncati extus
pubescentes modice costati.

1 18 ANNALS OF THK MISSOURI BOTANICAL GARDEN [Vol. 67

shaped, the style elongate, clavate at the apex, the stigmas 2. Fruits capsular,
narrowly oblong or cylindrical, septicidally bivalvate from the apex, the epicarp
separating from the endocarp; seeds numerous, oblong, the body small, the testa
membranous, produced at each end into a narrow wing.

a. Calyx cup ca. LI cm long, the teeth only minute points; corolla tube to 7.5 cm long

aa. Calyx cup 3.3-5 mm long, the teeth L5-3 mm long; corolla tube 6-10 cm long

1 . C. oralis

2. C. skinnen

1. Cosmibuena ovalis Standley, Contr. U.S. Natl, Herb. 18: 137. 1916. type:
Panama, Pittier 5074 (US, holotype; NY, isotype).

Trees to 30 ft tall, 4 inches d.b.h., terete, drying dark, the nodes well spaced.
Leaves rhomboid, oblong rotund or elliptic oblong, 12-17 cm long, 5.5-8.5 cm
wide, deltoid, rounded or obtuse at the apex, acuminate, the acumen vague,
short, basally cuneate or attenuate cuneate, the costa plane or prominulous
above, prominulous beneath, to 0.2 cm wide, the lateral veins 5-6, at first strict
and strongly ascending, arcuate toward the margin, the submarginal vein indis-
tinct, stiffly chartaceous, discolorous, glabrous above except the puberulent cos-
ta, glabrous beneath except puberulent along the lowermost part of the costa and
along the lateral veins; petioles to 1 cm long, to 0.15 cm wide in the middle,
puberulent, expanded at point of attachment; stipules not seen. Flowers 2 at the
end of the twiglet, erect, the calyx collar shaped, to 1.1 cm long, to 0.6 cm wide,
truncate, the teeth minute points to 0.5 mm long, acute, rigid, glabrous; corolla
tube narrowly cylindrical, to 7.5 cm long, to 0.3 cm wide in the middle, glabrous,
scarcely expanded at the apex, the lobes oblong, 2-2.5 cm long, 1-1.3 cm wide,
obtuse, glabrous; style exserted, probably to 8.5 cm long, densely golden villose.
Fruits not seen.

Cosmihuena ovalis is known only from Costa Rica and Panama. The above
description is sketchy, as only one collection was available except for the type.

CHiRiQuf: Rio Macho de Monte, 1 mi E of Cuesta de Piedra, 2800 ft, Tyson 904 (MO), cocle:
100-350 m, Ola, Pltticr 5074 (NY, US).

2. Cosmibuena skinneri (Oerst.) Hemsl., Biol. Centr. Amer., Bot. 2: 12. 1881.

Bucna skinneri Oerst. Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 26: 1852. type: Nic-
aragua, Oersted 11134 (not seen, holotype, photo, MO).

Cosmibuena paludicoUi Standley, Contr. U.S. Natl. Herb. 18: 137. 1916. type: Panama, Pittier 4107
(US, holotype; NY, isotype).

Trees to 25 ft tall, the branchlets terete, smooth, glabrous, often drying gray
or tan, frequently angular, the nodes usually well spaced. Leaves obovate oblong,
occasionally oblong subrotund, 6-16 cm long, 2.7-7.5 cm wide, obtuse or rounded
at the apex, cuneate at the base, the costa prominulous above and beneath, often
evanescent distally, 1-1.5 mm wide at the base, the lateral veins 5-7, prominu-
lous, evanescent, strongly ascending, coriaceous, glabrous; petioles to 2 cm long,
to 0.25 cm wide, lignose, glabrous; stipules deciduous, oblong rotund, ca. 1.5 cm
long, 1 cm wide, obtuse, coriaceous, glabrous, often with a thickened keel below
the middle. Inflorescences terminal, the peduncle short, to 1 cm stout, glabrous.

1980]

DWYHR— FLORA OF PANAMA (Family 179. Ruhiaceae)

119

Figure 27. Cosmihuena skinned (Oerst.) Hemsl. — A. Flowering branch {xVi). [After Croat
6i26.]— B. Fruiting branch {xVi). [After Croat 5057,]— C. Flower (x 1).—D. Seed (x2). [After Gentry

576L\

drying much darker than the stem tip, the flowers 3-6, ascending. Flowers 1-2
cm long (pedicels and hypanthium), ca. 0.4 cm wide, glabrous, marcescent; ca-
lycine cup 3.3-5 mm long, glabrous, coriaceous, the teeth 5, widely triangular,
disposed as points 1.5-3 mm long, acute or obtuse; corolla white, the tube nar-

120 ANNALS OF THF MISSOURI BOTANICAL GARDEN [Vol. 67

rowly cylindrical, 6-10 cm long, ca. 0.3 mm wide, drying brown, subcoriaceous,
glabrous, the lobes 5, oblong, 1.9-3.5 cm long, 0.9-1.2 cm wide, obtuse, often
glabrous, often drying yellow abaxially, brown adaxially; stamens 5, the anthers
narrowly oblong, 1.0-1.2 cm long, ca. 0.15 cm wide, the filaments short, inserted
near the middle of the corolla tube, anthers basifixed; stigmas 2, digitiform or
oblong, to 1 cm long, 1.8 cm wide, obtuse, the style slender, as long as the corolla
tube, ca. 1 mm wide, villose for ca. 6 mm below the stigmatic lobes, the latter
oblong, ca. 5 mm long, obtuse. Fruits pedicellate for ca. 2 cm, narrowly oblong,
often falcate, to 7.5 cm long, to 0.8 cm wide, terete in cross section, rounded at
the apex with an annular scar, the calyx rarely persistent, to 0.3 cm long, the
ovarian disc persistent as a cylindrical projection, to 2 mm long, ca. 1 .3 mm wide,
attenuate at the base, ligneous, glabrous, conspicuously lenticellate.

Cosmibuena skinneri ranges from Mexico to Colombia. Croat has noted on
labels that the flowers are pollinated by hawkmoths.

Panama: canal zone: Gatiin, Hi^gins 400 (MO), barro Colorado island: AvUes 78 (MO):
Ban^ham 473 (F); Croat 5057, 6326, 6554, 7833, 8519, 9029, 9853, 12285, 12444 (all MO); Foster 1912

(F); Kmf,^ht 8 (F); Shattuck 863 (MO); WooJnorth & Vestal 771 (MO), cocle: EI Valle, Allen 2798
(MO). 2 km from El Greco Hotel, El Valle, Correa & Dressier 997 (MO). Las Margaritas & El Valle,
Woodson et al. 1738 (MO), colon: Santa Rita Ridge, Correa & Dressier 695 (MO); DArcv & D'Arcy
6162 (MO). Maria Chiquita, E of Rio Piedras, Duver & Kirkhride 7773 (GH. F, K, MO, NY. PMA,
UC). Santa Rita Ridge, Dwyer & Gentry 9394 (MO). Vienta Frio, Pittier 4107 (US), darien: Rio
Pinas, Duke 10546 (MO). Panama: Cerro Campana, Correa & Dressier 1032 (MO); Duke 6008 (MO);
Dwyer & Kirkhride 7829 (MO); Ehinger 917 (MO); Gentry 5761 (MO); McDaniel 6921 (MO), san
blas: Puerto Obaldia, Gentry 1496 (MO), without exact locality: Duke 6193 (MO). Colombia:
choco: Between Camp Curiche and Quebrada Changame, 3.7 mi S of Camp Curiche, Duke <5 Idrobo
11574 (NY).

25. COUSSAREA

Coussarea Aubl., Hist PI. Guiane 1: 98. 1775. type; C vioJacca Aubl.

Trees or shrubs. Leaves often coriaceous; stipules usually entire and connate,
short, obtuse, often semiorbicular. Inflorescences terminal or rarely axillary, pa-
niculate or umbellate, many flowered, the peduncle usually present, the branch-
lets ultimately cymose; bracts slender or much reduced. Flowers usually 4-mer-
ous, rarely sessile, the bud rounded or truncate; hypanthium occasionally with
4 lobes or teeth; corolla funnelform or salverform, the throat naked, the lobes 4,
valvate; anthers subsessile; stigmas 2, the lobes filamentous, the ovary 2-celled,
the ovules 2, longer than broad, erect, connate by means of a thin, evanescent
septum. Fruits drupaceous, oval, 1-seeded by abortion of the second seed, the
seed vertical, ellipsoid, usually smooth, often one side convex.

Coussarea occurs throughout tropical America. It includes about 100 species.

One species of Viburnum, V. costaricanum (Oerst.) Hemsley, well collected
in Chiriqui, may be easily mistaken in the fruiting stage for a Coussarea, but the
fruit may be distinguished from Coussarea species by its persistent ovarian disc
which is triangular but conspicuously capitate at the apex, measuring ca. 0.4 mm
long.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhicueae) 121

Literature:

Dwyer, J. D. and M. Hayden. 1966. Notes on Coussarea (Rubiaceae) especially
the Panamanian Species. Ann. Missouri Bot, Gard. 53: 368-374.

a. Leaves coriaceous, inflorescence 3-5 cm wide.

b. Leaves with lateral veins 16-25; petioles less than 1 cm long; stipules 0.25-0.5 cm

long.

c. Leaves petiolate. the lateral veins ca. 16; calycine cup ca. 3 mm long

cc. Leaves sessile, the lateral veins ca. 25; calycine cup ca. 9 mm long

3. C. Jurifolia

5. C. garciae

bb. Leaves with lateral veins ca. 9; petioles 1-3 cm long; stipule 1-1.5 cm long.

d. Leaves 9.0-16.5 cm wide; fruits to 2 cm long 7. C, latifolia

dd. Leaves 1-5 cm wide; fruits to 1 cm long 6. Cjefcnsls

aa. Leaves chartaceous or if coriaceous (C paniculata) the inflorescence to 6 cm wide.

e. Leaves obviously petiolate.

f. Petioles to 0.5 cm long - 2. C. curvigemmia

ff. Petioles 0.6-5.0 cm long.

g. Calycine cup ca. 16 mm long; corolla tube ca. 33 mm long

4. C. cnncantha

Calycine cup 0.2-7.0 mm long; corolla tube less than 20 mm long,
h. Inflorescence patulous, open.

i. Leaves with lateral veins 11-15.

j. Calycine cup ca. 0.5 mm long -_ 14. C. vcragucnsis

ij. Calycine cup ca. 2.5 mm long 8. C. loftonii

ii. Leaves with lateral veins 5-10(-12).

k. Leaves with lateral veins 5-6 10. C. nehidosa

kk. Leaves with lateral veins 7-IO(-12).

I. Calycine cup ca. 16 mm long 4. C. cnncantha

II. Calycine cup 0.5-7.0 mm long,
m. Leaves with lateral veins ca. 12; petioles 2-5

cm long 1. C ccrroazulcnsis

mm. Leaves with lateral veins 8-10; petioles 0-1.5

cm long.

n. Calycine cup ca. 5 mm long; flowers

pink 13. C. rosco-cremea

nn. Calycine cup 1-2 mm long; flowers white.

o. Petioles (longest) 2.0-2.5 cm long;

fruits to 2.7 cm long 1 1. C. ncci

oo. Petioles (longest) ca. 1.5 cm long;

fruits 1.2-1,6 cm long

12. C. paniculata
p. Leaves coriaceous; calyx pu-

berulent, edentate.
pp. Leaves thinly chartaceous; ca-
lyx glabrous, often with teeth

__ 9. C. morii

hh. Inflorescence obviously contracted 15. C. viUosnIa

Leaves sessile - - 5. C. garciae

L Coussarea cerroazulensis Dwyer & Hayden, Ann. Missouri Bot, Gard. 53:
374. 1966. type: Panama, Dwyer 1368 (MO, holotype).

Trees 5-10 m tall, the branchlets terete, densely villosulose, the nodes well
spaced, often turgid with evident stipule bases. Leaves elliptic, 15-33 cm long,
5-14 cm wide, acute or conspicuously acuminate, the acumen slender, often
narrowly caudate, to 2 cm long, basally cuneate, the costa prominulous above.

122 ANNALS OF THE MISSOURI BOTANICAL GARDEN |Vol. 67

prominent beneath, the lateral veins ca. 12, widely arcuate, prominent beneath,
densely golden pubescent, often velutinous to the touch, coriaceous; petioles 2-
5 cm long; stipules not seen. Inflorescences terminal, sessile, to 12 cm long in
fruit, to 6 cm wide, with 3 pairs of opposite, well-spaced branches, the lowermost
to 2 cm long, each terminated by a 3-flowered cymule. Flowers with the hypan-
thium squarish in lateral view, coriaceous, densely golden villose, the calycine
cup cylindrical, about as wide at the base as at the apex, to 7 mm long, ca. 3.5
mm wide, coriaceous, eglandular within, the hairs outside ferrugineous, long and
spreading, the hairs within appressed, ferrugineous, the margin undulate, with 4
irregular triangular teeth, these often curved, to 1 mm long; corolla with the tube
narrowly cylindrical, ca. 1.7 cm long, to 3.5 mm wide, somewhat falcate, ex-
panded above the middle, coriaceous, densely appressed hairy outside, glabrous
within, the lobes 4, narrowly oblong, ca. 15 mm long; stamens 4, the anthers
narrowly oblong, ca. 7 mm long, sagittate at the base, the filaments plane, ca. 1
mm long, glabrous, attached near the middle of the tube; style slender, ca. 5 mm
long, the stigmas subulate, somewhat plane, ca. 5 mm long. Fruits globose, 1-3
cm long, densely puberulent, the persistent calyx narrowly cylindrical, to 1 cm
long, ca. 0.4 cm wide, the wall of the fruit 2-3 mm thick.

Coiissarca cerroazidensis is known only from Panama. Two striking features
of the species are the elongate petioles, to 5 cm long, and the elongate filaments
of the stamens which are almost as long as the elongate anthers,

Panama: Cerro Jefe, Croat 13052 (MO), 15189 (MO, NY). Without other locality, Duke 8167
(MO). Goofy Lake and Cerro Jefe, Blum et al. 2270 (MO). Cerro Jefe, Duke & Dwycr 15064 (MO);
Duke 8167 (MO). Cerro Azul, Dwyer 1368 (MO). Between Cerro Jefe and La Eneida, Dwver 8199
(MO, NY). Cerro Azul, Galle^os et al. 5174 (MO). Cerro Jefe, 2900 ft. Gentry 2147 (MO). 2-8 mi
S of Goofy Lake, Lewis et al. 2161 (DUKE, F, GH, K, MO, NY, PMA, UC, US). Gorgas Lab, ca. 25
km NE of Cerro Azul, 550 m, Mori & Kallunki 3267 (MO). NE of town of Cerro Azul, 20 km by road
from Panamerican Highway, Mori S: Kallunki 3654 (MO). 5-10 km NE of Altos de Pacora, ca. 750
m, Mori c^ Kallunki 4969 (MO).

2. Coussarea curvigemmia Dwyer, Phytologia 38: 215. 1978. type: Panama, Croat
14863 (MO, holotype).— Fig. 28.

Shrubs or small trees, to 5 m tall, the branchlets terete, glabrous, ashen gray
when dry, the nodes well spaced. Leaves oblong or lanceolate, 6.5-25.0 cm long,

2.0-13.5 cm wide, scarcely or strongly inequilateral, often falcate, acuminate, the

acumen straight or curved, to 1.5 cm long, obtuse, cuneate at the base or some-
times obtuse or lightly auriculate, the costa subplane to prominulous above,
prominulous beneath, the lateral veins 6-9, widely arcuate, the venules 1-3 be-
tween adjacent lateral veins, these branching and reticulate, the petioles lacking
or to 0.5 cm long, glabrous; stipules triangular, to 4 mm long, rounded toward
the apex, glabrous. Inflorescences solitary, terminal, glabrous to puberulent,
thrysoid paniculate, to 6 cm long, to 4.5 cm wide; peduncle slender, 1-2 cm long,
the primary branches alternate or opposite, to 2 cm long, ascending, the rachis
often with 2 pairs of terminal branches, alternate or opposite, flabellate in ap-
pearance, the flowers congested, sessile or subsessile. Flowers with the hypan-
thium oblong or oblong rotund, often shorter than the calyx cup, the latter cylin-
drical or collar shaped, 1.5-2.5 mm long, truncate, stiffly petaloid, glabrous or

] 980]

DWYER— FLORA OF PANAMA (Family 179. Rubiaccac)

123

Figure 28. Cotissarea cunigcmmia Dwyer. — A. Habit {xVi). — B. Fruiting branch (x>/^). — C.
Flower, opened to show stamens and style (x3.5). — D. Ovary cross section (x5). [After Croat 7308. \

124 ANNALS OF THE MISSOURI BOTANICAL GARDI-N [Vol. 67

puberulent on the outside; the teeth 5 or absent, triangular, to 0.5 mm long, often
with minute, glandular red spots on the margin; corolla white, the tube narrow
cylindrical, often falcate, 6-16 mm long, ca. 1.5 mm wide, petaloid, glabrous or
puberulent outside, glabrous within, the lobes 4, narrowly oblong, 7-8 mm long,
glabrous within, not cucullate; stamens 4, the anthers linear, 4.5-5.7 mm long,
acute with a minute apiculum, the filaments short, ca. 1 mm long, attached above
the middle of the tube; ovarian disc prominent, to 0.65 mm high, ca. 1 mm wide,
the style short or long, included or scarcely exserted, linear, 6-16 mm long, the
stigmas 2, linear oblong, 1.5-3.0 mm long, ca. 0.1 mm wide. Fmits ellipsoid,
often laterally compressed, 1.2-1.5 cm long, to 0.7 cm wide, glabrous, at first
fleshy and spongy, the pulp glistening white, the surface often lenticellate when
dry; seed solitary with a hard testa filling the cavity, the calyx persistent, to 1.5
mm long, to 1.2 mm wide.

Coiissarca ciirvlgemmia is known only from Panama and Costa Rica. Most
of the material cited below has previously been erroneously assigned to 0;//.v-
sarca impctiohins Donnell Smith which unlike this species has subquadran-
gular branches, a corolla tube up to 2 mm in width, much more elongate anthers
which are up to 8 mm long, and a fruit which in the dried condition is about 19
mm long. The type of Coussarca Unpetiolaris is Pittier 134008 (US, syntype)

from Costa Rica.

Most fruits on herbarium material have only the fibrous endocarp visible.

One collection, D\vyer2012 (MO) shows the fleshy outside of the fruit still present.
These fruits measure to 1.5 cm in length; those with only the endocarp present
measure to 1.2 cm in length.

BOCAS dilL toro: Almirante, Gentry 2770 (MO), canai_ zone: C-19 Road, Blum 1901 (MO).
Nuevo Emperador, Blum 2393 (MO). K-6 Road, Duycr 2853, 5021 (both MO). Madden Dam, Dnyer
9161 (MO); Harden 74 (MO); Kant 24 (MO). Boy Scout Camp Road near Madden Dam, Mori 4040
(MO). MaddenDam, Porter et al. 4051 (MO, SCZ, UC, VEN). Macapale Island, Tyson 5475 (MO).
BARRO COLORADO ISLAND! Croat 4892, 5465, 5873, 6201, 6282, 7308, 8600, 10295A. 10973, 14863 (all
MO); Ebingcr I7L 565, 610 (all MO); Foster 1779 (MO); Hayden 34 (MO); KdUp 39987 (MO);
Oppenheimer 66-11-2-1259 (MO); Shattuck 135, 621 (both MO); Starry 50 (MO); Wetmorc & Abhe
18 (MO), cocle: Cerro Pilon, Dwyer & Lallatliin 8630 (MO), colon: Santa Rita Ridge, Dnyer et a!.
8982 (MO), darien: Santa Fe, Duke <Sl Bristan 309 (MO). Yaviza. Duke 4875 (MO). El Real to
Pinogana, Duke 50I2A (MO). Santa Fe, Duke 12271, 14091 (both MO); Tyson et al. 4665, 4828 (both
MO). Rio Chucunaque, Stern et al. 845 (MO). Panama: Juan Mina, Bartlett & Lasser 16572 (MO);
Tyson 5459 (MO). Cerro Azul, Dwyer 7093 (MO), Cerro Jefe, Gentry 6781 (MO). Villa Rosario, near
Capira, Saldana 41 (MO).

3. Coussarea durifolia Dwyer.*' type: Panama, Licsner 1180 (MO, holotype).

Small trees to 4 m tall, the twiglets thick, ultimately somewhat angular,
smooth, glabrous, the nodes well spaced, somewhat turgid, the leaf scars trian-
gular or subrotund, ca. 0.5 cm long. Leaves obovate or obovate oblong, to 22 cm
long, to 12 cm wide, obtuse or rounded at the apex, acuminate, the acumen short,
to 5 mm long, cuneate or obtuse at the base, the costa prominulous or subplane
above, prominent beneath, convex, the lateral veins ca. 17, arcuate, the sub-

'*^ Coussarea durifolia Dwyer, spec. nov. Arhores parvae vel frutices. Folia obovata vel obovata-
oblonga, apicem verus obtusa vel rotundala, brevi-acuminata venis lateralibus ca. 17, coriacea in
sicco smargadina glabra; petiolis nullis vel ad 1 cm longis; stipulis ovato-triangularibus. Inflorescentiae
terminales, ramis paucis floribus patulis. Flores sessiles. Fructus non visi.

1980] DWYER— FLORA OF PANAMA (Family 179. /f^WrKCflf) 125

marginal vein undulate, 2-3 mm from the margin, the intervenal areas reticulate,
coriaceous, emerald green beneath when dry, glabrous; petioles absent or to 1
cm long, to 0.35 cm wide, glabrous when dry; stipules free, ovate triangular, to
1 cm long, basally ca. 0.65 cm wide, obtuse, coriaceous, with minute dots outside,
to 0.2 mm in diam. Inflorescences terminal, solitary, to 7 cm long, to 5 cm wide,
glabrous, drying black; peduncle to 2 cm long, to 0.35 cm wide, the branches
few, short, to 2 cm long, the flowers spreading. Flowers sessile, the perianth
parts carnose; hypanthium oblong, 2.5-4.0 mm long, minutely puberulent or gla-
brescent, the calycine cup turgid, collarlike, ca. 3 mm long, ca. 3.5 mm wide,
truncate, the teeth 4 or scarcely visible, ovate oblong, 0.8-1.5 mm long, reflexed,
obtuse; corolla cream colored, the tube narrowly hypocrateriform, ca. 1.4 cm
long, minutely puberulent outside, glabrous within, the lobes 4, narrowly oblong,
7-9 mm long; stamens 4, the anthers narrowly oblong, ca. 6 mm long, ca. 0.7
mm wide, the filaments short, ca. 1.5 mm long, plane, attached near the middle
of the tube; style slender, ca. 4 mm long, the stigmas 2, linear oblong, ca. 2 mm
long. Fruits not seen.

Coussarea durifolia is known only from Panama. It is readily recognized by
its coriaceous leaves which dry an emerald green beneath. The lateral veins are
unusually numerous for a Coussarea. The black punctations of the stipules are
probably diagnostic.

PANAMA: El Llano-Carti highway, 20 km N of El Llano, Dressier 4352 (MO). El Llano-Carti
road, 12 mi above Panamerican Highway, Liesner 1 180 (MO).

4. Coussarea enneantha Standley, J. Wash. Acad. Sci. 18: 282. 1928. type: Pan-
ama, R. S. Williams 841 (US, holotype; isotype, NY).— Fig. 29.

Shrubs 4(-7) m tall, the branchlets subterete, slender, the internodes well
spaced. Leaves elliptic oblong, 9-16(-24) cm long, 3.2-8.0(-13.0) cm wide, acu-
minate, the acumen to 1.5 cm long, usually falcate, obtuse, acute or cuneate at
the base, somewhat inequilateral, the costa prominent, the lateral veins (7-)10-
12, joining to form a usually distinct submarginal vein, the venules reticulate,
membranous to stiffly chartaceous, concolorous or discolorous, villosulose or
villose on veins and costa above and beneath; petioles slender or stout, 1.2-3.5
(-4.2) cm long, 0.1-0.2 cm wide, pubescent; stipules connate, the sheath ca. 3
mm long, pubescent. Inflorescences terminal, usually sessile or the peduncle to
1 cm long, trichotomous, the branches spreading, curved, to 8 cm long, slender,
ca. 1 mm wide, densely golden pubescent, each branch with 3 flowers at the
apex. Flowers sessile; hypanthium including calycine cup narrow cylindrical, to
16 mm long, ca. 3.3 mm wide, the cup more than twice the length of the hypan-
thium, villose outside, glabrous within, stiffly carnose; corolla white or cream
colored, the tube to 33 mm long, ca. 4 mm wide, densely pubescent outside,
sometimes glabrescent, the lobes 4, narrowly oblong, ca. 11 mm long, 2-3 mm
wide basally, glabrous adaxially; stamens 4, the anthers narrowly oblong, ca. 7
mm long, ca. 0.7 mm wide, obtuse, the filaments ca. 6.5 mm long, attached
submedially in the tube; stigmas 2, digitiform, ca. 3 mm long, the style ca. 13 mm
long; ovary oblong in cross section, thin walled. Fruits sessile or pedicellate for
3 mm, oblong, to 3 cm long, to 1.5 cm wide, tapering at the apex and base, the
calyx persistent, 1.0-1.7 cm long, densely pubescent.

126

ANNALS OF THF MISSOURI BOTANICAL GARDEN

[Vol. 67

Figure 29. Coussarea enneantha Standley. — A. Habit (x'/i). — B. Flower (x I). — C. Flower,
dissected showing stamens (xl). — D. Fruit (x 1). [After Duke 15707.]

1980] DWYER— FLORA OF PANAMA (Family 179, Ruhiaccae) 127

Coussarea cnneantha is known only from Panama. One collection, D^vyer <&.
Correa 8892, has extraordinarily large leaves, up to 24 cm long and 13 cm wide;
the indumentum is somewhat reduced on the lower surface of the blade and the
fruits are only up to 2 cm long.

BOCAS DEL TORo: Fish Creek Mountains, Wedel 2364 (MO), cocle: Cerro Caracoral. 900 m,
Duke & Dwxer 15139 (MO). Cerro Gaital Caracoral, Dwycr & Correa 8892 (MO). Cerro Pilon, Porter
et al. 4402 (MO), darien: Cerro Pirre, Bristan 536 (MO). Rio Pirre, Bristan 1462 (MO). Cerro
Campamiento, S of Cerro Pirre, Duke 15707 (MO). Cerro Pirre, 3500-4500 ft, Duke & Elias 13781
(MO, UC, VEN). Cana-Cuasi trail on Cerro Campamiento, E of Tres Bocas, Klrkbride & Duke 1239
(MO, NY). Cana, Williams 841 (NY, US). Panama: 6.5 km on road N of Goofy Lake, Nee 9301
(MO).

5. Coussarea garciae Standley, Publ. Field Mus. Nat. Hist. Bot. Ser. 22: 180.
1940. type: Colombia, KiUip & Garcia 33524 (F, holotype).

Shrubs or small trees 2.0-3.5 m tall, the branchlets stout, angular above,
lenticellate, the nodes somewhat turgid, glabrous, the stipular scars thin, ringlike.
Leaves sessile, obovate, to 50 cm long (or longer?), to 20 cm wide, obtuse or
rounded at the apex, occasionally short acuminate, tapering narrowly and cu-
neately toward the base and finally vaguely auriculate, the costa slender and
prominulous above, prominent beneath, to 0.3 cm wide, lignose, the lateral veins
ca. 25, to 3 cm apart, prominulous above, prominent beneath, the veinlets per-
pendicular, thin coriaceous, occasionally papyraceous, concolorous, drying
green, glabrous above and beneath; stipules free, deltoid, to 1.5 cm long, sub-

■

coriaceous, marcescent, glabrous. Inflorescences terminal, to 5 cm long, the cen-
tral axis suppressed or to 3 cm long, the branches few, short, the flowers densely
aggregated. Flowers with the hypanthium ca. 2 mm long, the calycine tube cy-
lindrical, ca. 7 mm long, rounded basally, somewhat turgid, glabrous, petaloid,
with a few, well spaced, reddish glands within, the calycine teeth 4, narrowly
oblong or narrowly ovate oblong, 10-12 mm long, ca. 2 mm wide at the base,
acute; corolla blue, the tube narrowly cylindrical, somewhat curved like an S, ca.
15 mm long, wider above the middle, ca. 2 mm wide at the middle, glabrous,
stiffly thin carnose, the lobes 4, narrowly oblong, ca. 6 mm long; stamens 4, the
anthers oblong, 4.5-6.0 mm long, subsagittate basally, the filaments slender, 3
mm long, attached near the middle of the tube; style slender: 4 mm long, the
stigmas digitiform, 2 mm long. Fruits on pedicels 0.3 cm long, elliptic oblong,
2.0-2.8 cm long, to 1.3 cm wide, obtuse at the apex and the base: light brown,
smooth, glabrous, the raphides conspicuous under magnification, ribbed, the per-
sistent calyx to 2 cm long, the lobes often withering, narrowly ovate lanceolate, to
12 mm long, glabrous.

Coussarea garciae is known only from Darien and the Dept, of Choco, Co-
lombia. Duke (1 1350) notes that the fruits are eaten in Colombia. 'Xolorin."

Panama: darien: Between the upper Rfo Membrillo and Camp 7 on the road to San Bias, Duke
10878 (MO). Hydro Camp No. 14, Rio Salaqui, 6 days upstream from Rio Sucio, Duke 11350 (MO).
Rio Tuquesa, lower Tuquesa Mining Co. camp, Mori 6982 (MO). Camp Summit, adjacent to Darien-
San Bias border. Sea Level Canal, Route 17, 1000-2000 ft, Oliver et al. 3679 (MO). Colombia:
Between Ri'o Nercua-Quebrada Barrial and Quebrada Ambrosio near Camp Curiche, 10-294 m, Duke
11435 (MO). Bahia Solano, dense forest along Quebrada Jellita, 50-100 m, Killip & Garcia 33524 (F,

US).

128 ANNALS OF THE MISSOURI BOTANICAL GARDHN [Vol.67

6. Coussarea jefensis Dwyer.^- type: Panama, Mori & Joly 7927 (MO, holotype).

Trees to 6 m tall, the branchlets terete, smooth, glabrous, drying dark, the
nodes well spaced, turgid, the leaf scars elevated, round, drying whitish with a
central circular vascular bundle. Leaves oblong rotund to oblong, 3.0-8.5 cm
long, 1-5 cm wide, rounded to obtuse at the apex, occasionally vaguely acumi-
nate, obtuse or rounded at the base, the costa prominulous above and beneath,
drying red, to 0. 15 cm wide proximally, the lateral veins ca. 9, arcuate, prominent
beneath, forming an undulate submarginal vein with usually 1 irregular interme-
diate vein between adjacent lateral veins, the intervenal areas patulous reticulate,
the margin slightly revolute, coriaceous, concolorous, drying dark, glabrous,
glandular shiny; petioles 1 .0-1.3 cm long, stiff, glabrous, to 1 .5 mm wide; stipules
connate, forming an expanded, coriaceous, glabrous and truncate cup, to 2.5 mm
long. Inflorescences (in fruit) terminal, paniculate, glabrous, drying dark, to 8.3
cm long, to 4.5 cm wide; peduncle 3.5-4.0 cm long, piano compressed, the low-
ermost branches opposite, paired, ascending, to 2 cm long, the upper branches
to 6, alternate; bracts and bracteoles minute, ca. 1 mm long. Fruits subsessile,
oblong, rarely vaguely curved, terete in cross section, to 1 cm long, drying black,
glabrous, vaguely 2-lobed, not costate; seed solitary, oblong, to 5 mm long, to 4
mm wide, obtusely 3-angled in cross section, with a thin horizontal ridge on the
adaxial surface.

Coussarea jefensis is immediately distinguished from all other Coussarea in
Panama by its thick-coriaceous leaves which are oblong rotund to rotund. The
elevated large leaf scars are diagnostic.

Panama: Cerro Jefe, ca. 2900 ft, Mori & J»l\ 7927 (MO).

7. Coussarea latifolia Standley, J. Wash. Acad. Sci. 18: 281. 1928. type: Costa
Rica, Tonduz. 9574 (US, holotype).

Trees to 15 m tall, glabrous throughout, the twiglets stout, 4-angled, lenticel-
late. Leaves broadly elliptic obovate to broadly ovate elliptic, 16.5-25.0 cm long,
9.0-16.5 cm wide, acute or obtuse at the base, the costa and lateral veins stout,
salient beneath, the lateral nerves 9, arcuate, papyraceous, concolorous, pale

green when dry; petioles stout, 1.7-2.5 cm long; stipules persistent above, semi-
orbicular, ca. 5 mm long, wider than long, intrapetiolar, shortly connate, broadly
rounded at the apex. Inflorescences terminal, glabrous, to 6 cm long, to 5 cm
wide, the peduncle to 2 cm long, to 0.35 cm wide. Flowers sessile or subsessile;
hypanthium short, glabrous, the calycine cup cylindrical, ca. 9 mm long, ca. 6
mm wide at the apex, the latter truncate, coriaceous, glabrous, dentate, eglan-
dular; corolla (seen only in part) narrow below, presumably falcate, coriaceous.

'" Coussarea jefensis Dwyer, spec. nov. Arhores ad 6 m altae ramulis laevibus glabris nodis
cicatricibus foliorum conspicue rotundis ornatis. Folia oblongo-rotunda ad oblonga, 3.0-8.5 cm longa,
1-5 cm lata, apice rotundata vel obtiisa venis iateralibus ca. 9, coriacea, glabra; petiolis 1.0-1.3 cm
longis: stipulis connatis vagina cupulata. ad 2.5 mm longa, truncata coriacea glabra. Inflorescentiae
terminales paniculatae, ad 8.3 cm longae, ad 4.5 cm latae, pedunculo 3.5-4.0 cm longo, ramulis
inferioribus ad 2 cm longis, superioribus ca. 6 alternatis; bracteis et bracteolis minutis. Friictiis
oblongi, ad 1 cm longi, glabri, vix 2-lobati.

19801 DVVYER— FLORA OF PANAMA (Family \79. Ruhiaccac) 129

glabrous outside, pubescent within; stamens 4, included in the tube, the anthers
lunate or S-shaped, 5-6 mm long, tapering at the apex, cordate at the base, the
filaments Vi to almost as long as the anthers, presumably attached at the middle
of the tube; ovarian disc prominent. Fruits oblong, to 2 cm long, often somewhat
falcate, obtuse or rounded at the apex, green to yellow, lenticellate or smooth,
the calycine cup short, smooth, glabrous.

Coussarea hitifolia is known only from Costa Rica, Panama, and Colombia.

BOCAS DEL TORo: Above Almirante. Gentry 2765 (MO). Fish Creek Hills. Chiriqui' Lagoon,
Wcdel 2218. 2463 (both MO), colon: 10 mi S of Portobelo, 2-4 mi from the coast, 10-2(X) m, Licsncr
1071 (MO). PANAMA: Cerro Jefe, ca. 700 m, Corrca & Dressier 1158 (MO). Between Cerro Azul and
Cerro Jefe, Mori & Kalliinki 2173 (MO).

8. Coussarea loftonii (Dwyer & Hayden) Dwyer, comb. nov.

Faramea loftonii Dwyer & Hayden, Phytologia 15: 56. 1967. type: Panama, Tyson et al. 3323 (MO,
holotype).

Shrubs or trees, to 10 m tall, the branchlets terete, drying yellow grey, gla-
brous, the nodes usually well spaced. Leaves salicoid, often falcate, 5-17 cm
long, 1.8-5.0 cm wide, the larger blades regularly 3 times longer than wide,
tapering acutely or deltoidly at the apex, acuminate, the acumen attenuate, usu-
ally narrowly falcate, to 1.5 cm long, 0.1-0.3 cm wide in the middle, acute at the
base, slightly inequilateral, the costa scarcely prominulous above, rimose, sub-
plane beneath, the lateral veins 9-12, prominulous, at first strict and then arcuate,
or strict to the margin, the intervenal areas markedly reticulate, papyraceous,
concolorous, glabrous; petioles to 1 cm long when present, narrowly alate; sti-
pules connate, often persistent, deltoid or compressed rotund, wider than long.

2.5-3.0 mm long, 3

Jnfl.

v_

to 5 cm long, to 5 cm wide, cymose paniculate, the peduncle to 2 cm long, ca.
1.5 mm wide, glabrous, the branches 3-4, the lowermost often paired, opposite
or subopposite, divergent or ascending, to 1 .5 cm long, the terminal cymules with
3-6 flowers; lowermost bracts rarely persisting, narrowly elliptic, to 15 mm long,
ca. 1.5 mm wide, glabrous. Flowers with the pedicels 1.5-3.0 mm long, glabrous;
hypanthium 0.5-1.2 mm long, smooth, glabrous, the calycine cup and teeth to
2.5 mm long, the cup with a few appressed glands, coriaceous, the teeth 4, sub-
ulate, to 0.7 mm long; corolla white, the tube narrowly cylindrical, to 9 mm long,
ca. 2 mm wide, glabrous, stiffly petaloid, the lobes 4, lanceolate, to 7.5 mm long,
tapering narrowly toward the apex; stamens 4, the anthers narrowly oblong, 4-
5 mm long, ca. 0.5 mm wide, obtuse, caudate at the base, dorsifixed, the filaments
ca. 1 mm long, attached near the middle or at the upper Vs of the tube; stigmas
2, subulate, to 2.2 mm long, the style 10-13 mm long. Fruits scarcely pedicellate,
oblong, to 1.7 cm long, ca. 1 cm wide, smooth, glabrous, white, drying yellow,
spongy within, the persistent calycine cup small, to 1.5 mm long, ca. 1 mm wide.

Coussarea loftonii is known only from Panama. It was originally described
from flowering material, but a recent collection, Lewis & Dressier 6567, has fruits
which are obviously those of a Coussarea and not of Faramea. Nee 9297 de-
scribes the flowers as very fragrant.

130 ANNALS OF THE MISSOURI BOTANICAL GARDIiN [Vol. 67

colon: Santa Rita Ridge, Correa & Dressier 667 (MO); Dressier 3852 (MO). Santa Rita Ridge
Road, Gentry et id. 8H64 (MO). El Llano-Carti road 9.6-11 km from Interamerican Highway, Mori

& Kallunki 3521 (MO). Santa Rita Ridge Road, 14 km from the Boyd-Roosevelt Highway, Mori &
Kallunki 4904 (MO). Panama: La Eneida, Lopez House, Correa & Dressier 818 (MO). Cerro Jefe,
Lewis ct Dressier 7567 (MO). NE of town of Cerro Azul, Mori ct Kallunki 3641, 3649 (both MO);
NEL 9297 Cerro Jefe, Tyson et al. 3323 (MO); Wilhnr & Weaver 11373 (MO).

9. Coussarea morii Dwyer/**^ type: Panama, Mori 7058 (MO, holotype).

Trees to 12 ft tall, ca. 20 cm d.b.h., the twiglets slender, smooth, glabrous,
the nodes well spaced. Leaves oblong, 10.0-24.5 cm long, 3.5-12,0 cm wide,
obtuse to widely cuneate at the apex, acuminate, the acumen to 1.5 cm long,
falcate, acute to obtuse at the base, the costa subplane above, subprominent
beneath, the lateral veins 7-10, arcuate, prominulous beneath, the submarginal
undulate vein vague, the intervenal areas patulous reticulate, thin chartaceous,
scarcely discolorous, drying green, glabrous except for tufts of hairs in the axils
of most veins; petioles 1.5-2.0 cm long, rigid, strict, glabrous, drying dark; stip-
ules not seen; glands at stipular scar position numerous, narrow oblong, ca. 1
mm long, drying red. Inflorescences terminal and axillary, cymose paniculate, to
19 cm long, to 6 cm wide; peduncle ca. 3 cm long, ca. 0.3 cm wide, angular and
compressed, drying black, glabrous; primary branches disposed as 3-4 pairs,
opposite or subopposite, well spaced, often at right angles to the rachis, the
lowermost to 2.5 cm long; cymes few, clustered at the tips of reduced secondary
branches; bracts and bracteoles apparently absent except for 2 mammiform swell-
ings in the axils of the uppermost leaves, 1 per axil, ca. 1 mm long, glabrous,
drying black. Flowers sessile or subsessile, externally glabrous; hypanthium to
2 mm long, the calycine cup and hypanthium chalicelike, the cup ca. 2 mm long,
turgid, petaloid, the teeth as vague points or distinctly 5, oblong, to 1 mm long,
obtuse, the cup with a few appressed glandular spots within; corolla white, the
tube narrow cylindrical to 5 mm long, to 1.3 mm wide in the middle, flaring at
the very base; stamens 4, the lobes 4, oblong, slightly longer than the tube, to 8
mm long, the anthers narrow oblong, ca. 4 mm long, exserted, the filaments to
2 mm long, attached below the mouth; style slender, to 6.5 mm long, glabrous,
the stigmas digitiform, to 1 mm long, as wide as the style; ovules 2, connate.
Fruits not known.

Coussarea morii is known only from Panama. It is named in honor of Scott
Alan Mori, distinguished student of tropical American plants. The new species
is closely allied to C. paniculata (Vahl) Standley, but differs in having both

'^'^ Coussarea morii Dwyer, spec. nov. Arhores ramulis glabris. Folia oblonga, 10.0-24.5 cm
longa, 3.5-12.0 cm lata, apice obtusa vel lato-cuneata, acumine ad 1.5 cm longo, venis lateralibus 7-
10, tenui-coriacea glabra; petiolis 1.5-2.0 cm longis. Inflorescentiae terminales et axillares, cymoso-
paniculatae, ad 19 cm longae, ad 6 cm latae, pedunculo ca. 3 cm longo glabro ramis primariis in jugis
3-4 dispositis. riores sessiles vel subsessilcs hypanlhio ca. 2 mm longo glabro cupula calycis ca, 2
mm longa, turgida petaloidea dentibus saepe inconspicuis vel alitcr ad 1 mm longis, obtusis glabris;
corolla tubo cylindrico, ca. 5 mm longo, leviter curvato exlus glabro intus villosuloso supra medium;
antheris 4 angusto-oblongis, ca. 4 mm longis exsertis filamentis ad 2 mm longis infra faucem affixis;
stigmatibus digitiformibus, ad 1 cm longis. Frmtus not visi.

1980J DWYHR— FLORA OF PANAMA (Family 179. Rublaceae) 131

terminal and axillary inflorescences which are often twice the length of those of
C panicidata\ usually a single small mammiform swelling in the axils of the
uppermost leaves subtending the peduncle; thin chartaceous rather than coria-
ceous leaves; glabrous hypanthium with the calycine cup glabrous and occasion-
ally with teeth to 1 mm long; corolla with oblong lobes which measure about 2
mm in diameter. Dr. Mori records that the tree is 12 meters in height. Apparently
C. paniculata does not exceed 8 meters in height (fide Steyermark, 1974).

darien: Rio Tuquesa, at middle Tuqesa Mining Company Camp called Charco Peje, tropical wet
forest, 250 m, Mori 7058 (MO).

10. Coussarea nebulosa Dwyer.^^ type: Panama, Croat 15868 (MO, holotype).

Shrubs 3.0-3.5 m tall, the twigs smooth, dark grey when dry, often rimose,
the nodes swollen, well spaced. Leaves oblong, 6.0-10.5 cm long, 2.5-3.5 cm
wide, acute toward the apex, acuminate, the acumen narrowly deltoid, to 1 cm
long, the costa plane or prominulous above, prominulous beneath, the lateral
veins 5-6, arcuate, extending to the margin, subchartaceous, somewhat discol-
orous, glabrous, the raphides minute, white, dense; petioles slender, to 1.2 cm
long, ca. 0.13 cm wide; stipules connate, the sheath slightly turgid, ca. 2 mm
long, each part of the sheath with 2 well-spaced awns shorter than the sheath.
Inflorescences terminal, solitary, glabrous, to 5 cm long, to 3 cm wide, the pe-
duncle to 2.5 cm long, the lowermost branches opposite, to 1.5 cm long, the
upper branches few; lowermost bracts subulate, to 3 mm long. Flowers sessile;
hypanthium subrotund, ca. 0.6 mm long, the calycine cup glabrous, 0.5 mm long,
lacking glands within, the teeth 5, triangular, 0.5 mm long; corolla white, ca. 25
mm long, the tube ca. 18 mm long, carnose, glabrous outside, villose inside, the
lobes 5-6 mm long; stamens 5, the anthers linear oblong, 4 mm long, 0.35 mm
wide, the filaments ca. 0.5 mm long, the style 19 mm long, ca. 0.3 mm wide, the
stigmas 2, ca. 2 mm long. Fruits not seen.

Coussarea nebulosa is known only from Panama. The new species has the
fewest lateral veins (5-6) among the Panamanian Coussarea; its connate stipules,
each part with 2 short awns, are unique among the Panama Coussarea, The type
was collected near Boquete at about 4,000 ft elevation. This is probably the
highest elevation at which a Panamanian Coussarea has been collected. The
specific name ''nebulosa" was selected as it suggests a cloud forest.

CHiRiQuf: Uncut cloud forest at Monte Rey near Boquete, Croat 15868 (MO).

'■^^ Coussarea nebulosa Dwyer, spec. nov. Frutices 3.0-3.5 m alti. Folia oblonga, 6.0-10.5 cm
longa, 2.5-3.5 cm lata, versus apicem acuta, acumine ad 1 cm longo venis lateralibus 5-6 subchartacea
glabra; petiolis ad 1.2 cm longis; stipulis connatis vagina subturgida, ca. 2 mm longis, parte utraque
setis duabus quam vaginam brevioribus. Inflorescentiae terminales solitarieae glabrae, 5 cm longae,
ad 3 cm latae, pedunculo ad 2.5 cm longo. Flores cupula calycis 0.5 mm longa, glabra eglandulosa
dentibus 5, 0.5 mm longis; corolla alba tubo ca. 18 mm longo, intus villoso, lobis 5-6 mm longis;
staminibus 5, antheris lineari-oblongis, 4 mm longis, filamentis ca. 0.5 mm longis. Fructus non visi.

132 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

11. Coussarea neei Dwyer.^^ type: Panama, Ncc 9820 (MO, holotype).

Trees or shrubs, 4-7 m tall, the twigs leafless except at tips, gray, cracking,
often flexuous, the nodes turgid. Leaves lanceolate or oblong-lanceolate, often
inequilateral and falcate, 5-16 cm long, 1.8-5.0 cm wide, acute, obtuse or rounded
at the apex, the acumen absent or conspicuous, to 1 cm long, acute to obtuse at
the base, the costa immersed above, prominulous or subprominent beneath, the
lateral veins 9-10, often widely arcuate, the intervenal areas spreading reticulate,
thin papyraceous, discolorous, often drying brown, arachnoid pubescent only on
the costa above, pubescent on the costa beneath, occasionally on the veins;
petioles to 2.5 cm long, 1.3 cm wide; stipules free, compressed rotund, to 0.4 cm
long, rotund at the apex, smooth, glabrous, ciliolate marginally. Inflorescences
terminal, solitary, ferrugineous villose or hirsute, 2.5-8.0 cm long, 2.5-9.0 cm
wide, the peduncle reduced, 0.5-0.8 cm long, ca. 1 mm wide, the branches 2,
opposite, the cymules terminal, the flowers few. Flowers (in bud), with the hy-
panthium oblong, to 3 mm long, ferrugineous hirsute, the calycine cup ca. 2 mm
long, subcarnose, villosulose within, the lobes 4-5, subulate, often scarcely in-
equilateral, to 3 mm long, erect, densely golden villose. Fruits sessile, narrowly
oblong, to 2.7 cm long, to 0.8 cm wide, drying black, densely golden puberulent.

Coussarea neei is known only from Panama. It is named in honor of Michael
Nee, who collected extensively in Panama in 1973-1974. It is closely related to
Coussarea enneantha Standley, as is evident from the structure of the inflores-
cence. The new species has a much shorter calycine cup with conspicuous ca-
lycine teeth.

darh'n: Serrania de Pirre, ascent of Serrania de Pirre above Cana gold mine, 600-1000 m, Croat
37737 (MO). viRAGUAs: Valley Rio Dos Bocas, between Escuela Agricola Alto Piedra and Calove-
bora, 15.6 km NW of Santa Fe, 450-550 m, Croat 27649 (MO). 5 mi NW of Sant Fe, above Rio
Primero Brazo, Licsner 823 (MO). 8.8 km from Fscuela Agricola Alto de Piedra, Mori & Kallunki
3216 (MO). 6-7 km W of Santa Fe, new road past agricultural school, 2900 ft, Ncc 9708 (MO), 9820
(MO, holotype).

12. Coussarea paniculata (Vahl) Standley, J. Wash. Acad. Sci. 18: 282. 1928.

Froclichia paniculata Vahl, Ecol. Amer. 3. 1796. not Moench. 1794. type: French Guiana, not seen,
Billardicria paniculata Vahl, Ecol. Amer. 10. 1796. type: French Guiana, ''In sylvis prope amncm

Galibiensem,'' not seen.
Faranica martini DC. ex Griseb., Fl. Brit. W. 1. 339. 1861, not DC. 1830. type: French Guiana, not

: photo in Delessert Herb. (F).
Coussarea froclichia A. Rich., Mem. Soc. Hist. Nat. Paris 5: 177. 1834. type: not seen.
C. daricncnsis Steycrmark, Ceiba 3: 20. 1952. type: Panama, Allen 4576 (F, holotype; isotype, NY).

Small trees. Leaves widely elliptic, 8.5-14.0 cm long, 3-6 cm wide, caudate
acuminate at the apex, the acumen to 1.8 cm long, cuneate, acute, often attenuate

^ Coussarea ncci Dwyer, spec. nov. Arhorcs vel frutices, 4-7 m altae. FoUa lanceolata vel oblongo-
lanceolata, saepe inaequilateralia et falcata, 5-16 cm longa, 1.8-5.0 cm lata, venis lateralibus 9-10,
tenui-papyracea supra in costa solum arachnoideo-pubescentia subtus in costa et saepe in venis
pubescentia; pctiolis ad 2.5 cm longis; stipulis compresso-rotundis, ad 0.4 cm longis. apice rotundis.

Inflorcsccntiac terminales solitariae ferrugineo-villosae vel hirsulae, 2.5-8.0 cm longae, 2.5-9.0 cm
latae, pedunculo 0.5-0.8 cm longo, ca. 1 mm lato, ramis 2 oppositis dispositis cymulis terminalibus

paucos flores ferentibus. Flores cupula calycis ca. 2 mm longa, lobis 4-5, ad 3 mm longis. Fructus
sessiles angusto-oblongi, au 2.7 cm longi, ca. 0.8 cm lati, in sicco nigri dense aureo-puberuH.

1980] DWYER— FLORA OF PANAMA (Family 179. Riihiaceae) 133

at the base, the lateral veins ca. 8, stiffly chartaceous, discolorous or concolorous,
glabrous except for a few hairs in the axils of the veins below; petioles to 1.5 cm

Infl.

9.0 cm

long, to 6 cm wide, the peduncle to 1.5 cm long, the branches few, paired,
perpendicular to the rachis, the lowermost to 2.5 cm long. Flowers subsessile or
on pedicels to 0.5 cm long; calycine cup glabrous, turgid, to 1 mm long, smooth,
the teeth scarcely measurable; corolla white, the tube to 3 mm long, the lobes
ca. 6.5 mm long; anthers linear, to 5 mm long, the filaments to 1 mm long attached
near the mouth. Fruits on pedicels to 0.5 cm long, ovoid, 1.2-1.6 cm long, 1.0-
1.5 cm wide, yellow, white or green, glabrous, drying black.

Coussarea paniculata is known from Panama, Trinidad and northern South
America. The isotype of C. darienensis is deposited in NY, not in the Missouri
Botanical Garden herbarium, contrary to Steyermark's indication in the original
description.

darien: Rio Chico, Yaviza, ca. 100 ft, Allen 4576 (F, NY). Puerto St. Dorothea, near Colombia-
Panama border on Pacific side, Dwyer 2222 (MO).

13. Coussarea roseo-cremea Dwyer.^*' type: Panama, Dressier 4334 (MO).

Shrubs to 3 m tall, the branchlets ultimately obtusely angular, smooth, gla-
brous, the lenticels minute, few. Leaves oblong, 6.5-13.0 cm long, 3.5-7.0 cm
wide, obtuse, rotund or widely deltoid at the apex, rounded or obtuse at the base,
acuminate, the acumen short, to 0.5 cm long, acute or obtuse, the costa promi-
nulous above and beneath, the lateral veins 7-8, widely arcuate, forming an
undulate submarginal vein, the intermediate veins 1-4, soon branching; petioles
moderately slender, stiff, glabrous, 1.0-2.5 cm long, to 0.12 cm wide; stipules
often short connate, persistent, hemispherical, turgid, to 0.4 cm long, each part
with 2 awns. Inflorescences solitary, terminal, cymose paniculate, to 6.5 cm long,
to 5.5 cm wide, the peduncle to 2.5 cm long, the lowermost branches few, 1-3,
opposite or verticillate, to 3 cm long, the flowers terminal, few. Flowers sessile;
hypanthium subrotund, ca. 1 mm long, shiny, glabrous, the calycine cup to 0.5
mm long, subturgid, petaloid, glabrous, eglandular within; corolla pink, the tube
narrow cylindrical, ca. 15 mm long, wider above the middle, puberulent inside
and out, petaloid, the lobes 5, linear subulate, ca. 10 mm long, 1 mm wide at
base, the margins contiguous above the middle, flat, venose near the mouth,
petaloid; stamens 5, the anthers narrowly oblong, 3.3 mm long, subacute, not
exserted, the filaments short, 1.1 mm long, attached above the middle of the tube;
ovarian disc 0.4 mm long, 0.9 mm wide, the style slender, 15 mm long, the stigmas
2, subulate, 1.5 mm long. Fruits not seen.

^''Coussarea roseo-cremea Dwyer. spec. nov. Friitices ad 3 m alti. Folia oblonga, 6.5-13 cm
longa, 3.5-7 cm lata, apice obtusa, rotundata vel lato-deltoidea, acumine ad 0.5 cm iongo, basi
rotundata vel obtusa venis lateralibus 7-8; petiolis 1-2.5 cm longis; stipulis saepe brevi-connatis,
persistentibus hemisphaericis turgidis ad 0.4 cm longis parte utraque aristis duabus praedita. Inflo-
rescentiae solitariae terminales cymoso-paniculatae, ad 6.5 cm longae, ad 5.5 cm latae, pedunculo ad
2.5 cm Iongo, floribus terminalibus paucis. Flores cupula calycis ad 0.5 mm longa, dentibus nullis;
corolla roseo-cremea, tubo ca. 15 mm Iongo, lobis 5 lineari-subulatis ca. 10 mm longis: antheris
angusto-oblongis, 3.3 mm longis, filamentis 1.1 mm longis. Friictus non visi.

134 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

Coussarea rosco'crcmca is known only from the single collection in Panama.
It is closely related to Coussarea nehidosa Dwyer which also has connate stipules
with awns, few lateral veins in the leaf blades and a very reduced calycine cup.
Coussarea roseo-cremea, however, is a lowland species with pink flowers, small-
er anthers and corolla lobes almost equal to the corolla tube in length. The Latin
name roseo-cremea refers to the flower color.

Panama: EI Llano-Carti Road, 6-10 km N of El Llano, Dressier 4334 (MO).

14. Coussarea veraguensis Dwyer. •*^ type: Panama, Mori & Kallunki 4800 (MO,
holotype).

Trees to 4 m tall, the branchlets rigid, angular, glabrous, sharply ascending,
the nodes well spaced, somewhat turgid. Leaves narrowly or widely elliptic, often
slightly falcate, rarely inequilateral, 14-20 cm long, 3.5-10.0 cm wide, deltoid
toward the apex, acuminate, the acumen to 2 cm long, often ultimately acute,
cuneate or obtuse at the base, the lateral veins 11-14, arcuate, disappearing
toward the margin, the intervenal areas reticulate, the raphides minute, white,
rigidly membranous, concolorous, glabrous or puberulent below on the costa and
veins; petioles thick, 0.5-1.0 cm long, to 0.25 cm wide, puberulent; stipules de-
ciduous, not seen. Inflorescences terminal, racemelike, to 5 cm long, to 3 cm
wide, the peduncle slender, 1-2 cm long, subtended by 2 large leaflike bracts 2.0-
5.5 cm long, 0.8-1.5 cm wide, each bract with an erect linear-oblong process in
the axil, to 0.4 mm long, obtuse, black, glabrous, the flowers 15-20, either solitary
on the rachis or in 2-3 sessile clusters. Flowers with the pedicels short, 1-3 mm
long, glabrous; hypanthium oblong, ca. 2 mm long, glabrate, the calycine cup
cupuliform, somewhat turgid, to 0,6 mm long, minutely puberulent only on the
margin, inside with 10 rotund glands of various size, dark red; corolla white,
the tube narrowly cylindrical, ca. 7 mm long, 1.5 mm wide, subfalcate in
the bud, narrow toward the base, carnose, glabrous, the lobes 4, carnose, nar-
rowly oblong, to 5 mm long, stamens 4, the anthers narrowly rectangular, ca. 4
mm long, obtuse, sagittate at the base, the filaments arising near the middle of
the tube; disc of the ovary compressed rotund, to 0.8 mm long, the style plane
compressed, ca. 5 mm long, the stigmas 2, fingerlike, to 2 mm long. Fruit not
seen.

Coussarea veraguensis is known only from Panama. It is distinguished by its
racemelike inflorescence in which the flowers arise either in solitary fashion or
are disposed as clusters on 2 or 3 reduced branches of the rachis. The 2 large

■*^ Coussarea veraguensis Dwyer, spec. nov. Arbores ad 4 m altae. Folia angusto- et lato-elliptica,

14-20 cm longa, 3.5-10 cm lata, versus apicem deltoidea acumine ad 2 cm longo, plerumque ultime
acute, basi cuneata vel oblusa venis laleralibus 11-14, rigido-membranacea. glabra vel puberula;
petiolis 0.5-1 cm longis. Infloresventiae terminales racemo similes, ad 5 cm longae, ad 3 cm latae,
pedunculo gracili, 1-2 cm longo, bracteis basi pedunculi 2 magnis foliis similibus, 2-5.5 cm longis,
floribus 15-20 solitariis in rhachidi vel 2-3 fasciculis sessilibus dispositis. Flores pedicellis 1-3 mm
longis; cupula calycis cupuliformi, ad 0.6 mm longo, solum in margine minulo-puberula intus decern
glandulis rotundis praedita; corolla alba; tubo angusto-cylindrico, ad 7 mm longo, glabro, lobis 4, ad
5 mm longis: staminibus 4, antheris angusto-rectangularibus ca. 4 mm longis. Fructus non visi.

19801 DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 135

leaflike bracts have a distinct oblong process at the base; the corolla tube appears
to be shortest (only 7 mm long) among the Panamanian Coussarea except for C
villosula,

Coussarea veragiiensis appears to be closely related to C Carolina Standley,
a Costa Rican species, and to C. panicidata (Vahl) Standley, which occurs in
Panama and in Costa Rica.

VERAGUAS: NW of Santa Fe, 1 km from Escuela Agricola Alto de Piedra, on the slopes of Cerro
Tute, Mori & Kallunki 4800 (MO).

15. Coussarea villosula Dwyer, Ann. Missouri Bot. Gard. 53: 105. 1966. type:
Panama, Duke 5338 (MO, holotype).

Small trees, the branchlets terete, nodose, minutely puberulent. Leaves lan-
ceolate, to 20 cm long, to 8 cm wide, acuminate at the apex, acute basally, the
lateral veins ca. 15, ca. 1 cm apart, the costa prominulous above, prominent
proximally beneath, the margin delicately revolute, rigidly papyraceous, glabrous
above, sparsely villosulose and conspicuously reticulate beneath; petioles 1-4 cm

long, somewhat thick, puberulent; stipules triangular, to 8 mm long, hirsute.
Inflorescences terminal, 2-4 cm long, narrow, to 2.5 cm wide, densely sericeous
villose; peduncle slender, 2-3 cm long, the cymules 3-4-flowered; bracts subulate,
to 2 mm long. Flowers with the buds fusiform, to 0.5 cm long; calycine cup bell
shaped, to 2.5 mm long, ca. 1.6 mm wide, carnose, densely appressed villose
outside, the hairs to 0.8 mm long, the teeth 5, scarcely measurable or to 0.5 mm
long; corolla white, the tube about 3,2 mm long, densely appressed villose out-
side, glabrous within, carnose, the lobes 5, imbricate, ca. 1 mm long, falcate and
obtuse at the apex; anthers subsessile, somewhat wedge shaped, to 1.3 mm long;
style slender, to 1 mm long; stigmas small; ovules 2, brown when dry, separated
by a thin septum. Fruits not seen.

Coussarea villosula is known only from Panama. Its spindle shaped buds and
greatly contracted hairy inflorescence readily distinguish it.

cocle: F1 Valle from village to La Mesa, ca. 2000 ft, Dwver & Corrca 7925 (MO), dariln:

Cerro Pirre, from Rio Pirre S to El Real, 750-1030, Duke 5338 (MO).

26. COUTAREA

Coutarea Aubl., Hist. PI. Guiane 314. 1775. type: C. speciosa Aubl.

Shrubs or small trees. Leaves membranous, glabrate, petiolate; stipules short,
acute. Inflorescences terminal or axillary, cymose, the flowers occasionally sol-
itary (not Panama). Flowers with the calyx 5-8-lobed, deciduous in fruit; corolla
funnel shaped or campanulate, the tube asymmetric, curved, gibbous ventricose,

the lobes 5-8, short, plicate imbricate; anthers exserted, the filaments free, not
attached to the corolla tube; ovules numerous, the ovary 2-celled. Fruits capsular,
compressed oval, usually coriaceous and conspicuously lenticellate, dehiscing
longitudinally; seeds broadly winged.

Coutarea ranges from Mexico to Argentina, with about 7 species in the genus.
I have some doubts as to whether this genus belongs in the Rubiaceae. The unique

136 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

corolla shape, the staminal filaments being free of the corolla tube, and the unsusal
shape of the fruit are the principal characters which render the genus suspect.

1. Coutarea hexandra (Jacq.) K. Schum. in Mart., Fl. Bras. 6(6): 196. 1889.

Portlandia hexandra Jacq., Enum. PL Carib. 16. 1760. type: Colombia, Cartagena, not seen.

Coutarea flavcsccns Moc. & Ses., DC, Prodr. 4: 350. 1830. typc: not seen.

C. hexandra puheseens K. Schum. in Mart., Fl. Bras. 6(6): 198. 1889. type: not designated.

Small trees or shrubs, the twiglets terete, glabrous, relatively smooth, the
lenticels often abundant and conspicuous, the branchlets rigid or subarcuate, the
nodes usually well spaced. Leaves usually ovate rotund to ovate, 4-12 cm long,
2-6 cm wide, round or occasionally deltoid at the apex, acuminate, the acumen
short, subfalcate, acute or more often acuminate, basally rounded, obtuse or
truncate, often vaguely inequilateral, the venation immersed or prominulous
above, prominulous beneath, the costa slender, the lateral veins 6-12, scarcely
arcuate, well spaced, stiffly papyraceous, glabrous above, glabrous to golden
pubescent beneath, the hairs curled, scattered along the veins, often tufted in the
axils; petioles short, to 1.5 cm long, glabrous or often golden villulose; stipules
soon deciduous, widely triangular or compressed hemispherical, 2-4 mm long,
acute. Infloreseences terminal and axillary, glabrate or minutely puberulent, the
peduncle absent or short, the branches 3-5, often lenticellate, terminated usually
by 3-flowered cymules or to single flowers; bracteoles linear subulate or ovate
subulate, several located at the base of the pedicel; bracts lanceolate or subulate;
pedicels short. Flowers large, showy, hypanthium oblong rotund or oblong, ca.
6 mm long, smooth, puberulent, the calycine cup ca. 1.5 mm long, glabrous
within, with patches of oblong glands at the junction of the calycine lobes, these
ca. 0.75 mm long, obtuse, the lobes 4-6, subulate, ca. 5.5 mm long, obtuse,
marginally pubescent; corolla obovate oblong, slightly falcate and thus zygo-
morphic, to 8 cm long, 1.8-3.0 cm wide, obtuse at the apex, attenuate acute at
the base, petaloid, scattered pubescent outside, puberulent within, the lobes 2.0-
2.5 cm long, obtuse or acute; stamens 6, the anthers linear oblong, 1.7-2.2 cm
long, ca. 1.5 mm wide, sagittate at the base, basifixed, the filaments free of the
corolla tube, ca. 7.5 cm long; style linear, 7.5-9.5 cm long, ca. 1.5 mm wide, the
stigmas 1.2-1.7 cm long, scarcely wider than the style, the ovary wall oblong in

cross section, with conspicuous ducts. Fruits capsular, oblong obovoid or ob-

ovoid rotund, 2.5-5.0(-7.0) cm long, 2.0-2.8 cm wide, strongly compressed, lig-
nose, bisculcate, dark brown, lustrous, glabrous, with numerous white, punctate
lenticels; seeds round or oblong, 2-15 mm long, brown, the body 2-5 mm long,

alate, the wing membranous, entire or undulate.

Coutarea hexandra is a common tree or shrub in the American tropics ex-
tending from Mexico to Argentina. The fruits of Mori et aL 4097 A are noteworthy
being ca. 3 times longer than wide, the longest to 7 cm long. Dr. Thomas Croat
(pers. comm.) notes that the flowers are bat pollinated. ''Nino Muerto'';

^^Quina.^^

BARRO COLORADO island: AvUes X-5 {U0)\ Croat 5165, 7354. H700 {jaW MO); Foster 1074 {U0)\
Gentry 1988 (MO); Zetek 3904 (MO), canal zonl: Pipeline Road, Croat 16683 (MO). Fort Amador,
D'An'y 9264 (MO). Island beyond Fort Amador, D'Arcy & D'Arey 6119 (MO). Ft. Kobbe, Duke

1980]

DWYHR— FLORA OF PANAMA (Family 179. Ruhiaceae) 137

4257 (MO). Farfan Beach, Dwyer 3067 (MO); Dwycr cl al. 457 (MO); Dwycr et al. 4693 (MO, SCZ).
Madden Dam, Dwyer & Haydcn 10 (MO). Farfan Beach, Dwycr <& Hayden 7524 (COL, DUKE, K,
MO, UC). Pipeline Road, Gentry 1447 (MO). Farfan Beach, Gentry 5501 (MO). Ancon Hill, Holdridge
6534h (MO). Farfan Beach, Lewis et al. (GH, K, MO, NY, UC, US); Tyson 1814 (MO), chiriqui:
San Felix to Cerro Flor, Allen 1953 (MO), cocll: Nueva Gorgona, D'Arcy & D'Arcy 6283 (MO). 5
mi S of Anton, Tyson & Blum 2572 (MO). La Vinta, Woodson et al. 1706 (MO), colon: 4 km NW
of Salamanca, 13 km NE of Buenos Aires, Nee 9067 (MO), darihn: Rio Tuqueza below Quebrada
Venado, Bristan 1078 (MO). Isla Saboga, Dnke 10357 (MO). Punta Garachine, Duke 10490 (MO). 1-
4 mi N of Pucro, Dnke 13028 (MO). Trail, Rio Pucro to Quebrada Maskia, Duke 13071 (MO).
HhRRHRA: Pese, Allen 796 (MO). Panama: Burunga, Austin 28 (MO). Morro Island, DArey &
D'Arcx 6785, 6786 (both MO). Nueva Gorgona, Duke 4532 (MO). E of Bejuco, Duke 4548 (MO).
El Llano, Duke 5864 (MO). Taboga Island, Dwyer 2807, 3067 (both MO). Cerro Azul, Gentry &
Dwyer 5508 (MO). Juan Diaz, Hunter 252 (MO). Playa de Venado, Mejia 2 (MO). 6 mi SW of
Arraijan, road to Bique. A^^^^^ 6898 (MO). Las Sabanas, A'/z/Vr 6757 (MO). Nuevo Emperador, vicinity
of Arraijan, Rivas de Hoyos 6 (MO). N of Chorrea, Tyson 5936 (MO). S of La Capitana, Tyson 6765
(MO). Chepo. Tyson 6776 (MO). Arraijan, Woodson et cd. 1386 (MO). Isla Taboga, Woodson et al.
1495 (MO). Pacora and Chepo, Woodson et al. 1668 (MO).

27. CRUSEA

Crusea Cham. & Schlecht., Linnaea 5: 165. 1830. type: C. rubra (Jacq.) Cham

& Schlecht.

Herbs, often perennial, the stems usually 4-angled. Leaves mostly lanceolate,
the lateral nerves conspicuous; petioles adnate to the stipular sheath, the latter
setiferous. Inflorescences capitate, surrounded by foliaceous bracts. Flowers
with the 4 calycine lobes alternating with minute glands, the calyx persistent in
fruit; corolla funnelform, glabrous within, the 4 lobes valvate; ovary 2-celled, the
ovules 1 per locule. Fruits capsular, didymous, the mericarps or cocci indehis-
cent, thin, the endocarp transparent, smooth, ecostate, the ventral face grooved,
narrow, the flanks of endosperm folded against the deep hilar ridge, the seeds 1
per locule, the testa cells contorted.

The genus Crusea, according to Anderson (1972), extends from '^Southern

Mexico through Me

ama."

Literature:

Anderson, W. R. 1972. A monograph of the genus Crusea (Rubiaceae). Mem.
N.Y. Bot. Garden 22(4): 128.

a. Corolla red, rose or violet; calycine cup 3-4 mm long; anthers ca. 3 mm long

1. C. coccinea var. chiriquiensis

aa. Corolla white; calycine cup less than 1 mm long; anthers ca. I mm long

2. C. longiflora

1. Crusea coccinea DC, Prodr. 4: 567. 1830, typii: Pavon, not seen.— Fig. 30.

la. Crusea coccinea DC. var. chiriquiensis W. R. Anderson, Mem. N.Y. Bot.
Garden 22: 45. 1972. type: Panama, Woodson & Schery 358 (F, holotype;
GH, MO, isotypes).

Upright or creeping herbs, stems angular, ribbed, glabrous, drying purple.
Leaves oblong or ovate oblong, 1.5-6.0 cm long, 0.5-2.0 cm wide, acute at the

138

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 67

FiGURh 30. Crusca coccinea var. chinquiensis W. R. Anders. — A. Habit (xVi). — B. Flower
(xL55).— C. Calyx (x3.5).— D. Ovary in longisection (x3.5). [After D'Arcy & D'Arcv 6519.]

1980] DWYER— FLORA OF PANAMA (Family 179. «HW«crrtc) 139

truncate

apex, occasionally distinctly acuminate, the acumen to 1.2 cm long, basally cu-
neate to acute, often slightly decurrent on the petioles, the costa slender, often
indistinguishable from the lateral veins, plane or immersed above, prominent
beneath, to 0.7 mm wide, the lateral veins usually 5, strongly arcuate ascending,
arising at a sharp angle, immersed or plane above, prominent beneath, charta-
ceous, not strongly discolorous, the upper surface striate, white hirtellous be-
tween the veins, white hirtellous beneath; petioles 0.5-0.8 cm long, to 0.1 cm
wide, these adnate to the lip of the stipular cup; stipules cyHndriform, to 1.5 cm
long, to 0.5 cm wide, membranous, glabrous or occasionally villosulose, the
sheath extending beyond the next superior node, with a terminal fringe of 1-12
glandular tipped linear setae, often hirtellous, to 1.4 mm long, ca. 0.2(-0.5) mm
wide. Inflorescences terminal, sessile, the flowers several, ascending, the hypan-
thiums hidden by the terminal stipules and the leaf bases. Flowers with the hy-
panthium turbinate, 3-5 mm long, ca. 2 mm wide at the
lycine cup cupuliform or campanulate, 3-4 mm long, ca. 3 mm wide at the apex,
glabrous, petaloid, the lobes 4, unequal, lanceolate, to 8 mm long, ca. 1.2 mm
wide at the base, acute, glabrous except hirsute marginally, usually with 2-3
small glands at the junction of 2 adjacent lobes, oblong, to 1 mm long, obtuse;
corolla red, rose or violet, to 35 mm long, the tube narrowly funnel shaped,
sharply attenuate below middle, to 0.3 cm wide near the mouth, glabrous within
and outside, the lobes 4, elliptic or ovate oblong, 8-1 1 mm long, to 5 mm wide,
with a few white hairs abaxially and terminally; stamens 4, exserted, the anthers
oblong, ca. 3 mm long, twisted after shedding pollen, versatile, the filaments
slender, plane, equal to the corolla lobes, to 7 mm long, attached near the mouth;
style filamentous, as long as the corolla tube, minutely puberulent on the upper
V3, the stigmas 2, slender, ca. 1.8 mm long. Fruits not seen.

Crusea coccinea var. chiriquiensis is known from Costa Rica and Panama.
Croat 15956, with only flower buds, is described on the label as "white.

^ ^

CHiRiQUi: Rio Chiriqui Viejo Valley, Allen 1369 (MO). Cerro Punta, Blum et aL 2426 (MO). NW
side of Cerro Pando cloud forest, Croat 15956 (MO). Cerro Respinga above Cerro Punta. D'Any
10709 (MO): D'Arcy & D'Arcy 6519 (MO). Cerro Punta, /?/V(t.'UY/y & Solis 2391 (MO, UC). Rfo
Chiriqui Viejo, Monte Lirio, Seihert 220 (MO). Casita Alta to Cerro Copete, Woodson & Schery 358
(MO).

2, Crusea longiflora (R. & S.) Anderson, Mem. New York Bot. Card. 22: 89. 1972.

Spcrmacoce longiflora Willd. ex R. & S., Syst. Veg. 3: 531. 1818. type: Mexico, Humboldt &

Bonpland (B-Willd., not seen).
Crusea hrachyphylla Schlecht. & Cham., Linnaea 5: 165. 1830. type: Schiedc & Deppe 532h (F,

isotype).
Spermacoce tenuiflora Ses. & Moc, Fl. Mex. 24. 1893. type: not seen.
Crusea chiapensis Brandegee, Univ. Cal, Publ. Bot. 6: 374. 1917. type: Mexico, Purpus 7526, not

seen.
Crusea hrachyphylla fo. microcephala Loes., Verh. Bot. Vereins Prov. Brandenburg 65: 115. 1923.

type: Seler & Seler 121 1, not seen.

Herbs to suhshruhs. 0.5-50.0 cm tall, branched or unbranched, the stems
slender, the hairs appressed, deflexed, the nodes well spaced. Leaves sessile,
lanceolate, occasionally ovate oblong, 0.5-3.0 cm long, 0.2-0.6 cm wide, acute
or deltoid at the apex, acute at the base, the costa slender, plane or immersed

140 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

above, prominulous beneath, the lateral veins (l-)2-5, strongly arcuate ascend-
ing, without a submarginal undulate vein, the intervenal areas smooth, the vein-
lets not evident, chartaceous to thinly succulent, concolorous, the hairs above
strigose, appressed, golden colored or white, subulate, 0.2-0.7 mm long, the hairs
beneath as above but mostly on costa and veins, the margin somewhat thick,
minutely ciliolate or with short, thick, ascending hairs; stipules to 4 mm long,
truncate, the setae 3-5, erect, the hairs stiff, to 1.7 mm long. Inflorescences
terminal, 0.5-4.0 cm in diam., the uppermost leaves involucrate, the stem im-
mediately below the involucre of leaves with numerous, weak, reflexed hairs.
Flowers subsessile, the hypanthium oblong, shorter than the calyx, minutely
puberulent; calycine cup 0.2-0.5 mm long, the lobes 4, subulate, erect, slightly
spreading, 1.6-3.2 mm long, usually slightly unequal, the hairs slender, stiff, to
0.8 mm long, with a minute gland in the sinus of 2 adjacent lobes; corolla white,
the tube slender, 0.6-0.7 cm long, ca. 0.2 mm wide, minutely puberulent to
glabrous outside, glabrous within, the lobes 4, oblong, 1.0-2.2 mm long, obtuse;
stamens 4, exserted, the anthers oblong, ca. 1 mm long, the filaments threadlike,
to 1.8 mm long, attached at the mouth; style filamentous, about as long as the
tube, the stigmas 2, 0.3-1.0 mm long, slightly wider than the style. Fruits not

seen.

Crusea longiflora extends from Mexico to Panama. Superficially the two Pan-
amanian collections seem to belong to different species but floral dissections
indicate that they are conspecific. On the dwarf plants of Davidse & DArcy
10348, the leaf blades show a reduction of lateral veins to 1-2 pairs,

CHiRiQuf: Large old lava flow, 3 km NE of El Hato del Volcan, base of Volcan de Chiriqui, 1-

3 km E of highway, 1500-1800 m, Davidse & D'Arcy 10348 (MO). Volcan Chiriqui above Boquete,
roadside, DAnx 9793 (MO).

28. DECLIEUXIA

Declieuxia H.B.K.,^^ Nov. Gen. Sp. PI. folio 3: 275, tab. 28L 1819; quarto 3
352, tab. 28L 1820. type: Z). chiococcoldcs H.B.K.

Herbs or subshrubs, sometimes cricoid in habit. Leaves occasionally verti-

cillate, often coriaceous, frequently sessile; stipules interpetiolar, often subulate.
Inflorescences terminal or axillary, cymose, usually branching dichotomously, or
spikelike and racemiform; bracts often absent. Flowers with the calycine cup
reduced, the lobes usually 4, equal or unequal; corolla variable in color, infun-
dibuliform or cylindrical, the lobes 4, valvate; ovary 2-celled, the ovules 1 per
cell, attached basally. Fruits a schizocarp, at first fleshy, didymous, indehiscent,
compressed, the thin endocarp usually hard; seeds compressed.

Declieuxia ranges from Mexico, through Central America, the West Indies,
and South America. There are about 40 species in the genus.

^^ For synonymy see Kirkbride (1976).

1980] DWYER— FLORA OF PANAMA (Family 179. Riihiaccae) 141

Literature:

Kirkbride, J. H., Jr. 1976. A revision
New York Bot. Card. 28(4): 1-87.

Mem

Willd

Fig. 31.

Hoiistoniafniticosa Willd. exRoem. & Schult.,-'''Syst. Veg. 3: 527. 1818. type: Venezuela. Hiimhohlt
Si Bonpland 223 (B-Willd., holotype; P, isotype, neither seen).

Herbs, woody, to 3.5 m tall, the branchlets arising from the base or above,
rarely restricted terminally, smooth, glabrous, delicately alate, the nodes well
spaced. Leaves sessile, occasionally 3 per node, lanceolate, occasionally slightly
falcate, to 5 cm long, to 1.2 cm wide, acute at the apex, obtuse to acute at the
base, the costa prominulous above and beneath, the lateral veins ca. 5, arcuate,
prominulous above and beneath, the intervenal areas reticulate, usually concol-
orous, lustrous above, marginate; stipules deciduous. Inflorescences terminal,
cymose paniculate, a peduncle like axis (really a primary cyme) often present, to
2 cm long, or the stem tip with 3 primary cymes, each with an axis terminated
by a sessile flower and 2 lateral branches, these branching into small cymules,
the more profusely branched inflorescences often flat topped; bracteoles tending
to persist, subulate. Flowers with the hypanthium subrotund, truncate, ca. 0.8
mm long, glabrous, the calycine teeth 4, erect, linear oblong, ca. 0.8 mm long,
obtuse, glabrous; corolla white, the tube cylindrical, ca. 6 mm long, petaloid,
densely villose below the middle within, the lobes oblong, to 2 mm long, glabrous,
not cucuUate; stamens 4, the anthers exserted, oblong, to 1 mm long, dorsifixed,
the filaments to 1.5 mm long, attached in the middle of the tube or below; ovarian
disc bilobed, pulvinate, the stigmas 2, slender, to 0.3 mm long, the style slender,
to 2.5 mm long, attenuate basally, the ovary irregularly ribbed, 2-carpellate. Fruits
sessile, rotund and ca. 2 mm in diam. or bilobed (2 carpels developing) and then
wider than long, drying black, marcescent, often reticulate, glabrous, usually
marginate; bracteoles persistent, subulate.

Declieuxia fniticosa ranges from Mexico south to Brazil.

CHiRiguf: San Felix to Cerro Flor, 100-800 m, Allen 1930 (MO). Savanna below Boquete, ca.
800 m, D'Arcv 9773 (MO). 2 mi S of Boquete, 3-4000 ft, D'Arcy & D'Arcy 6309 (MO). Boquete,
4000 ft, Davidson 701 (MO); Dnycr & Haydcn 7593 (MO); Partch 69-80. 69-128 (both MO). Boquete,
Llanos Francia, Stern et al. 1152. 1952 (both MO). Boquete. 1200-1500 m, Woodson & Sclwry 791
(MO), cocle: Nata, Allen 827 (MO). Santa Clara Beach, Croat 9588 (MO). Penonome, Ehinger
(MO). PANAMA: Taboga Island, Allen 118. 1271 (both MO). Las Uvas, Bartlett & Lasser 16650 (MO).
Hacienda La Joya, Dodge et al. 16897 (MO). Savanna near Rio Pacora and Chepo Highway. Duke
5898 (MO). Savannalike areas, road to Cerro Campana, Duke 6029 (MO). Forest next to grasslands.
Cerro Campana, 2400-2700 ft. Duke 8701 (MO). Tocumen Airport, D^xyer 1251. 4428 (both MO).
Cerro Campana, Dwxer et al. 4847 (MO, SCZ, UC); Dnwer & Kirkbride 7824 (MO). Nuevo Arraijan,
Guerra 24 (MO). Cerro Campana, Lcnis et al. 3100 (COL, DUKE, K. MO, UC, VEN). Taboga
Island, Lon^ield 489 (MO). Cerro Campana, 2600 ft, MvDaniel 6824 (MO); Nee 6872 (MO); Witli-
erspoon & Witherspoon 8365 (MO). Taboga Island, Woodson et al. 1467 (MO), veraguas: Yeguada,

38 For synonymy see Kirkbride (1976) p. 17.

142

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 67

Figure 31. Dedieuxia fruticosa (R. & S.) Kuntze.— A. Habit (x.6)— B. Flower, opened to
show interior (x6.0). [After Allen 1271.]

1980] DWYER— FLORA OF PANAMA (Family 179. /?///;/«(■('«(') 143

Hladik 383 (MO). San Jose on Santiago-Santa Fe Road, 18 km S of Santa Fe, 400 m, Mcc 8180 (MO).
Canazas, Tyson 3740 (MO).

29. DEPPEA

Deppea Schlecht. & Cham., Linnaea 5: 167. 1830. type: D. erythrorhiza Cham

& Schlecht.

Choristes Benth., PI. Hartw. 63. 1840. type: C. ohtusifolia Benth.

Infl.

minal or axillary, the flowers rarely solitary, disposed in cymes or umbels, ebrac-
teolate. Flowers pedicellate; calyx 4-lobed, the lobes persistent; corolla rotate,
the tube often shorter than the lobes, glabrous within, the lobes 4, contorted;
stamens 4, exserted, the filaments short, attached near the corolla mouth; style
slender, the stigmas capitate or the 2 lobes evident, the ovary 2-celled, the ovules
numerous, imbricate, the placentas peltately adnate to the septum. Fruits cap-
sular, small, turbinate or obovoid, scarious to coriaceous, costate, loculicidally
dehiscent from the apex; seeds numerous, subglobose, minute, obtusely angular,
the testa hard, granulate.

Deppea ranges from Mexico to Venezuela. The genus includes about 20
species.

a. Flowers with hypanthium sparsely villose: fruits turbinate, 0.4-0.5 cm long. 8-costate; high

mountain species, 1500-4500 m 1. D. grandifolia

aa. Flowers with hypanthium glabrous; fruits rotund, to 0.2 cm long, 12-costate; low elevation

species 2. D. panamcnsis

1. Deppea grandiflora Schlecht., Linnaea 19: 748. 1847. syntypes: Mexico, Lei-
bold 235, 253, 263, none seen. — Fig. 32.

D. coslaricensls Polak., Linnaea 41: 566. 1877. type: Costa Rica, Polakowski 148 (not seen: photo,

F).

D.floribundu Hemsl., Diag. PI. Nov. 31. 1879. type: Bottcri 937 (not seen; P, holotype; photo, MO).

D. longipes Standley, Contr. U.S. Natl. Herb. 18: 138. 1916. type: Panama, Pitticr 3070 (US, ho-
lotype).

Trees 3-8 m tall, the branchlets terete, smooth, glabrous to puberulent. Leaves
oblong, oblong rotund or oval, 6-17 cm long, 2.0-6.5 cm wide, deltoid at the
apex, acuminate, the acumen to 1.5 cm long, acute, cuneate to acute at the base,
the costa plane above, prominulous beneath, the lateral veins 8-12, arcuate,
prominulous above and beneath, chartaceous, scarcely bicolorous, glabrous
above except on veins, appressed villose on the veins beneath, the hairs often
tufted in the vein axils; petioles to 4.5 cm long, to 0.2 cm wide, wiry, often
arcuate and deflexed, densely pubescent; stipules deciduous, minute, broadly
deltoid and glanduliferous (fide Standley). Inflorescences terminal or pseudo ter-
minal, occasionally longer than the uppermost leaves, paniculate, the peduncles
3_4 cm long, villosulose, terminated by 3 primary branches, these to 2.5 cm long,
strongly ascending, in turn branching, ultimately producing cymes. Flowers with
the hypanthium oblong or turbinate, 1.6-2.0 mm long, ribbed, sparsely villose,
the calyx cup scarcely measurable, the teeth reduced to 5 points, 0.2-0.6 mm

144

ANNALS OF THi: MISSOURI BOTANICAL GARDIiN

[Vol. 67

FiGURF 32. Dcppca grandiflora Schlecht.— A. Fruiting branch (x'/i). [After Tyson & Loftin
5964. ]_B. Inflorescence in bud (xi/2). [After Wilbur 10291. \

long, acute; corolla yellow, the tube compressed cylindrical, ca. 2 mm long,
usually wider than long, petaloid, glabrous, the lobes 4, imbricate, oblong, 3.5-
5.0 mm long, ca. 4 mm wide, longer than the corolla tube, venose, glabrous within
except granulose puberulent in the middle; stamens ?4, the anthers oblong, 3.0-
4.2 mm long, ca. 0.6 mm wide, basally bilobed, the filaments to 3 mm long,
attached at the apex of the tube; ovarian disc platelike, the stigmas 2, clavate or
rotund, ca. 1.3 mm long, the style slender, to 7 mm long, the ovary oblong, the
wall thin, the ovules numerous on intrusive placentas. Fruits turbinate, 0.4-0.5
cm long, ca, 0.35 cm wide at apex, truncate, the calycine cup thickened and
persistent as a rim, smooth, glabrous, 8-costate, the lateral split not extending
below the terminal ringlike rim.

Dcppea grandiflora ranges from Mexico to Panama.

1980] DWYER— FLORA OF PANAMA (Family 179. RubUiceac) 145

CHiRiQuf: Boquete, Chiquero, Davidson 541 (MO). Volcan de Chiriqui, Davidson 1049 (MO).

Camp Aguacatal, Chiriqui' Volcano, 2100-2200 m, Pitticr 3070 (MO). CerroPunta toward Bajo Grande
Wilbur et al. 10921 (MO).

2. Deppea panamensis Dwyer,^** spec. nov. type: Panama, Mori & Kalliinki 4965
(MO, holotype).

Trees to 8 m, the branches numerous, the branchlets often drying yellow gray,
the bark rough, cracking, glabrous, the nodes crowded at the apex of the twigs.
Leaves lanceolate or obovate oblong, 8-15 cm long, 2.5-5.0 cm wide, obtuse,
deltoid or subobtuse at the apex, acuminate, the acumen to 1 cm long, acute to
attenuate acute at the base, the costa prominulous above and beneath, often
drying dull red beneath, the lateral veins 9-11, prominulous, arcuate, the sub-
marginal undulate vein well set in from the margin, the intervenal areas smooth,
mosaiclike under magnification, the veinlets immersed, stiffly chartaceous, con-
colorous, glabrate above, minutely puberulent on the costa and the veins beneath,
minutely white barbate in the leaf axils; petioles slender to 2.5 cm long; stipules
apparently connate, cylindrical, 2.5-3.5 mm long, truncate or each part with a
notch extending to the middle, coriaceous, minutely puberulent. Inflorescences
terminal, pyramidal paniculate, puberulent, to 10 cm long, 8-10 cm wide, the
peduncle suppressed, 3 branches arising from the twiglet tip, the middle one
projecting upward as a rachis, or with the peduncle 2-4 cm long, slender, deli-
cately winged, to 1.2 mm wide, bearing several well-spaced branches along its
length, at least those below the middle of the rachis opposite, the uppermost
often alternate, the inferior branches to 10 cm long, branching, the secondary
branches producing several few-flowered cymules, or occasionally the flowers
solitary. Flowers not seen except for the hypanthium, calyx and ovarian disc,
these measuring ca. 1.5 mm long, glabrous; hypanthium turbinate, the calycine
cup scarcely measurable, the teeth ca. 0.15 mm long. Fruits pedicellate, rotund,
to 2 mm long, glabrous, shiny, loculicidally dehiscent almost to the base, the 4
resulting segments each with 3 delicate ribs, venose, the persistent ovarian disc
in the unopened fruit moderately prominent, umbonate; seeds oblong or trape-
ziform, to 0.25 mm long, somewhat piano compressed, verrucose, honey colored.

Deppea panamensis is known only from Panama. It is distinguished by its
minute fruit which separates to the base into 4 parts.

Panama; 5-10 km NE Altos de Pacora, Mori & Kallnnki 4965 (MO).

30. DIDYMAEA

Didymaea Hook, f., Benth. & Hook, f., Gen. Pi. 2(1): 150. 1873. type: D, mex-
icana Hook, f.

Herbs, the stems weak, 4-angled. Leaves small, petiolate; stipules connate,
subulate, persistent or deciduous. Inflorescences axillary, the pedicels often elon-

"^^^ Deppea panamensis Dwyer, spec. nov. Arhores ad 8 m. Folia lanceolata vel obovato-oblonga,
8-15 cm longa, 2.5-5.0 cm lata, acumine ad 1 cm longo. venis lateralibus 9-11, chartacea, in venis
et costa subtus puberula; petiolis ad 2.5 cm longis; stipulis fortasse connatis vagina cylindrica truncata
vel relusa. Infloresentiae terminales ad 10 cm longae, 8-10 cm latae. Florcs hypanthio turbinato
dentibus calycis ca. 0.15 mm longis. Frnctus rotundi, ad 2 mm longi, glabri.

146 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

gating in fruit. Flowers with the hypanthium turbinate globose, the calycine cup
truncate; corolla campanulate or rotate, the lobes 4, valvate; ovary 2-celled, the
style thick; ovules 1 per cell, attached laterally to the septum. Fruits didymous,
fleshy, globose, one lobe often abortive.

Didymaca is a genus of about 5 species. The genus has been poorly collected.
The reader is referred to Standley & Williams (1975) where 2 new species are

described and the genus is discussed in some detail.

Literature:

Standley, P. C. & L. O. Williams. 1975. Didymaca Hooker f. pro parte, Flora
of Guatemala. Fieldiana: Botany 24: 60-63.

1. Didymaea alsinoides (Schlecht. & Cham.) Standley, Field Mus. Nat. Hist.,
Bot. Ser. 18: 1291. 1938.

Ncrtcra alsimndes Schlecht. & Cham., Linnaea 6: 413. 1831. type: not indicated.

Didymaca alsinoides var. aitstralis Standley, Field Mus. Nat. Hist., Bot. Ser. 18: 1292. 1938. type:

" Costa Rica, Brade & Brade 2143 (F, holotype).
Didymaca alsinoides var. mollis Standlev, Field Mus. Nat. Hist., Bot. Ser. 18: 1292. 1938. ty

Costa Rica, Standley & Valerio 43949 (US, holotype).

Herbs, slender, fragile, the branches 4-angled, delicately porcate, the grooves
densely hairy, the nodes well spaced. Leaves oy^X^ or oblong ovate, 1.0-3.5 cm
long, 0.5-1.5 cm wide, acute at the apex, rounded or obtuse at the base, occa-
sionally truncate, usually inequilateral, with 1-2 pairs of veins arising from the
base or with 1 pair of basal veins and 1 pair along the costa, membranous, con-
colorous, densely villose above and beneath; pedicels to 1 cm long, pubescent;
stipules free, linear oblong, to 2.5 mm long, curled. Pedicels to 2 mm long.
Flowers with the hypanthium ca. 0.5 mm long, glabrous, the raphides scattered,
the calycine cup 0.2 mm long, the lobes 4, subtriangular, ca. 1 mm long with a
small gland at the junction of 2 calycine lobes, the remaining floral parts not seen.
Fruits didymous, oblong reniform, to 0.65 cm long, drying black, lustrous, pu-
bescent; pedicels to 2 cm long.

Didymaea alsinoides is known only from Costa Rica and Panama.

In the material examined I found it possible to separate glabrous fruited ma-
terial {Cochrane et aL 6290; Croat 13674) from non-glabrous {Croat 13726\
DArcy 9996), thus seemingly supporting Standley 's two varieties: var. alsinoides
and var. australis. But Davidson 395, cited by Standley under var. australis, has
some fruits sparsely hairy and others seemingly glabrous. In speaking of the
undersurface of the blades of var. australis Standley says that they are ''subtus
glabra vel ad nervos hispidula." The leaves of Davidson 395 are hispidulous
beneath, the hairs not being restricted to the costa and veins. In other respects

the leaves and fruits of all the material cited above are so similar that I see no
reason for maintaining the two varieties.

CHiRiQDf: Cerro Horquela, cloud forest, 1750-2100 m. Cochrane ct aL 6290 (MO). Las Cumbres
Cerro Punta, Croat 13674, 13726 (both MO). Alio Respinga and above, ca. 2800 m, D'Arcy 9996
(MO). Bajo Chorro, Boquete, 6000 ft, Davidson 395 (MO).

1980J DWYER— FLORA OF PANAMA (Family !79. Ruhiaceae) 147

31. DIDYMOCHLAMYS
Didymochlamys Hook, f., Icon. PI./?/. 1122. 1872. type: D. whitei Hook. f.

Epiphytic herbs. Leaves sessile or subsessile, alternate, anisophyllous, pre-
sumably distichous, the lateral veins evanescent; stipules small, dimorphic, sub-
ulate or linear lanceolate, apparently not occupying a fixed position with relation
to the petiole attachment. Inflorescences terminal, pedunculate, the latter ter-
minated by 2 large subrotund bracts enclosing 2 or more flowers. Flowers with
the calycine lobes 5, subulate, erect; corolla funnelform, pubescent at the base
within near the filament attachment point, the lobes themselves lobed basally;
stamens 5, unequal, the anthers ovate or oblong, apiculate, the filaments attached
near the base of the tube; ovary 2-celled, the placentas with numerous ovules,
arising from the base of the septum. Fruits capsular, laterally compressed,
2-celled, dehiscing at the apex, the placentas attached basally to the septum-
seeds numerous, minute, hairy, the hairs tufted at 1 point on the seed.

Didymochlamys is known from Panama, Colombia and Venezuela. There are
two species in the genus.

Literature:

Hooker, J. D. 1872. Didymochlamys whitei Hook. f. in Hooker's Icones Plan-

tarum 21-22, /?/. 1122, London.

1. Didymochlamys connellii N. E. Brown, Trans. Linn. Soc. London, Bot, 6: 34.

pL 5. fig. 1-9. 1901. type: British Guiana- Venezuela border, McConnell &

m +

Quelch 714, not seen. — Fig. 33.

Epiphytic herbs, the stems glabrous, minutely lenticellate, to 20 cm tall.
Leaves sessile or subsessile, alternate, anisophyllous, the larger blades disti-
chous, narrowly falcate oblong, occasionally vaguely narrow trapeziform, 4.5-
10.0 cm long, 0.8-1.7 cm wide, the smaller blades narrowly subulate, to 10 mm
long, occupying varying positions on the stem, the blades of the larger leaves
with the costa plane beneath, to 1.5 mm wide, the lateral veins not visible (made
visible by soaking), ca. 8, arcuate, forming a distinct, undulate, submarginal vein
well in from the margin, with a slender vein paralleling the margin ca. 0.1 mm
from the margin, chartaceous, concolorous, glabrous, smooth, minutely silver
scaly under a lens, the margin vaguely revolute; stipules triangular, ca. 1 mm
long, often deciduous. Inflorescences terminal, to 3.5 cm long, the peduncle 1.0-
1.5 cm long, terminated by 2 large, subrotund bracts, to 1.5 cm wide, minutely
acuminate, glabrous, membranous, replete with cystolilhs, blue violet or some-
times green, enclosing 2 flowers, the bracteoles several, subulate, unequal, short-
er than the calyx. Flowers at anthesis exceeding the bracts, sessile, the hypan-

thium to 2 mm long, to 1.8 mm wide, glabrous, the calycine cup absent, the lobes
5, subulate, subequal, ca. 1 mm long, glabrous; corolla bluish, the lobes white,
to 1.8 cm long, the tube funnelform, to 5 mm wide above the middle, to 2 mm
wide below the middle, glabrous outside, a few hairs within below the filament
attachment, the lobes 5, the basal portion of each lobe ovate oblong, to 2 mm

148

ANNALS OF THE MISSOURI BOTANICAL GARDEN

IVol. 67

Figure 33. Didymochlamys conncUii N. E. Brown. Habit (x-34). [After Forero & JaramiUo
2647^ Colombia.]

1980] DWYER— FLORA OF PANAMA (Family 179. /?(//?/«cTrt(') 149

long, with 2 lateral lobes, oblong, 2.5-3.0 mm long, infolded in the bud, curled
and flaring at anthesis; stamens 5, 2 being slightly shorter, not exserted, the
anthers oblong or ovate oblong, ca. 1.7 mm long, obtuse and minutely retuse, the
filaments with the free portion longer than the anther; ovarian disc 2 or 4 lobed,
the style slender, ca. 9 mm long, glabrous, the stigmas 2, ca. 1 mm long. Fruits
not seen.

Didymochlamys conncllii is known only from Guyana, Venezuela, and Pan-
ama. My description states that the leaves are anisophyllous, which appears to be
the case. The larger blades are obviously distichous. The smaller subulate leaves
have an intricate phyllotaxy. The more frequent situation is to have smaller leaves
in a non-opposite position, but occasionally they are directly opposite the larger
blade and like the latter have 2 stipules at the base, the minute stipules being the
counterparts of the stipules subtending the larger blades. Whether this alternate
arrangement fits the Fibonnaci series pattern has to be determined.

In Brown's original description of D. connellii, he states: "Leaves alternate
. . , stipule solitary, 3-4 lin. long, linear." Hooker described the only other known
species of Didymochlamys, D. nhitci Hook, f., from Colombia. This species is
based on a collection from "the dense woods at Quibdo, on the river Atrato, an
affluent of the Magdalena, growing on stumps of trees, R. B. White Esq/' Hooker
stated (my translation) that the stipules are well set apart and stipules like the
leaves are seemingly remote; some membranous, located on the dorsal surface
of the prostrate stem, linear lanceolate, entire or 2-fid (or 2-nate), the petiole
remote or close [to the stipule]; the others coriaceous or horny when dry, minute,
clawed, almost axillary. Hooker's plate #1122 is certainly that of DidymocMa-
mys\ figure 1 in the plate shows the base of what I designate as a "large blade"
and directly opposite 2 subulate structures which I designate as "small blades."
It is not clear whether the artist has shown the triangular and minute stipules
which I assume are present in D. whitei and which Hooker described in his

diagnosis.

Steyermark (1974) says: "Estipulas solitarias o en pares con dos formas; una

axilar en la base del peciolo, diminuta y en forma de garra, la otra solitaria o en
pares, alargada, linear o linear-lanceolada, entera o bifida." While this considers
what I call a small blade to be a stipule, it is much clearer than Hooker's de-
scription. The sketch in Steyermark's work is in general very good except that
the subulate stipules are shown opposite the large leaves with the small leaves
lacking stipules, a feature subject to question.

A critical point in determining terminology is that the small leaves may them-
selves be subtended by minute stipules; therefore, I regard the awl shaped struc-
tures as leaves and not as stipules. In fairness to the argument, the small leaf

lacks venation.

In the Dressier collection I found no evidence of the small leaf or so-called
stipule being bifid. In soaking the stem of Dressler's collection in a 2% technical
detergent (Aerosol OT) solution, 1 found that the small leaf or stipule turned
white in contrast to all the other vegetative parts. This is of uncertain significance.

cocle: Aserradero El Cope, 8 km N of El Cope. ca. 900 m, Dressier 5640 (MO).

150 ANNALS OF THH MISSOURI BOTANICAL GARDKN [Vol. 67

32. DIODIA
Diodia L., Sp. PI. 1: 104. 1753; Gen. PI., ed. 5. 45. 1754. type: D. virginiana L.

Annual or perennial herbs, the stems terete or 4-angled. Leaves opposite or
seemingly verticillate, sessile or short petiolate; stipules connate, united with the
petioles, the sheath filiform setose, the processes often ciliate. Inflorescences
axillary or in terminal spikes, the capitate heads borne in the axils of reduced
leaves. Flowers sessile, usually 4-merous, the calycine tube 2-6-lobed, often with
small interdental glands; corolla infundibuliform to hippocrateriform, the lobes
4-6; stamens 4-6, the anthers exserted, dorsifixed, the filaments attached at the
mouth; ovary mostly 2-celled, the ovules solitary, adnate to the septum in the
middle. Fruits partly superior, the 2 cocci indehiscent, the endocarp thick and
woody, smooth or costate, the ventral face plane, the endosperm relatively re-
mote from the hilar ridge.

Diodia occurs in Mexico, Central America, the Antilles, and South America.
Some species are found in the United States and in Africa. The genus has about
50 species.

Literature:

Steyermark, J. 1972. The Botany of the Guayana Highland — Part IX. Mem
N.Y. Bot. Garden 23: 788-805.

a. Leaves 1.2-3.0 cm wide.

b. Inflorescences flagelliform in appearance and with obviously reduced leaves or bracts;

leaves with lateral veins not deeply immersed above 1. D. dcnudata

bb. Inflorescences not flagelliform, not with obviously reduced leaves or bracts; leaves

with lateral veins obviously immersed beneath 2. D. sarmcntosa

aa. Leaves 0.2-0.4(-0.8) cm wide 3. D. teres

1. Diodia denudata Standley, J. Wash. Acad. Sci. 15: 105. 1925. type: Panama,
StancUey 28123 (US, holotype).— Fig. 34.

Vines, subshrubs or shrubs, the stems spreading, the nodes usually well
spaced, the short axillary branchlets often simulating crowded leaves, glabrous,
delicately alate on the angles. Leaves lanceolate or obovate-lanceolate, 4.0-7.5
cm long, 2-3 cm wide, acute at the apex, acute to attenuate acute at the base,
often inequilateral, the costa immersed or plane above, prominulous to subplane
beneath, the lateral veins ca. 7, strongly arcuate, ascending with 1 intermediate
vein occasionally found between 2 laterals, papyraceous, scarcely discolorous,
smooth when dry, minutely puberulent or scabridulous on costa and veins be-
neath, glabrate above; petioles slender, to 1.5 cm long or sometimes the blades
lightly decurrent; stipules triangular or compressed rotund, the sheath shorter
than the setas, the latter unequal puberulent. Inflorescences axillary, capitate, to
0.4 cm wide, the heads arranged along the length of a whiplike branch, the latter
to 45 cm long, the internodes elongate, the subtending leaves ?bracts lanceolate,
0.5-2.0 cm long. Flowers minute; hypanthium ca. 0.3 mm long, the calycine cup
minute, glabrous, the lobes 4, sometimes 1 or 2 of these about Vi the length of
the longer lobes, ovate subulate, to 0.4 mm long, scabridulous ciliate; corolla

1 980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

151

JL^

FiGURL 34. Diodia denudata Standley. — A. Habit {y-Vi). — B. Fruit in ventral and dorsal views
(xI2'/2). [After Croat 13191,]

152 ANNALS OF THE MISSOURI BOTANICAL GARDEN |Vol. 67

white, the tube ca. 0.5 mm long, the lobes 4, about equaling the tube; stamens
4, attached at the mouth; style ca. 0.5 mm long, the stigmas to 0.3 mm long.
Fruits sessile, subrotund, to 1.3 mm long, often slightly wider than long, with a
distinct median furrow, scabridulous, delicately red farinose or delicately retic-
ulate, otherwise smooth, each valve indehiscent, the calyx persistent; seed ob-
long, to 0.8 cm long.

Diodia denudata is known only from Panama and Colombia.

BARRO COLORADO ISLAND: AvUc's 94 (MO); Croat 4373. ^738 (both MO), canal zonk: Rio
Mcndosa, Vi km downstream from Pipeline Road, Nee 9577 (MO). Fort Lorenzo, Piper 5895 (US).
Rio Tapia, StamUey 28123 (MO. US), col on: Santa Rita Ridge, Croat 13191 (MO); Lewis et at. 5238
(MO), darikn: Santa Fe, Duke 9488 (MO). Panama: Rio Tocumen, StamUey 29364 (US). Colombia:
CHOco: Punta Las Barcas and Turbo, Duke 9695 (MO). Madure.x Logging Camp, Duke 9977 (MO).

2. Diodia sarmentosa Swartz, Prodr. Veg. Ind. Occ. 30. 1788. type: Jamaica,
Snartz, not seen.

Herbs or shrubs, often elongate, branching freely, obtusely angular, puberu-
lent, the nodes well spaced. Leaves sessile or subsessile, elliptic oblong or ovate
oblong, 3-5 cm long, 1.2-1.8 cm wide, acute to deltoid at the apex, often vaguely
acuminate, cuneate at the base, the costa immersed above, prominulous beneath,
the lateral veins 4-5, strongly arcuate ascending, immersed above, subplane to
prominent beneath, the intervenal areas generally smooth beneath, the veinlets
evanescent, the margin revolute, stiffly chartaceous, concolorous but appearing
bullate above and rough to the touch, scabridulous above and beneath, the hairs
above short, white, calcareous punctate, strigose, the hairs beneath more slender
and somewhat longer; stipules with sheath turgid, stiff, to 3 mm long, the setas
subulate, slender, to 7 mm long, pubescent. Inflorescences in the axils along the
length of the branches, the flowers congested, forming a spikelike mass ca. 0.5
cm in diam. Flowers not seen. Fruits oblong or turbinate, 5 mm long, truncate,
smooth, glabrate to puberulent, often lightly punctate and bearing a few faint
longitudinal lines, the calyx persistent or not, the teeth ovate triangular, Vi the
length of the valves, the inner face of valve like a drum head, white, stiffly
membranous, with a delicate median vein.

Diodia sarmentosa is a common weed in the West Indies, throughout Middle
America, and northern South America. It also occurs in the Pacific islands, the
Malaya Peninsula and tropical Africa. Steyermark (1972) in his key to Diodia
stresses several diagnostic floral characters: "corolla 2-5 mm long . . . style sub-
capitate, shortly bilobate . . . calyx lobes linear-lanceolate, 1.0-2.2 mm long."

canal zone: Coco Solo, Dwyer & Duke 7919 (COL, MO, UC). barro Colorado island: Croat
5129, 5276. 5525 (all MO), vlraguas: Mouth of Rio Concepcion, Lewis et aL 2781 (MO).

3. Diodia teres Walt., Fl. Carol. 87. 1788. type: "In sabulosis sylvarum a Virginia
ad Carolinam" Walter (BM, not seen).

D. prostrata Swartz, Prodr. Veg. Ind. Occ. 30. 1788. i ype: not ;

Herbs, often woody at the base, prostrate and forming mats, or erect and to
0.3 m tall, the branches terete or angular, delicately 4-ribbed, white villose, the

19S0] DWYER— FLORA OF PANAMA (Family 179. Riihiaceae) 153

hairs to 1.2 mm long, with smaller crinkly hairs interspersed. Leaves sessile,
spreading, occasionally deflexed, subulate, narrowly oblong or narrowly ovate
oblong, 1.5-2.5 cm long, 0.2-0.4(-0.8) cm wide, subequilateral, tapering acutely
at the apex, the terminal seta to 1.5 mm long, white, basally obtuse or truncate,
the costa subimmersed above, prominulous beneath, the lateral veins evanescent,
stiffly papyraceous, white punctate and hispid above and beneath, the hairs to 2
mm long, erect or curled, the margin frequently revolute, scabridulous, the hairs
usually associated with marginal setas; stipules connate, cuplike, the setas prom-
inent, stiff, ascending, linear, ca. 10, subequal, to 6(-7) mm long, usually 2-3
times the length of the sheath, the latter with few, elongate, stiff, reflexed, white
hairs, these usually associated with the marginal setas, the hairs at the base of
the sheath absent, or if present, shorter and weaker. Inflorescences axillary, the
flowers few. Flowers with the hypanthium oblong, to 1.8 mm long, truncate,
densely hirsute, the calycine cup scarcely measurable, the teeth 4, unequal, sub-
ulate, to 2 mm long, acute, petaloid, densely ciliate with 1-2 glandular, subulate
structures, ca. 0.1 mm long in the sutures; corolla white or pink, the tube cam-
panulate, to 5 mm long, petaloid, glabrous within except for a ring of hairs ca,
1.5 mm from the base of the tube, with scattered uncinate hairs outside, the lobes
4, oblong or deltoid, to 2 mm long, acute; stamens 4, the anthers oblong, to 1.2
mm long, obtuse, dorsifixed or subversatile, the filaments short, attached just
below the middle of the throat; stigmas 2, the style filamentous, to 7 mm long,
the ovary thick walled. Fruits sessile, indehiscent, of 2 mericarps, turbinate, ca.
3 mm long at maturity, hispid; seeds red brown, reticulate, the calyx deciduous.

Diodla teres is common throughout Middle America and South America. It
is found in the United States, ranging from Connecticut south to Florida and
Texas and westward to Oklahoma.

CHiRiQL'i: CeiTO Horqueta. ca. 5000 ft. Dwyer et al. 458 (GH. MO. US), cocle: Nata. Allen 835
(MO). E of Nala, Duke 12384 (MO). Penonome, Ehnif^cr 1006 (MO), herrera: Las Minas and Pese,
Btifch et aL 1323 (F, GH, K, MO, NY, UC, US); Duke 12301 (MO). Las Minas, La Pena, Stern et
al. 1778 (MO). Ocu, Tyson et al. 2857 (MO). V2 mi N of Macaras, Ri'o La Villa. Tys(m el al. 3131
(MO, SCZ). VKRAGUAs: 5 mi NE of La Mesa, Blum & Tyson 657 (MO). 10 mi W of Santiago, Duke
13658 (MO). Isla de Coiba, Duyer 1675 (MO). 2 mi W of Santiago, Dnjer et al. 7535 (GH, MO, US).
50 km E of Santiago, Folsoni 3911 (MO). 1-2 mi above Santa Fe, Gentry 3047 (MO). V2 km W of
Highway 50, turnoff to Ocii, Nee 8001, 8009 (both MO). Just S of Santa Fe, 450 m. Nee 8026 (MO).

33. DRESSLERIOPSIS

Dressleriopsis Dwyer, gen. nov.^^ type: D. panamensis Dwyer.

Suhs/iruhs, Leaves often cordate toward the base, petiolate; stipules free,
triangular. Inflorescences axillary, the flowers disposed in globose heads, the
peduncle absent, the bracteoles subulate. Flowers subsessile, the calycine cup

^' Dressleriopsis Dwyer, gen. nov, Suffrudees. Folia saepe versus basin cordata petiolata; stipulis
liberis triangularibus. Infloreseentiae axillares floribus in caespites globosas dispositis, pedunculo
nullo; bracteolis subulatis. Flores subsessiles cupula calycina intus glanduHs ornata lobis 4; corolla
tubo cylindrico intus glabro lobis 5; staminibus 5 antheris oblongis filamentis brevibus; disco ovarii
pulviniformi: stylo gracile; stigmatibus 8; ovario 8-locellato ovulis 8 uno utrumque loculum complente
placentatione axili. Fruetus bacati pyrenis 8 lobis calycis persistentibus.

154 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

with 4 lobes, with glands inside; corolla tube cylindrical, the lobes 5; stamens 5,
the anthers oblong, the filaments short; style slender, the stigmas 8; ovary
8-locellate, the ovules 8, each filling a locule, the placentation axile; ovarian disc
cushion shaped. Fruits baccate; pyrenes 8; the lobes of the calyx persistent.

The 8-carpellate nature of the flowers distinguishes the genus from all other
New World genera of the Rubiaceae. It is named for Robert L. Dressier.

1. Dressleriopsis panamensis Dwyer.^- type: Panama, Dressier 3560 (MO, holo-
type). — Fig. 35.

Suhshruhs to 1 m tall, often with a pair of slender, rigid, opposite branches.
Leaves oblong, to 19 cm long, to 9 cm wide, deltoid to subobtuse, short acuminate,
basally cordate, truncate, or auriculate, the lateral veins ca. 15, arcuate, forming
a conspicuous submarginal vein, the intervenal areas, spreading reticulate, rigidly
chartaceous, discolorous, pilose petioles to 6 cm long; stipules triangular, free,
ca. 7 mm long, ca. 6 mm wide, carnose, glabrous inside, diffusely hirsute out-
side. Inflorescences axillary, the flowers congested into sessile globose heads 2 cm
in diam.; bracteoles subulate, to 2 mm long. Flowers subsessile; hypanthium
subrotund, to 2 mm long hirsute, the hairs septate, lobes 4, to 4 mm long, ca. 1.6
mm wide, basally attenuate glandular inside at the base; corolla white, cylindrical,
ca. 3.5 mm long, glabrous within, the lobes 5, often horizontally expanded, oblong,
ca. 4 mm long, villose, attenuate; upwards anthers 5, slightly exserted, subsessile,
oblong, ca. I mm long, glabrous; disc cushion shaped, ca. 0.5 mm wide, the
locules 8, a single linear oblong axile ovule filling each locule, the style narrowly,
to 5 mm long, the stigmas 8, oblong, radiate. Fruits (?immature) berrylike,
globose, to 1.1 cm in diam., not ribbed, compressed at the apex, the internal
pulp with abundant raphides, the pyrenes pyrenes 8, pear shaped or somewhat
hemispherical, ca. 3 mm long, hard, orange, with an oblong scar on the concave
side.

Dressleriopsis panamensis is known only from Panama.

colon: Between Cerro Jefe and La Eneida, 900 m, Correa & Dressier 1 121 (MO). Santa Rita
Ridge, ca. 800 ft, Duyer ct al. 9035 (MO). Panama: Llano-Carti Road, 7-12 km from Panamerican

Highway, Croat 25164 (MO). La Eneida, Dressier 3560 (MO). 16 km above Panamerican Highway.
road from El Llano to Carti-Tupile, Kennedy el al. 2433 (MO). El Llano-Carti Road, 10 km from
Panamerican Highway, Mori & Kallunki 2340 (MO). El Llano-Carti Road. Mori ct Kallanki 2852,
2856 (both MO). El Llano-Carti Road, 8 km N of Panamerican Highway at El Llano, Nee & Warm-

hrodt 10343 (MO).

^2 Dressleriopsis panamensis Dwyer. spec. nov. Snffrntires ad 1 m alti. Folia oblonga acuminata
versus basin sacpe cordata, venis lateralibus ca. 15. arcuatis, rigido-chartacea discoloria pilosa
praecipuc in costa venisque; stipulis triangularibus, ca. 7 mm longis, ad 7 mm latis, extus diffuso-
hirsutis; petiolis ad 6 cm longis, dense pilosis. Inflorescentiae sessiles. globosae in axillis inferioribus
superioribusque dispositae: bracteolis subulatis, ad 2 mm longis. Flores subsessiles hypanthio hirsulo
lobis calycis 4, ad 4 mm longis; corolla tubo cylindrico, ca. 3.5 mm longo, intus glabro lobis 5
saepe applanatis, ad 4 mm longis, utrinque villosis; anthcris subsessilibus, ca. 1 mm longis;
loculis ovarii 8 placentatione axile, ovulis 8; stylo lineari, ad 5 mm longo; stigmatibus 8, oblongis,
radiate dispositis. Frintns globosi. ad 1.8 cm in diam.. dense hirsutis pyrenis 8 pyriformibus, ca. 3
mm longis.

1980 J

DWYER— FLORA OF PANAMA (Family 179. Rubiaceae)

155

Figure 35. Dressleriopsis panamensis Dwyer. Habit (x.45). [After Mori & Kalliinki 2852.]

34. DUROIA

Duroia L.f,, Suppl. PI. 209. 1781. Nomen cons, typf: D. crlopila L.f.

Dioecious trees or shrubs. Leaves usually coriaceous; stipules caducous,
oblong, connate, calyptrate. Corolla lobes contorted in bud; ovary 1-4 celled,
many ovulate, Fruit baccate, 1-4 celled; seeds compressed, horizontal.

Duroia includes about 20 species occurring in much of tropical America.

1. Duroia hirsuta (P. & E.) Schum. in Mart., Fl. Bras. 6: 367. 1889.

Tree or shrub \ branches hirsute. Leaves oblong obovate, villous, 10-28 cm
long. Inflorescences golden hirsute. Calyx truncate, the lobes linear, 5-12 mm
long; corolla white, the tube 2-3 cm long, pubescent, the lobes oblong. Fruits
golden hirsute, 2-6 cm long.

Not yet collected in Panama, Duroia hirsuta ranges from Guatemala to Costa
Rica and is expected in Panama.

156 ANNALS OF THH MISSOURI BOTANICAL GARDEN [Vol.67

35. ELAEAGNIA
Elaeagnia Wedd., Monogr. Cinch. 94. 1849. type: E. utilis (Goud.) Wedd.

Trees or shrubs. Leaves coriaceous, petiolate; stipules free or connate. Inflo-
rescences large, terminal, paniculate or rarely racemose. Flowers small, the ca-
lycine cup 5-lobed, inflated, persistent; corolla reduced, infundibuliform, cylin-
drical or somewhat bell shaped, the throat villous, the lobes 5, recurved,
contorted in bud; stamens 5, the anthers oblong, dorsifixed, the filaments exsert-
ed, geniculate at the base; ovarian disc orbicular, the style short, the stigmas 2,
obtuse, the ovules numerous, the placentas peltately affixed to the septum. Fruits
capsular, small, loculicidally dehiscent into 2 valves, these split apically; seeds
minute, the testa membranous.

Elaeagnia, a genus of about 15 species, ranges from Central America to South
America. It also occurs in Cuba.

Literature:

Steyermark, J. S. 1965. Botany of the Guayana Highland— Part VI. Mem. N.Y.
Bot. Garden 12: 186-192.

a. Leaves short pilose beneath; petioles less than 1 cm long; fruits ca. 4.5 mm long

1. E. aiirhidatii

aa. Leaves glabrous beneath; petioles 2.0-2.5 cm long: fruits ca. 7 mm long 2. E. nitidifolia

1. Elaeagnia auriculata Hemsl., Diag. PI. Nov. 32. 1879. type: Costa Rica, not
seen. — Fig. 36.

Shrubs or trees, the branchlets 4-angled, glabrous to pubescent, often rimose.
Leaves elliptic, ovate elliptic to obovate elliptic, often rounded, to 40 cm long,
to 25 cm wide, rounded to acute at the apex, often abruptly acuminate, rounded
or subauriculate at the base, the costa slender, prominulous, the lateral veins ca.
15, angular ascending, subcoriaceous, glabrous to densely short pilose above,
more so beneath; petioles absent or to 1 cm long; stipules deciduous, oblong,
4.5-5.0 cm long, ca. 2 cm wide, rounded at the apex, truncate at the base. Inflo-
rescences often exceeding the leaves, often epedunculate or seemingly with 3

peduncles arising from the apex of the twiglets, puberulent, the rachis twisted,
plane, the branches opposite, well spaced, the flowers solitary or in few-flowered
cymules. Flowers on slender pedicels, 2-5 mm long; hypanthium compressed
oblong, ca. 1 mm long, pilose, the calycine cup ca. 0.8 mm long, scarious, eglan-
dular within, the calycine lobes 5, expanded, compressed rotund, ca. 1 mm long;
corolla white, the tube short, ca. 1 mm long, petaloid, venose, glabrous outside,
densely short bearded at the point of filament attachment, the hairs forming a
ball-like bulge, the lobes 5, oblong, ca. 1 mm long, rounded; stamens 5, exserted,
the anthers oblong, ca. 2 mm long, the filaments to 2.5 mm long, glabrous distally;
style to 2.5 mm long, the stigmas 2, fleshy but subplane, to 2 mm long, to 1 mm
wide. Fruits oblong or oblong rotund, ca. 4.5 mm long, puberulent or glabrous,

dark red.

Elaeaf^nia auriculata is known from Honduras, Costa Rica, and Panama. The
single collection from Panama is in fruit with only one leaf present and the inflo-

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

157

Figure 36. FJacagnia aurlcuhita Hemsl.— A. Habit (x>/2).—B. Flower (x5). |After Dcke 6580.]

rescence in poor condition. The flower is described above from Molina 6102 (F)

from Costa Rica.

dariln: Cerro Pirre, W slope, Diike 6580 (MO).

2. Elaeagnia nitidifolia Dwyer.^'^ type: Panama, Corrca ct aL 1608 (MO, holo-

type).

^^ Elaeagnia futidifolia Dwyer, spec, nov. Arhores ramulis glabris in sicco nitido-glabris. Folia
oblonga, 7-23 cm longa, 4.0-8.5 cm lata, apice obtusa vel lato-cuneata acuminata basi acuta venis
lateralibus 12-14 tenui-coriacea glabra minuto-farinoso-punctata; petiolis 2.0-2.5 cm longis, glabris;
stipulis deciduis fortasse connatis cylindricis, ad 0.5 cm longis, truncatis coriaceis glabris. Inflorcs-
centiae terminales cymoso-paniculatae, 12-16 cm longae, ad 10 cm latae, glabrae, in sicco rubrae;
pedunculo ad 8.5 cm longo ramis paucis oppositis vel suboppositis bene distantibus; floribus in cymis
saepe unilateraliter dispositis. Flores pedicellati cupula calycis crateriforme, ca, 2 mm longa, edentata
coriacea glabra: corolla alba tubo ad 1.5 mm longo. extus glabro intus barbato lobis 2-plo longioribus;
antheris oblongis, ca. 1.5 mm longis, exsertis, filamentis ca. 2.5 mm longis. Fructus turbinati, ad 7
mm longi, loculicide dehiscentes glabri cupula calycis ad 2 mm longa persistenti seminibus multis ca.
0.1 mm longis, reticulalis.

158 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vol. 67

Trees, to 6 m, the branchlets often drying green, glabrous, subterete, the
petiole scars conspicuous, crowded, the terminal bud black, shiny, covered with
a resinous exudate. Leaves ascending, oblong, 7-23 cm long, 4.0-8.5 cm wide,
obtuse to widely cuneate above, acuminate, the acumen triangular, to 0.5 cm
long, acute at the base, often ultimately slightly contracted, the costa prominulous
above, prominent beneath, drying dull, shiny red, the lateral veins arcuate, 10-
14, prominulous above and below, the submarginal undulate vein obvious or not,
the reticulate intervenal areas small but conspicuous, the smaller veins pinnati-
form, thin coriaceous or coriaceous, usually slightly discolorous, drying tan be-
neath, glabrous, minutely farinose punctate under magnification above and below,
the margin thinly callose marginate; petioles 2.0-2.5 cm long, rigid, stout, shiny,
rubescent; stipules persistent only at twiglet tips, connate, located immediately
above the point of petiole attachment, perhaps eventually splitting, cylindrical,
somewhat oblong in outline, to 0.5 cm long, truncate, coriaceous, glabrous, cov-
ered with a glistening resin. Inflorescences terminal, cymose paniculate, some-
what oblong in outline, 12-16 cm long, to 10 cm wide, glabrous, with 3 peduncles
terminating the twiglet, or perhaps the peduncle suppressed and 3 branches aris-
ing directly from the twiglet tip, the median peduncle or branch 7.0-8.5 cm long,
rigid, ascending, piano compressed, somewhat twisted, drying shiny deep red,
the lowermost branches 2.5-3.0 cm long, 7-9 cm between the first and second
pair of branches, the smaller branches from these 2.5-3.0 cm long, opposite or
subopposite, well spaced, the flowers usually 12-15 per branchlet, the cymes 3-
4-flowered, usually unilaterally disposed, the flowers occasionally solitary. Flow-
ers pedicellate, the pedicels reflexed, to 5 mm long, glabrous, stiff; hypanthium
scarcely measurable, acute at the base, glabrous, the calyx cup bowl shaped, ca.
2 mm long, truncate, edentate, coriaceous, glabrous; corolla white, the tube cy-
lindrical, ca. 1.5 mm long, glabrous outside, densely barbate within, the hairs
thin, white, the lobes 5, oblong, ca. 2 times the length of the tube, obtuse, glabrous
except densely barbate at the base within; stamens 5, exserted, the anthers ob-
long, ca. 1.5 mm long, sub-basifixed, the filaments ca. 2.5 mm long, glabrous,
affixed near the mouth; ovarian disc pancakelike, marcescent on drying; style ca.
4 mm long, glabrous, the stigmas 2, ca. 1.5 mm long, scarcely wider than the
style. Fruits turbinate, to 6 mm long, truncate, glabrous, drying brown, withered
reticulate on drying, loculicidally dehiscent, the orifice of the pericarp appearing
gaping and bilabiate within the calycine cup, the latter arising slightly below the
rounded apex of the pericarp, extending for ca. 1.5 mm, to 5 mm broad; seeds
numerous, oblong to hemispherical, compressed, ca. 0.2 mm long, reticulate.

Elaeagnia nitidifolia is known only from Panama but is to be expected in the
Dept. Choco, Colombia. The present distribution in Panama is noteworthy: Cerro
Jefe and the Panama-Colombia frontier. A study of ''mountain hopping'' species
such as the present one will provide an interesting study as far as the Republic
of Panama is concerned. It appears to be closely related to the Venezuelan species
E. rniz-teranii Steyermark, which differs in having much larger leaves and a
ciliolate calyx. Steyermark's species'^^ has the calyx dentate, while the flowers in

^^ See Fig. 65, p. 406, Flora de Venezuela, primera parte.

19801 DWYER— FLORA OF PANAMA (Family 179. Ruhiuccae) 159

the new species, as observed on Correa et al. 1608, are toothless. The other

Mori

sistent calyx cup usually deeply split; this I regard as a product of the fruit's
maturation.

darien: Serrania del Darien, Panama-Colombia border, Cerro Mali base camp, 1500 m, Gentry
& Mori 13785 (MO). Panama: Cerro Jefe, 1000 m, Correa et ol. 1608 (MO).

36. EXOSTEMA

ifl'

Solendra Hook, in Benth. & Hook., Gen. PI. 2: 12, 43, 1873, not Solenandria P. Beauv. ex Ventanat

(1803). type: 5. ixoroides J. D. Hooker.
Steudelago Kuntze, Rev. Gen. PI. 298. 1891. Substitute name for Solenandra Hooker, 1873.

Shrubs or trees. Leaves petiolate or subsessile; stipules entire or bifid, often
persistent. Inflorescences terminal, paniculate or corymbose, flowers occasion-
ally solitary and axillary (not Panama spp.); bracteoles present or absent. Flowers
pedicellate, calyx 4-5-lobate, the lobes often narrow; corolla usually elongate,
the lobes (4-)5, usually elongate and narrow with 2 exterior in bud; stamens 5,
usually exserted, the filaments linear, connate at the very base and appearing free
of the tube; ovarian disc annular, the style slender, usually exserted, the capitate
stigma entire or 2-Iobed, the ovules numerous, the placentas peltately adnate to
the septum. Fruits capsular, 2-celled and 2-valvate, septicidally dehiscent with
each valve entire or bivalvate; seeds usually numerous, imbricate, the body ob-
long, compressed, the testa membranous, produced into a wing.

Mexico

West Indies.

1. Exostema mexicanum A. Gray, Proc. Am. Acad. Arts 5: 180. 1861. type:
Mexico, Ervendberg 125 (not seen).

Trees to 9.5 m tall or shrubs, the twigs terete, smooth, drying gray or gray
green, lenticellate, glabrous, the nodes 5-7 cm apart. Leaves oblong, obovate
oblong or widely ovate, 5.5-17 cm long, 2-9.5 cm wide, deltoid to obtuse at the
apex, often tapering into a vague acumen or somewhat attenuate acuminate, the
basal acumen to 1.5 cm long, ultimately obtuse, widely cuneate or rounded,
occasionally truncate, the costa plane or immersed above, prominulous beneath,
the lateral veins about 10, the intervenal areas spreading reticulate, chartaceous,
glabrous above and beneath, often minutely pubescent in the axils beneath; pet-
ioles to 1.5 cm long, strict or curved, stout; stipules (fide Standley) 2.5-4 mm
long, triangular, acuminate or long cuspidate. Inflorescences terminal and/or ax-
illary, glabrous, thrysoid cymose, shorter than or occasionally as long as the
uppermost leaves, the peduncles absent and then with 3 branches arising from
the tip of the twig, or distinctly pedunculate, the peduncles 2-5 cm long, the
branches few, well spaced, divergent, the first and second pairs opposite, the
cymules terminal, disposed in a fan or brush-like pattern. Flowers white, short
pedicellate; hypanthium oblong, ca. 2 mm long, glabrous or puberulent, the ca-
lycine cup scarcely measurable with patches of minute reddish glands within, the

160

ANNAl.S OF THE MISSOURI BOTANICAL GARDEiN

[Vol. 67

Fir.LiRi: 37. Exostema mcxicanum A, Gray.— A. Habit (x'/z). — B. Inflorescence {xVi). [After
Dwyer S: Eiuis 7510. \—C. Flower ( x2. 5).— D. Stamens, opened (x2.5). — E. Ovary, opened to show
ovules (x5).— F. Seed (x3). [After Dwyer 8391. \

calycinc teeth 5-6, subequal, widely or narrowly subulate; corolla with tube
curved, narrow cylindrical, 7-10 mm long, ca. 0.5 mm wide, petaloid, puberulent
outside, glabrous within, the lobes 6, linear oblong, ca. 12 mm long, ca. 1.5 mm
wide, obtuse, glabrous within; stamens 6, the anthers linear oblong, 3-4 mm long,
basifixed, the filaments linear, to 16 mm long, free from corolla tube, attached
basally in a ring, ca. 1 mm long; style cylindrical, linear, to 24 mm long, thicker
above the middle, the stigmas clavate, ca. 0.6 mm long. Fruits septicidally bival-
vate, each valve splitting medially, fusiform oblong or obovate oblong, to 12 mm

I9K0I DVVYF:R— FLORA OF PANAMA (Family 179. Ruhiaceac) 161

long, 3-4 mm wide before splitting, lignose, glabrous, white punctate lenticellate;
seeds subfusiform, 7-9 mm long, 2-3 times longer than wide, membranous, the
body plane, oblong rotund, ca. 2 mm in diam., the wing at 1 end subtriangular
or oblong, obtuse, bifid at other end, the processes divergent, one oblong and
obtuse, the other shorter, subtriangular and acute.

The species is known from Mexico, Guatemala, Belize, Honduras, El Sal-
vador, and Panama.

CANAL zoNt: Madden Dam, Ihvycr & Ellas 7502. 7508 (both MO), 7510 (MO, NY): Dn'ycr839l
(MO), 9164 (MO, NY); Ihrycr & Nee 11979 (MO). Panama: Vera Cruz, Brisfan P'I9 (MO). Between
Rio Bayano dam and Canitas, DArcy 9385 (MO).

37. FARAMEA
Faramea Aubl., Hist. PI. Guiane 1: 102. 1775. type: F, corymhosa Aubl.

Trees or shrubs. Leaves often coriaceous; stipules usually entire, free or con-
nate, usually awned, the base usually ovate triangular. Inflorescences terminal
or axillary, often disposed as loose decussate panicles or umbellate cymes, ulti-
mately cymose; bracts present or absent, occasionally with 2 large bracts at the
base of the inflorescence; pedicels often stout, stiff. Flowers 4(-5)-merous, ca-
lycine cup truncate, teeth present or absent; corolla tubular or salverform, the
lobes usually 4, valvate in bud, usually wider than the tube; pollen dimorphic;
stigmas disposed as 2 filamentous or clavate lobes, ovary 2-celled, becoming
l-locular with the ovules collateral or connate at the apex of the incomplete
septum. Fruits drupaceous, depressed oval to rotund, 1-seeded by fusion of the
ovules; seed horizontal, usually with a deep ventral groove.

Faramea is well distributed in Central America and tropical South America.
It also occurs in the West Indies. There are about 125 species in the genus.

a. Leaves conspicuously callose marginate.

b. Leaves oval (rarely a few oblong oval), 4-9 cm long, acuminate, the acumen less than

0.9 cm long 10. F. oralis

bb. Leaves oblong or obovate oblong, 10-15 cm long, cuspidate, the acumen 1-1.5 cm

long 8. F. luteovirens

aa. Leaves not callose marginate.

c. Leaves with 2 submarginal veins.

d. Leaves 3-7 cm long: flowers ca. 4 per inflorescence 17. F. woodsonii

dd. Leaves 7-30 cm long; flowers many per inflorescence.

e. Leaves not glaucescent or pruinose beneath: calycine teeth obsolete.

f. Leaves oblong, ovate oblong or elliptic. 4-7(^10.5) cm wide: calycine

cup minute; corolla white, blue or deep purple 14. F. suerrensis

ff. Leaves narrowly elliptic, to 3.5 cm wide; calycine cup ca. 2 mm long:

corolla bluish white 13. F. scalaris

ee. Leaves glaucescent or pruinose beneath; calycine teeth to 1 mm long —

3. F, biiUata

cc. Leaves with 1 distinct submarginal vein or lacking a submarginal vein.

g. Leaves bullate 11. F. papulata

gg. Leaves not bullate.

h. Leaves with lateral veins 6-9; inflorescence racemose .. 2. F. bocataurcnsis
hh. Leaves with lateral veins 10-20; inflorescence not racemose.

i. Stipules 2.5-3 cm long: leaves with lateral veins ca. 20.

j. Stipules free; inflorescence an ovate, blunt head

4. F. caput-anguis

ij. Stipules connate; inflorescence spreading.

162 ANNALS OF THE MISSOURI BOTANICAL GARDHN [Vol. 67

k. Calycine teeth absent; calycine cup ca. 0.6 mm long

__ 16. F, tcrreyeae

kk. Caylcine teeth ca. 0.5 mm long; calycine cup ca. 1.5 mm

long 6. F. jefensls

ii. Stipules 0.3-2 cm long; leaves with lateral veins 10-19.

L Leaves oval or suboval 10. F. ovaUs

II. Leaves not oval but elliptic or oblong,
m. Leaves with lateral veins 14-17.

n. Calycine cup ca. 0.3 cm long; corolla tube ca. 15 mm

long: anthers ca. 4 mm long: stipules ca. 0.4 cm

long L F, altipetens

nn. Calycine cup 1-1.2 cm long; corolla tube 25-30 mm

long; anthers ca. 8 mm long; stipules 1-1.5 cm long

12. F. paiiciflora

mm. Leaves with lateral veins 8-13.

o. Corolla tube ca. 6.5 mm long; leaves drying emerald

green 7. F. Ucsncri

oo. Corolla tube 7-17 mm long; leaves not drying em-
erald green.

p. Anthers ca. 8 mm long 9. F. occiclentalis

pp. Anthers to 4 mm long.

q. Corolla tube 12-17 mm

15. F. tahiDiancandn

qq. Corolla tube 7-11.5 mm long.

r. Inflorescences ca. 4-flowered; leaves

3-7 cm long 17. F. wooJsonli

rr. Inflorescences many-flowered; leaves

5-16 cm long 5. F. curycarpa

Mar

lotype).

Trees to 4 m tall, the branchlets smooth, glabrous, piano compressed, twisted,
the nodes well spaced. Leaves oblong lanceolate, 11-17.5 cm long, 3-5.5 cm
wide, rounded or obtuse toward the apex, caudate acuminate, the acumen to 1,2
cm long, ultimately obtuse, basally cuneate, usually lightly inequilateral, the costa
subplane to prominulous on both sides, the lateral veins ca. 14, widely arcuate,
dull yellow, the submarginal vein undulate, to 3 mm from the margin, the inter-
venal areas conspicuously reticulate, the smallest veinlets conspicuously promi-
nulous, coriaceous, concolorous, dark yellow or yellow brown when dry, gla-
brous; petioles ca. 1 cm long, rigid, stipules free, ovate triangular ca. 0.4 mm
long, lacking terminal setas. Inflorescenees terminal, solitary, glabrous, panicu-
late, to 9 cm long, to 10 cm wide; peduncles to 3 cm long, terminated by 4-5
branches, these radiately disposed, each branch with 1-2 pairs of opposite, an-
gular ascending branchlets terminated by 1-2 few-flowered cymules; pedicels 2-
5 mm long, drying black. Flowers glabrous; hypanthium ca. 1 mm long; calycine
cup narrowly cylindrical, to 4 mm long, eglandular within, truncate at the apex,
often undulate, the margin occasionally with shallow notches, teeth absent; co-

^'' Farafuea altipetens Dwyer, spec. nov. Arhorcs ad 4 m altae, ramulis glabris ultime tortis.

FaJia oblongo-lanceolata, venis lateralibus ca. 14 interveniis conspicue reticulatis venulis minimis
prominulis; stipulis ca. 0.4 mm longis. Inflorcsccntiae terminales solitariae, pedunculis 4-5 ramis
radiate dispositis terminatis ramo utroque 1-2 jugis ramulorum oppositorum cymulis paucis. Flares
glabri; corolla alba lubo ca. 15 mm longo, lobis 4 ad 7 mm longis; antheris subsessilibus, ca. 4 mm
longis. Frtictus non visi.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhicueac) 163

rolla white, the tube narrowly cylindrical, ca. 15 mm long, the lobes 4, crassate,
triangular subulate, to 7 mm long; stamens included, 4, the anthers subsessile,
linear ovate, ca. 4 mm long, minutely apiculate, inserted near the middle of the
corolla tube; style ca. 9 mm long, the stigmas 2, slender, ca. 2 mm long. Fruits
not seen.

Faramca altipetens is known only from Panama but is to be expected in
nearby Colombia. It is marked by the combination of short stipules and anthers
and a relatively elongate corolla tube. The smallest veinlets are prominulous with
the intervenal areas having a striking reticulate appearance. It is closely related
to F. pauciflora Dwyer in which the flowers are white and few, ca. 3 per cyme.
F. pauciflora, however, has much larger stipules, corolla tube, and anthers.

darien: Cerro Tacarcuna, 1500 m. Gentry & Mori 14013 (MO).

2. Faramea bocataurensis Dwyer. ^^ type: Panama, Kirkbridc & Duke 806 (MO,
holotype).

Small trees, to 5 m tall, the branches smooth, glabrous, the nodes well spaced.
Leaves lanceolate, 6-11.5 cm long, 2-2.7 cm wide, acute at the apex, acuminate,
the acumen to 2 cm long, ca. 0.15 cm wide in the middle, acute or subobtuse at
the base, the costa prominulous, the lateral veins 6-9, arcuate, prominulous, with
1-4 smaller veins between a pair of adjacent veins, these subparallel but soon
branching, the intervenal areas spreading reticulate, chartaceous, bicolorous, gla-
brous; petioles slender but stiff, ca. 0.5 cm long; stipules connate, the sheath
cylindrical, ca. 0.7 cm long, stiffly petaloid, glabrous, each part with an awn

0.8 mm long. Inflorescences terminal, solitary, glabrous, to 3.5 cm long, epedun-
culate, cymose racemose, the branches to 5 mm long or absent, the cymules few
flowered, bracteoles deciduous, subulate, 0.8-2 mm long; pedicels ca. 4 mm long.
Flowers purple, glabrous; hypanthium subrotund, ca. 0.5 mm long, the calycine
cup turgid, to 0.35 mm wide, smooth, petaloid, eglandular within, with 4 subulate
lobes, to 0.8 mm long; corolla (only one seen in bud) to 7.5 mm long, the tube
cylindrical, rigidly carnose, to 6 mm long, ca. 1 mm wide toward the base, the
lobes 4, narrowly ovate lanceolate, ca. 2 mm long; stamens 4, the anthers nar-
rowly oblong, ca. 4 mm long, obtuse and subsagittate at the base, the filaments
slender, ca. 1 mm long, affixed near the middle of the tube, the disc of the ovary
to 1.3 mm long, 3 times longer than wide, the style linear, ca. 6 mm long, the
stigmas 2, narrowly oblong, ca. 1 mm long. Fruits not seen.

Faramea bocataurensis is known only from Panama. While the flowers are
known only from the bud stage, the calycine cup is remarkably short as are the

^^" Fanunea bocataurensis Dwyer, spec. nov. Arhores parvae ad 5 m altae. Folia lanceolata, 6-
11.5 cm longa, 2-2.7 cm lata, apice acuta caudato-acuminata, acumine ad 2 cm longo, basi acuta vel
subobtusa, venis lateralibus 6-9: petiolis ca. 0.5 cm longis; stipulis connatis vagina cylindrica, ca. 0.7
cm longa, parte utraque seta ad 0.8 mm longa. Inflorescentiac terminales solitariae glabrae, ad 3.5
cm longae, epedunculatae cymoso-racemosae ramis solum 3-4 mm longis vel nullis, cymulis floribus
paucis praedilis; bracteolis saepe deciduis, subulatis, 0.8-2 mm longis. Flares glabri; pedicellis ca.
4 mm longis, cupula calycis turgida, ad 0.35 mm lata, ca. 0.5 mm longa, dentibus 4, ad 0.8 mm longis;
corolla purpurea, ad 7.5 mm longa; antheris lineari-oblongis, ca. 4 mm longis. Fnwtus non visi.

164 ANNALS OF THE MISSOURI BOTANICAL (]ARDEN [Vol. 67

calycine teeth. The leaves are as narrow as any species of Panamanian Faramea
and have the fewest lateral veins. The reduced racemelike inflorescence is note-
worthy.

BOCAS DLL TORo: E slopc of La Zorra to divide, Kirkhtulc & Diikc 806 (MO).

3. Faramea bullata Standley, Field Mus. Nat. Hist., Bot. Ser. 4: 294. 1929. type:
Panama, Cooper 507 (F, holotype; NY, isotype).

Trees or shrubs, 2-4.5 m tall, glabrous, the branchlets smooth, the nodes well
spaced. Leaves oblong, 15-21.5 cm long, 5.5-7 cm wide, acuminate, the acumen
ca. 2 cm long, basally cordate or occasionally rounded, the costa prominent, the
lateral veins ca. 15, immersed above, with 1-2 somewhat irregular intermediate
veins between pairs of lateral veins, the submarginal veins 2, the outermost ca.
1.5 mm from the margin, the other ca. 5 mm from the margin, subcoriaceous,
shiny green above, bullate, glaucescent and pruinose beneath; petioles ca. 0.5 cm
long, glabrous; stipules presumably connate basally, each part oblong rotund, ca.
8 mm long, tapering acutely toward the apex, the awn subulate, to Vs the length
of the sheath, coriaceous, glabrous. Infloreseences cymose paniculate, to 7.5 cm
long, often almost as wide as long; peduncle thick or slender, to 3 cm long, the
branches few, angular ascending; pedicels 2-6 mm long. Flowers with hypan-
thium and calycine cup short, the teeth triangular or subulate, to 1 mm long;
corolla blue or bluish white, the tube ca. 9 mm long, ca. 1.2 mm wide in bud.
Fruits compressed rotund, to 1.5 cm wide, smooth, ca. twice as long as wide,
drying black, smooth, glabrous; "seed gray, deeply and narrowly exsculpate at
the base, smooth, ca. 1 cm long" (fide Standley).

Faramea hitllata is known only from Panama. In preparing the above descrip-
tion I have drawn freely from Standley's original description.

BOCAS DiiL TORo: Kankintoc, 10 mi above Holstein near Almirante, Cooper 507 (F, NY, US).
Punta Pena, Chiriqiiicito, Lewis ct al. 2145 (F. MO), vpraguas: Caribbean slope above Rio Primero
Brazo. 5 mi NW of Santa Fe, LicMwr 965 (MO).

4. Faramea caput-anguis Dwyer & Hayden, Ann. Missouri Bot. Gard. 55: 39.
1968. type: Panama, Dwyer & Ganger 7375 (MO, holotype).

Trees to 8 m tall, the branchlets smooth, drying black and glabrous, the in-
ternodes to 7 cm long. Leaves oblong, to 26 cm long, to 12 cm wide, acute or
acuminate at the apex, the acumen to 1.5 cm long, acute at the base, the costa
subplane above, prominent beneath, the lateral veins ca. 20, arcuate, prominent,
1-2 cm apart, the venules pinnatiform, thin coriaceous, often yellow green be-
neath, minutely yellow puberulent; petioles 2-4 cm long, smooth; stipules per-
sistent, triangular, to 3 cm long, to 0.8 cm wide, acute, lignose, glabrate. In-
florescenees usually 1 per axil; peduncles divergent, stiff, smooth, glabrous,
expanding terminally, to 9 cm long; heads ovoid, apically blunt, solitary, cymose,
at first gemmiform and fusiform oblong, to 2 cm long, ca. 1 cm wide, the outer
bracts ca. 5, widely fusiform oblong, glabrous, foliose; flowers 7-9 per head, each

1980] DWYER— FLORA OF PANAMA (Family 179. Rithiaccac) 165

pair with 2 to several bracteoles. Flowers glabrous, the calyx narrowly campan-
ulate, to 8 mm long, the lobes 4, unequal, subulate, the shorter lobes ca. 4 mm
long, the longer lobes ca. 8 mm long; corolla white, tube ca. 25 mm long, narrow
medially, dilated above, the lobes 4, lanceolate, to 7 mm long, acute; stamens 4,
the anthers linear oblong, ca. 5.5 mm long, the filaments affixed near the middle
of the tube; ovules orbicular. Fruits oblong, to 1.5 cm long, to 1 cm wide including
persistent calyx, smooth, glabrous, glandular shiny, drying black, the calyx 2-

mm long.

5

Faramea caput-angiiis is known only from Panama.

Panama: Cerro Jefe, Busey 791 (MO). La Eneida, Cerro Jefe, Correa & Dressier 954, 1120 (both
MO). Cerro Jefe, Duver et al. 7248 (MO): Dwyer & Gcuiiier 7375, 7388A (both MO): Dwyer et cil.
8236 (MO). 9.4 km N of Goofy Lake, 900 m, Folsoin et al I963A (MO). Cerro Jefe, iMteyii c6
Foster 1140 (MO).

5. Faramea eurycarpa Donnell Smith, Bot. Gaz. 44: 113. 1907. type: Costa Rica,
Toncluz 12880, not seen.

F. phitycorpa Dwyer & Hayden, Ann. Missouri Bot. Card. 54: 140. 1967. type: Ecuador, Owenhy
2711 (MO, holotype).

Trees to 4 m tall, the ultimate branchlets becoming angular, smooth, glabrous,
the nodes well spaced. Leaves oblong or oblanceolate, 5-16 cm long, 1.5-5.5 cm
wide, rounded or obtuse at the apex, rarely cuneate, contracted into a slender
acumen, the acumen to 1.3 cm long, ultimately obtuse, basally cuneate to obtuse,
rarely truncate or auriculate, often inequilateral, the costa prominulous, the lat-
eral veins ca. 13, at first nearly perpendicular, then widely arcuate, united to form
a distinct undulate submarginal vein, to 0.3 cm from the margin with 1-2 smaller
veins between 2 lateral veins, the intervenal areas spreading reticulate, charta-
ceous to papyraceous, concolorous, glabrous, the margins delicately revolute;
petioles slender to stout, to 0.5 cm long, glabrous; stipules connate, the 1 at the
twig tip dome shaped, to 6 mm long, the awn shorter than the body, the inferior
stipules more cylindrical, to 7 mm long, the sheath separating above into 2 tri-
angular parts, each with an awn ca. 2 mm long, glabrous. Inflorescences terminal,
solitary, oblong rotund to subcuneate, to 9 cm long, to 9 cm wide, paniculate,
the peduncle 2-5 cm long, the lowermost branches opposite, to 3 cm long; the
upper branches disposed as 2-3 pairs, usually opposite, the branchlets terminating
in a fan of few-flowered cymules. Flowers with pedicels ca. 1 cm long, stiff;
hypanthium ca. 0.75 mm long, glabrous, the calycine cup compressed cup shaped,
shorter than the hypanthium, scarious and eglandular within, the teeth 4-5, ir-
regular, unequal, triangular or subulate, to 0.8 mm long, acute; corolla purplish
or white with a purplish apex, narrowly cylindrical, to 1 cm long, glabrous, car-
nose; corolla lobes 4, ovate, to 3 mm long, occasionally slightly acuminate; sta-
mens' subsessile, the anthers 4, narrowly oblong, 3-4 mm long, minutely apicu-
late, attached near the middle of the tube; style slender, ca. 8 mm long, the
stigmas fingerlike, slightly unequal, to 1 mm long, scarcely wider than the style.
Fruits drying black, bright blue, compressed rotund, wider than long, to 1.3 cm
wide, smooth, glabrous, the calycine scar ringlike, scarcely elevated.

166 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

Faramca curycarpa is known from Costa Rica and Panama. It is also known
from Colombia and Ecuador. Leon 537 (MO) from the Dept. Choco, Colombia,
seems to be F. curycarpa, although it reaches 10 m in height, has a strongly
contracted inflorescence, and stipules to 1 cm. Forcro ct at, 3073 (MO), also from
Dept. Choco, superficially resembles F. curycarpa but has corolla lobes to 15
mm in length.

coclr: La Mesa, Dwyer83I8 (MO), colon: Santa Rita Ridge, Dwyer 8554 (MO), darien: 1-3
mi N of Paya, Duke 14005 (MO). Panama: Cerro Campana, Allen 1877 (F, MO, US); Correct &
Dressier 291 (MO); Croat 14217. 17240, 22779 (all MO); Duke 10737 (MO, OS); Dwver et al. 4718
(MO); Gentry 1831 (MO, NY), 4904. 4952 (both MO); Kirkhride & Havden 277 (MO, NY); Kirkhride
337 (MO, NY); Lewis et al. 3049 (MO, SCZ, UC, VEN). El Llano-Carti Road, 21,1 km from Pana-
merican Highway, Mori & KaUunki 5112 (MO). El Llano-Carti Road, 11-12 km from Panamerican
Highway, Mori et al. 6885 (MO). Cerro Campana, Mori & Bolten 7703 (MO). El Llano-Carti Road,
16 km N of Panamerican Highway, Nee & Dressier 9356 (MO). El Llano-Cartf Road, Nee et aL 8769
(MO). Cerro Campana, 825-1000 m. Nee <^ Stockwel! 1 1581 (MO); Porter et al. 4162 (COL, DUKE,
MICH, MO, NY, SCZ, UC, VEN).

6. Faramea jefensis Dwyer & Hayden, Phytologia 15: 56. 1968. type: Panama,

Tyson ct al. 3396 (MO, holotype; NY, isotype).

Trees or shrubs, the branchlets strongly arcuate, ultimately angular, black
when dry, the nodes somewhat turgid, the internodes to 5 cm long, minutely
ciliolate. Leaves narrowly elliptic, 9-15 cm long, 2-5 cm wide, long cuspidate,
the cusp to 1.5 cm long, ca, 1.5 mm wide, basally cuneate, often inequilateral,
the costa prominulous above and beneath, the lateral veins ca. 20, scarcely ar-
cuate, delicately prominulous beneath, the submarginal vein irregular undulate,
ca. 3 mm from the margin, thin coriaceous, concolorous or discolorous, drying
a dull yellow beneath, glabrous, the margin scarcely thickened; petioles to 0.8
cm long; stipules connate, cylindrical, ca. 2.5 cm long, the sheath separating into
2 triangular parts ca. Vi the length of the sheath, minutely puberulent. Inflores-
cences rarely reduced to a single flower, usually cymose thrysoid, to 2.5 cm long,
to 3 cm wide, the peduncle scarcely developed, crassate, ca. 3 mm wide, the
branches few, to 1.5 cm long, ascending; bracts foliaceous, enclosing the juvenile
inflorescence, ovate, 2-4 cm long, 1-2 cm wide, cuspidate, glabrous, variegated
medially, the margins green; pedicels to 3 mm long. Flowers with the hypanthium
and calyx to 6 mm long, glabrous, the calycine cup cupular, to 1.5 mm long, ca.
2 mm wide, the teeth 4, minute; corolla dark blue, the tube cylindrical, 9-12 mm
long, the lobes 4, crassate, triangular in cross section, to 8 mm long; stamens 4,
the anthers 3.5-4 mm long, inserted below the middle of the corolla tube; style
ca. 8 mm long, the stigmas linear, ca. 2 mm long, erect, minutely papillate. Fruits
sessile or subsessile, compressed rotund, 0.3-0.5 cm long, 1-1.6 cm wide,
smooth, glabrous, drying black or yellow, the calycine cup scar annular, ca. 2
mm wide.

Faramea Jefensis is known only from Panama.

Panama: 6 miles above Goofy Lake, road to Cerro Jefe, Croat 15222 (MO). Cerro Jefe, 2900 ft,
Dnyer et aL 7279 (MO); Dwyer ct Gauffer 7328 (GH, MO, NY), 7338A (MO): Foster ^ Kennedy
1875 (MO); Tyson et al. 3396 (MO).

1980) DWYER— FLORA OF PANAMA (Family 179. Ruhiaccae) 167

7. Faramea liesneri Dwyer.

47

(MO)

Shrubs to 3V2 m tall, the branchlets terete, smooth, glabrous, uhimately an-
gular, the nodes well spaced. Leaves oblong, 4.5-9.5 cm long, 1.3-4 cm wide,
obtuse or widely deltoid at the apex, acuminate, the acumen often barely dis-
cernible, ultimately obtuse, basally acute, the costa prominulous above, the lat-
eral veins ca. 13, perpendicular to the costa, forming an undulate submarginal
vein ca. 1,5 mm from the margin, the margin lightly revolute, subcoriaceous,
glabrous, concolorous, drying emerald green or whitish green; petioles to 0.8 cm
long, rigid, glabrous; stipules connate, glabrous, with each part oblong rotund,
to 4 mm long, with a median keel continuing into a terminal rigid bristle shorter
or longer than the body, coriaceous. Inflorescences terminal or axillary, umbel-
liform, to 6 cm long, 1.5-4 cm wide, glabrous, the peduncle 1-2.5 cm long, the
cymes radiately disposed; pedicels slender, 0,5-1 cm long. Flowers with the
hypanthium oblong rotund, to 1.2 mm in diam., the calycine cup much reduced,
truncate, erose and minutely puberulent marginally, the lobes inconspicuous, to
0.3 mm long, truncate or obtuse; corolla violet blue, the tube narrowly cylindrical,
5-6.5 mm long, ca. 0.1 cm wide, thin carnose, glabrous inside and out, the lobes
4, unequal, narrowly oblong, to 6 mm long, to 2 mm wide, obtuse, glabrous;
stamens 4, the anthers oblong, ca. 3 mm long, obtuse, attached above the middle
of the tube; style Hnear, to 2.5 mm long, the stigmas 2, to 0.8 mm long, ovarian
disc to 0.6 mm long, rounded. Fruits on pedicels to 1.5 cm long, equal in length,
slender, radially disposed, the pedicel scars obvious, ringlike, the bracteoles few,
stipulelike, the body of the fruit round to compressed rotund, ca. 0.5-1.2 cm in
diam., smooth, glabrous, often emerald green when dry, the calycine cup small.

Faramea liesneri is known only from Panama. It is named in honor of Ronald
Lee Liesner, Missouri Botanical Garden, who has contributed much to our
knowledge of the flora of Panama. It is closely related to F. woodsonii Standley,
a small-leafed species, which has only stiffly chartaceous leaves and has a much
reduced inflorescence in which the flowers number about four. Faramea liesneri
has stiffly coriaceous leaves which dry a striking emerald green. This, coupled
with a much reduced, marginally pubescent calycine cup, as well as much reduced
calycine lobes, serves to distinguish the new species.

Panama: Road to Carti, 8-19 km N of El Llano, 350-500 m, Busey 913 (MO): Folsom et al.
1495 (MO); Mori 7733 (MO); Mori & KaUunki 2265. 2889 (both MO).

8. Faramea luteovirens Standley, Contr. U.S. Natl. Herb. 18: 138. 1916. type:
Panama, Pittier 4038 (US, holotype).

Trees to 10 m tall, the twiglets smooth, shiny, terete, finally angular, glabrous,

^'^ Faramen liesneri Dwyer. spec. nov. Fruticcs ad 2 m alti. Folia oblonga, 4.5-9.5 cm longa,
E3-4 cm lata, acuminc brevi, venis lateralibus ca. 15, subcoriacea glabra in sicco smaragdina vel
albido-smaragdina: petiolis ad 0.8 cm longis: stipulis connatis glabris parte utraque oblongo-rotunda,
ad 4 mm longis, in medio carinata seta rigida quam vagina brcviori vel vix longiori. Inftorescentiae
terminales vel axillares, ad 6 cm longae, umbelliformes cymis radiate dispositis. Flores pedicellis 0.5-
1 cm longis; cupula calycis brevissima lobis inconspicuis, ad 0.3 mm longis; corolla tubo angusto-
cylindrico, 5-6.5 mm longo, lobis 4, ad 6 mm longis; antheris ca. 3 mm longis. Fructus ca. 0.5 cm
diam,, smaragdini.

168 ANNALS OF IHH MISSOURI BOTANICAL GARDEN |Vol. 67

the nodes well spaced. Leaves oblong, obovate oblong, oblong rotund, to 15 cm
long, to 8 cm wide, obtuse, rounded or truncate at the apex, cuspidate, the cusp
to 1.5 cm long, often falcate, cuneate to acute, rarely truncate at the base, equi-
lateral, the costa plane to prominulous above, prominent beneath, the lateral
veins 10-13, widely arcuate, prominulous above and beneath, the intervenal areas
reticulate, the veinlets often disappearing, the margin conspicuously thickened,
coriaceous, glabrous, concolorous or discolorous, usually drying lemon yellow;

petioles stiff, to 1 cm long, glabrous; stipules appressed, deltoid to compressed
hemispherical, 2-4 mm long, the awn subulate, 3.5-5.5 mm long, coriaceous,
glabrous. Inflorescences axillary or occasionally terminal, the peduncle usually
suppressed (or if present to 2 cm long), branches few or absent, the flowers 1-2
per axil; pedicels strongly ascending, to 5 cm long, stiff, lignose, glabrous, ex-
'panded at the apex. Flowers with hypanthium oblong, ca. 9 mm long, coriaceous,
glabrous, the calycine cup ca. 4 mm long with patches of minute appressed glands
within, these covered with a membranous ''skin,'' the calycine teeth absent, the
margin undulate; corolla white, the tube narrowly cylindrical, falcate, ca, 2.2 cm
long, ca. 4.5 mm wide medially, contracted below the middle, stiffly petaloid or
subcoriaceous, glabrous, the lobes 3-4, lanceolate, 2,2-3.0 cm long, ca. 4.5 mm
wide at the base, obtuse, coriaceous, glabrous, the margins deeply infolded; sta-
mens 4, narrowly oblong, ca. 10 mm long, ca. 1.3 mm wide, obtuse, the filaments
slender, dorsifixed, ca. 4 mm long, inserted on the lower V3 of the corolla tube;
stigmas 2, erect, linear, to 2.2 mm long, the style slender, ca. 22 mm long, ca.
0.35 mm wide, the ovary thick walled. Fruits subglobose, 8-1.5 cm in diam.,
crowned with a calycine cup, smooth, glabrous, drying black.

Farcunea luteovirens is known from Panama and Colombia. The label on Duke
10055 indicates that the flowers turn maroon. ''Huesito'' Duke 14241 .

CANAL zone: Old Fort Lorenzo, Croat 154/1 (MO). Fort Sherman, Dwyer 7180 (MO). Pipeline
Road, vicinity of Ri'o Mendoca, Gentry 7469 (MO). Fort Lorenzo, Johnston 1514 (MO). Road to
Battery Vll, Johnston 1662 (MO). Rio Petitpie, Fort Sherman to Gatun Locks, Mori ct KaUunki 2701
(MO). Headwaters Rio Providencia, Nee ct Gentry 8656 (MO). Gatun, Standley 27177 (MO). Fort
Sherman, Standley 31006 (US, S); Tyson & Dwyer 1187. 1189 (both MO). Fort San Lorenzo, Tyson
2230 (MO). BARRO COLORADO ISLAND: Croat 5217. 6725 (both MO), chiriolji': 4.1 mi from Boquete,
Road to David, Kirkhride 102A (MO). Cano Quebrado, Pittier 6664 (US), colon: Near home of P.
H. Zimmerman, Achiote, Folsom et al. 3326 (MO). Loma de la Gloria, Fato, Pittier 4038, 4098 (both

US), darifn: Rio Tuira and Rio Paca, Duke 5024 (MO). Asnati, Duke 10055 (MO). Pinas, Duke
10603 (MO). Near Estero Grande off Ri'o Maresa, Duke 10954 (MO). Ri'o Perrecenega and Rancho
Frio, Duke & Elias 13883 (MO). Morti Hydro, Rio Morti, Duke 14212, 14241 (both MO). La Palma,
Pittier 6697 (US), san blas: Isia Sokatupu, Duke 8956 (MO). Playon Chico, Stier 187 (MO). Isia
Pino, near Miilatupo, Elias 1701 (GH, MO). Rio Mulatupo, headwaters, Elias 1764 (GH, MO). Co-
lombia: choco: Municipio de Rfo Sucio, Parque Nal. Los Katvos. Camino Tilupo Via Alto Sautata,
Leon 536 (MO).

9. Faramea occidentalis (L.) A. Rich., Mem. Soc. Hist. Nat. Paris 5: 176. 1834.
Fig. 38.

Ixora oeeidentalis L., Syst. Nat. ed, 2: 893. 1759. type: not seen.
Tetranieriuni odoratissin\um DC., Prodr. 4: 496. 1830. type: not seen.

Faramea zx'teki Standley, Contr. Arnold Arbor. 5: 147. 1933. r^ pe: Woodworth & Vestal 641 (F,
holotype).

Trees to 10 m tall, the branchlets terete, smooth, glabrous, ultimately angular.
Leaves oblong or elliptic, occasionally ovate oblong, 8-20 cm long, 2.5-11 cm

19801

DWYHR— FLORA OF PANAMA (Family 179. Riihiaccuc)

169

Figure 38. Faramea occidentalis (L.) A. Rich.— A. Flowering branch (xVi). [ Stern et al.
24.]— B. Fruit (xVi). [After Stimson 5385.]

wide, rounded or obtuse at the apex, acuminate, basally cuneate or truncate, the
costa subplane to prominuious above, prominulous to prominent beneath, the
lateral veins 8-10, widely arcuate, usually with 1-5 irregular and soon evanescent
veinlets between adjacent lateral veins, the intervenal areas spreading reticulate,
slightly marginate, papyraceous to thin coriaceous, glabrous above, sometimes
sparsely pubescent on the veins beneath; petioles stiff, often twisted, to 1.5 cm
long, to 0.2 cm wide, glabrate; stipules widely deltoid, 1-2 cm long, 0.5-0.8 cm
wide at the base, the terminal awn linear, about the same length or twice the
length of the body, glabrous. Inflorescences terminal, occasionally axillary, pa-
niculate, spreading, shorter than the terminal leaves, glabrous, the peduncle oc-
casionally absent, mostly 1-2.5 cm long, the primary branches usually 2(4), either
strongly ascending or the branches strict arcuate, divergent or deflexed, the cy-

170 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vol. 67

mules few, sparsely flowered; pedicels 1-2 cm long. Flowers with the hypanthium
oblong or rounded, the calycine cup cylindrical, ca. 2.2 mm long, stiffly carnose,
glabrous with scattered glands within, these oblong, ca. 0.1 mm in diam., the
teeth absent or evanescent; corolla white, the tube narrowly cylindrical, 1.7-2.2
cm long, carnose, glabrous, the lobes 4, lanceolate or ovate lanceolate, 1.1-1.8
cm long, ca. 0.45 cm wide; stamens 4, the anthers barely exserted, narrowly
oblong, ca. 8 mm long, 0.8 mm wide, minutely apiculate, versatile, the filaments
slender, ca. 2 mm long, inserted ca. 5 mm below the corolla mouth; ovarian disc
prominent, ca. 0.5 mm long, the style slender, as long as the corolla tube or less
than 14 the length, the stigmas 2, slender, erect, 2.5-5.5 mm long, obtuse, the
ovarian wall very thick. Fruits rotund or occasionally compressed rotund, often
wider than long, to 1.3 cm in diam., crowned with a cylindrical calycine cup, to
2 mm long, glabrous, drying black.

Faramca occidentalis ranges from southern Mexico to Panama; it also occurs

in the West Indies and throughout tropical South America. ''Huesito,'' "Benja-
min,^' "Palo Escrito" {Howell 119).

CANAL zone: Pina, near Waterways Exp. Sta., Duke 9252 (MO). Galiin, Dnyer 1781 (MO).
George Greene Memorial, Las Cruces Trail, Dwyer & Elias 7522 (MO, UC). Boy Scout Road, Madden
Dam, Dwycr 11974 (MO). Obispo, Hayes 66 (BM). Pipeline Road, 4 km NW of Gamboa, Nee 7076
(MO). Road S-8, Quebrada La Furnia, L5 km SW of mouth of Chagres River, Ncc 8636 (MO). Rio
Indio de Gatiin, Pitticr 2783 (BM, US). Madden Forest, Stimson 5385 (MO). Gamboa, Tyson et al.
4571 (MO). Madden Lake, Boy Scout Camp, Txson 5446 (MO), barro coi orado island: Croat
4024A, 4131 A, 63W, 6597, 9302, 14570, 14923,' 14961 , 14992, 17055 (all MO); Dwxcr 1407 (MO);
Kni^^ht 69-36 (MO); Oppcnhcimcr 66-11-2-0905, 66-10-20-0911, 66-8-28-1528, 67-1-17-1004 (all MO);
Foster 827 (MO); Rohyns 65-32 (MO); Wooclworth & Vestal 641 (F); Zetek 3992 (MO), colon; Rio
Culebra, Fittier 4164 (US), cocle: Lower Rio Anton, near El Valle, 800-1000 m, Allen 113 (MO). El
Valle, Alston 1849 (US), dariln: Rio Chico, Allen 4640 (BM). Rio Bayano above confluence with
Rio Chepo, Duke 3979 (MO). 0-4 mi up Rio Sabano, near Santa Fe, Duke 4153 (MO). Rio Tuira, 2
mi upstream from Boca de Cupe, Duke 5380 (MO). El Llano, Duke 5519 (MO). Without other locality,
Duke 8347 (MO). Rio Chucunaque, between Ri'o Membrillo and Rio Subcuti, Duke 8610 (MO). Near
Estero Grande off Ri'o Marea, Duke 10958 (MO). Rio Corso headwaters off Rio Pacora, ca. 500 m,
Duke 11943 (MO). Quebrada Maskia at Rio Pucro, Duke 13099 (MO). Cativo Swamp near Tusi-
juanda, 1 day up Ri'o Balsa from Manene, Duke 13534 (MO). Tumagantf, ca. 300 m, Duke 14155
(MO). Without other locality, Duke 15787 (MO). 1-3 mi N of Paya, Kirkhride & Duke 14013 (MO,
US). Payita, Stern et al. 248 (MO). Between Rio Escucha Ruido & Rio Tuira, Stern et al. 576 (MO).
2 mi E of Santa Fe, Tyson et al 4673, 4807 (both MO), los santos: Loma Prieta, 800-900 m, Duke
11843 (MO). Panama: Bayano Cuipo Forest, 34.6 km E of Bayano Bridge, Folsom 3533 (MO). Rio
Pasiga, first sharp bend. Gentry 2263 A (MO). 6 km SW of Arraijan, old road to Bique, Nee 6901
(MO). El Llano-Carti Road, 2.3 mi N of Panamerican Highway, ca. 250, Nee & Dwyer 9242 (MO).
Ri'os Pacora & Corso to headwaters of Rio Corso, Oliver 2374 (MO, UC, VEN). Juan Diaz, Standlex
30581 (MO). Macapale Island, Madden Lake, Txson 5468 (MO).

10. Faramea ovalis Standley, Contr. U.S. Natl. Herb. 18: 138. 1916. type: Pan-
ama, Pittier 3159 (US, holotype; C, BM, isotypes).

Trees 3-13 m tall, the twiglets slender, the nodes well spaced, glabrous.
Leaves small, oval, oblong rotund, or lanceolate, 4-9 cm long, 1.2-4.8 cm wide,
rounded, obtuse, or cuneate at the apex, abruptly short acuminate, the acumen
0.3-1.0 cm long, cuneate to obtuse at the base, the lateral veins 6-12, widely
arcuate, thin coriaceous, concolorous, glabrous, callose marginate; petioles to 1
cm long; stipules hemispherical, 3-9 mm long, the terminal awn longer than the
body. Inflorescences to 5 cm long, to 4 cm wide, few flowered, the pedicels

1980] DWYHR— FLORA OF PANAMA (Family 179. Ruhiaccac) 171

umbelliform from the apex of a shorter peduncle. Flowers with the pedicels ca.
1 cm long; calycine cup undulate; corolla purplish or white, tinted pink, the tube
ca. 12 mm long, the lobes 4-6 mm long. Fruits not seen.

Faramea ovalis is known only from Panama. The small, often oval leaves are
unique among the Faramea of Central America. Only one other species of Far-
amea in Panama has strongly marginate leaf blades, F. hiteovirens Standley. The
notes on the type collection indicate ''trunk straight, ca. 6 m, the crown pyram-
idal.'' Davidson 530, unlike the type, has lanceolate leaves. In addition the notes
on the label indicate that it is 30-40 ft in height and has a corolla tube which is
white and tinted pink. While these 3 differences from the type suggest that Da-
vidson 530 is a second species, I prefer to regard it as F, ovalis.

CHiRiQUi: Chiquero, Boquete District, 5500 ft, Davidson 530 (MO). Cerro Horqueta, 1200-1700
m, Pitricr 3159 (BM, C, US).

11. Faramea papulata Dwyer & Hayden, Phytologia 15: 57. 1967. type: Panama,
Tyson et al 2218 (MO, holotype).— Fig. 39.

Shrubs to 4 m tall, glabrous, the branchlets somewhat plane, the nodes well
spaced. Leaves elliptic, 12-14.5 cm long, 5.5-7.5 cm wide, rounded at the apex,
acuminate, the acumen 0.3-0.8 cm long, ultimately obtuse, basally rounded or
widely cuneate, the costa prominent beneath, the lateral veins 10-12, arcuate,
the submarginal vein conspicuous, ca. 0.5 cm distant from the margin, the inter-
venal areas prominent, coriaceous, shiny, bullate or papillate, bicolorous, occa-
sionally drying yellow; petioles 1-1,5 cm long; stipules connate at the base, di-
morphic, some small and compressed rotund, others elliptic, to 0.7 cm long,
elongate cuspidate, the cusp to 0.25 cm long, the stipular scars to 0.4 cm long.
Inflorescences terminal, umbelliform paniculate, 5.5-9 cm long, to 7 cm wide,
the peduncle smooth, to 4.5 cm long, ca. 0.3 cm wide, the primary branches 4,
equal, rigid, ascending sharply, the terminal branches ternately disposed, equal,
the flowers often ternately disposed on each branchlet; pedicels to 4 mm long.
Flowers with pedicels ca. 2 mm long, the hypanthium urceolate, ca. 1.2 mm long,
black when dry; calycine cup crownlike, ca. 2.2 mm long (on immature fruit to
4 mm long), brown, the margin undulate, the teeth evanescent or to 0. 1 mm long;
corolla white and purple white, drying black, the tube narrowly cylindrical, to 13
mm long, scarcely expanded toward the apex, glabrous, thickly petaloid, the
lobes 4, narrowly ovate triangular, to 13 mm long, carnose, glabrous; stamens 4,
included, the anthers 4, oblong, to 5.3 mm long, the connective apically produced
1 mm, the filaments short, inserted below the middle of the corolla tube; ovules
2, hemispherical, the septum evanescent; style ca. 3 mm long, the stigmas fin-
gerlike, ca. 1.8 mm long, scarcely wider than the style; ovarian disc ca. 1 mm
long, drying black. Fruits rotund, to 0.5 cm in diam., smooth, glabrous, black
when dry.

Faramea papulata is known only from the vicinity of Cerro Jefe, Province of
Panama.

Panama: Cerro Jefe, Dwxcr & Haxdcn 4392 (MO). 9.4 km N of Goofy Lake. Folsom et al. 1977
(MO). Cerro Jefe, Mori 652L 7991, 7998 (all MO); Mori & Kallunki 3630 (MO); Tyson et al. 2218,
3284, 3381 (all MO).

172 ANNALS OF THE- MISSOURI BOTANICAL GARDEN [Vol. 67

12. Faramea pauciflora Dwyer/^ type: Panama, Croat 27575 (MO).

Trees to 4 m tall, the branchlets angular, twisted, somewhat smooth, the nodes
well spaced. Leaves elliptic oblong, 15-19 cm long, 6-7.5 cm wide, rounded or
obtuse at the apex, the acumen slender, to 1 .5 cm long, ultimately obtuse, cuneate
or acute at the base, the costa somewhat prominulous above, subprominent be-
neath, the lateral veins 14-17, arcuate, prominulous, forming an undulate mar-
ginal vein, rigidly papyraceous, somewhat discolorous, minutely puberulent on
the veins and the costa; petioles to 1.5 cm long, ca. 0.13 cm wide, black when
dry, minutely puberulent; stipules free, ovate triangular, to 1.5 cm long, 0.4-0.6
cm wide at the base, subcoriaceous, glabrescent, minutely rugulose, the apex
drawn into a short bristle. Inflorescences axillary, the flowers 3 per cyme, or the
flowers solitary; peduncle to 4 cm long, to 0.15 cm wide, black, glabrous; bracts
and bracteoles not seen, perhaps regularly absent; pedicels perpendicular to the
axis of the inflorescence or sharply ascending, to 4 cm long, ca. 0.1 cm wide,
glabrous. Flowers with hypanthium similar to the pedicel but slightly expanded,
glabrous, to 2.5 mm wide, the calycine cup cylindrical, to 1.2 cm long, to 0.5 cm
wide, conspicuously truncate, the lobes 4, subulate, to 8 mm long, with 2 longer
than the others, acute; corolla white, aromatic, the tube to 3 cm long, ca. 0.25
cm wide in the middle, expanded toward the apex, stiffly petaloid, glabrous on
both sides, the lobes 4, these erect, narrowly subulate, to 1.3 cm long, to 0.35
cm wide, glabrous; stamens 5, sessile, narrowly oblong, ca. 8 mm long, obtuse
at the apex, sagittate at base, inserted above the middle of the corolla tube; style
not seen. Fruits not seen.

Faramea pauciflora is known only from Panama. It is readily distinguished
by its few-flowered inflorescences, its elongate calycine lobes and corolla tube,
and its elongate anthers.

VERAGUAs: Valley Ri'o Dos Bocas, Road toward Escuela Agricola Alio Piedra and Calovebora,
450-550 m, Croat 27575 (MO).

13. Faramea scalaris Standley, Contr. U.S. Natl. Herb. 13: 139. 1916. type:
Panama, Pittier 3265 (US, holotype; isotypes F, NY).

Shrubs 4-6 ft tall, the branchlets smooth, glabrous, the nodes well spaced.
Leaves sessile or subsessile, glabrous, narrowly elliptic, 5.5-15 cm long, 1.3-3.5
cm wide, tapering at the apex, acuminate, the acumen falcate, to 1.5 cm long,
usually ultimately obtuse, basally cuneate to obtuse, the costa prominulous above
and beneath, the lateral veins 8-13, perpendicular, scarcely arcuate, the inner
submarginal vein more or less parallel to the margin, to 4 (-7) mm from the

^'^ Faramea pauciflora Dwyer, spec. nov. Arborcs ad 4 m altae. FoUa elliptico-oblonga, 15-19
cm longa, 6-7.5 cm lata, apice rotunda vel obtusa, acumine ad 1.5 cm longo, basi cuneata, venis
lateralibus 14-17 rigido-papyracea in venis costaque minuto-puberula; petiolis ad 1.5 cm longis; sti-
pulis liberis ovato-triangularibus, ad 1.5 cm latis, apice seta brevi praeditis. Jnflorescentiae axillares
floribus 3 in cymam imam dispositis, pedunculo ad 4 cm longo. Florcs pedicellis ad 4 cm longis;
cupula calycis ad 1.2 cm longa, truncata dentibus 4 subulatis, ad 8 mm longis, inaequalibus; corolla
alba, tubo ad 3 cm longo, lobis 4 ad 1.3 cm longis; antheris sessilibus angusto-oblongis, ca. 8 mm
longis. Fructus non visi.

19801

DWYER— FLORA OF PANAMA (Family 179. Riihiaccae)

173

Figure 39. Fciramea papiilata Dwyer & Hayden.— A. Habit (x'/2).— B. Flower (xl.5).— C
Flower opened to show interior (x 1.5).— D. Fruit (x2). [After Mori 7998.]

margin, prominulous beneath, the outer submarginal vein close to the margin,
less prominulous and conspicuous than the inner submarginal vein, with 1-3
smaller veins paralleling the adjacent lateral veins, these soon branching, stiffly
chartaceous, concolorous, the margin revolute; stipules persistent, free, rectan-
gular or deltoid, to 0.5 cm long with a short median awn arising from a median
keel, stiffly chartaceous, glabrous. Inflorescences terminal, the peduncle 3-4 cm
long, ca. 0.15 cm wide, flowers ca. 15, aggregated into a head subtended by 2

174 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

ovate rotund bracts, to 1.5 cm long, the terminal awn about as long as the body.
Flowers subsessile; hypanthium and calycine cup ca. 2 mm long, the teeth tri-
angular, 0.2-0.4 mm long; corolla bluish white, the tube narrowly cylindrical, 1-
1.2 cm long, ca. 0.1 cm wide, the lobes 4, 0.3-0.5 mm long. Fruit not known.

Faramea scalaris is known only from high elevations in the Province of Chi-
riqui. It may be conspecific with the well-collected F. sucrrcnsis (Donnell Smith)
Donnell Smith. Unfortunately the fruit of F. scalaris is not known.

CHiRiQuf: Bajo Chorro, near Boquete, 6000 ft, Davidson 94 (US), 430 (MO), 436 (US). Between
Alto de las Palmas and Cerro Horqueta, Pittier 3265 (F, NY, US).

14. Faramea suerrensis (Donnell Smith) Donnell Smith, Bot. Gaz. 44: 112. 1907.

F. triiwrvia K. Schum. & Donnell Smith var. siiern-nsix Donnell Smith, Bot. Gaz. 31: 115. 1905.
type: Costa Rica, Donnell Smith 6589, not seen.

Trees to 15 ft tall, the branches stout terete, ultimately angular, glabrous, the
nodes well spaced. Leaves oblong, ovate oblong or elliptic, to 14(30-) cm long,
to 7(10.5-) cm wide, usually IVi times longer than wide, acute to obtuse at the
apex, cuspidate, the cusp often falcate, acute, obtuse or truncate, to 2 cm long,
occasionally vaguely auriculate at the base, the costa prominulous above, prom-
inent beneath, the lateral veins 13-15, usually with a smaller vein intermediate
between 2 laterals paralleling these, at first strict, then arcuate and joining a
prominent marginal vein, 0.5-0.7(1.0-) cm within the margin, usually with a sec-
ond less prominent vein ca. 2 mm from margin the intervenal areas spreading
reticulate, occasionally bullate, chartaceous to subcoriaceous, concolorous, gla-
brous; petioles absent or to 2 cm long, glabrous; stipules connate, cylindrical,
oblong rotund, oblong rectangular to hemispherical, 6-7 mm long, 5-7 mm wide,
minutely mucronate or retuse, glabrous. Inflorescences terminal, cymose cor-
ymbose, glabrous, to 9 cm long, sometimes epedunculate with 2 branches arising
from the tip of the twiglet, sometimes long pedunculate, the peduncle to 5 cm
long and the secondary branches ascending and flabellate or penicillate. Flowers
with the pedicels slender, to 2.5 mm long; hypanthium oblong rotund, ca. 1.2 mm
long, glabrous, the calycine cup rimlike ca. 0.3 mm long, glabrous but with a few
glandular spots within, the teeth minute, to 0.3 mm long; corolla white (in bud),
blue or purple, the tube narrow, to 6 mm long, ca. 1.2 mm wide, petaloid, gla-
brous; stamens 4, the anthers oblong, ca. 2.2 mm long, ca. 0.4 mm wide, minutely
apiculate, the filaments attached near the middle of the tube; ovarian disc oblong
rotund, ca. 1 mm long, the style short, the ovules ca. 0.3 mm in diam. Fruits
reniform or compressed oblong, to 0.7 cm long, to 1.5 cm wide, about twice as

wide as long, apiculate, smooth, often delicately ribbed, farinose or minutely
rugulose.

Faramea suerrensis is known from Costa Rica, Panama, and Colombia (Cho-
co). The fruit of Duke & Lallathin 14977 (MO) is reported as blue.

BOCAS DEL TORo: Punta Pena. vicinity of Chiriquicito, Lewis el al. 2145 (MO). Chiriqui Lagoon,
Wedel 1430 (F). Water Valley, Wedel 1511 (MO, US), 1610 (MO), chiriqui: Branch in road to Cerro
Colorado and Escopeta above Rio San Felix near town of San Felix, ca. 13 mi N of Rio San Felix,
800-1200 m, Croat 33428 (MO), cocle: Cerro Pilon, Duke tC Lallathin 14977 (MO). EI Cope, El

1980] DWYHR— FLORA OF PANAMA (Family 179. /?»/i(«r«/f) 175

Potroso, Atlantic slope of Alto Calvario, Folsom & Lantz IS87 (MO). Cerro Pilon, LaUathin 5093
(MO), colon: Camp Bejita, District Donoso, Holdridge 6251 (MO). Santa Rita Ridge, Mori & Kal-
Iiinki49l9 (MO); Mori & Crosby 6425 (MO), veraguas: 6.4 km outside Santa Fe, road to Agricultural
School, Folxom 2965 (MO). Cerro Tute, Folsom & Edwards 3336 (MO). Colombia: choco: Alto de
Buey, 500-1200 m, Gentry & Forcro 7287 (MO).

15. Faramea talamancarum Standley, Field Mus. Nat. Hist., Bot. Ser. 4: 332.
1929. type: Costa Rica. Carleton 135 (F, holotype, US).

Trees to 15 ft tall, the branches slender, terete, smooth, glabrous, the nodes
well spaced. Leaves oblong to obovate oblong, to 17 cm long, to 6 cm wide,
cuspidate, the cusp to 1.5 cm long, erect or falcate, to 0.25 cm wide, obtuse,
cuneate or acute at the base, scarcely inequilateral, the costa slender, prominu-
lous above and beneath, the secondary veins 10-15, almost perpendicular, the
submarginal vein not elevated or impressed, membranous to chartaceous, con-
colorous; petioles to 1 cm long, glabrous; stipules persistent, connate, the sheath

cylindrical, 2

Infl

paniculate, cymose corymbose, 8-10 cm long, glabrous, sometimes the peduncle
absent and usually 3 branches arising from the stem tip, sometimes 2-3 cm long,
ca. 0.15 cm wide, the branches spreading or strongly ascending, in 2 or more
pairs, opposite, 2-4 cm long. Flowers with the pedicels slender, 8-10 mm long;
hypanthium compressed rotund, to 0.7 mm long, carnose, glabrous, the calycine
cup inflated, glabrous, ca. 0.6 mm long, with several rubescent glandular spots
within, the teeth 4, deltoid, to 0.3 mm long, acute; corolla blue, the tube narrow
cylindrical, expanding apically, 10-17 mm long, the lobes oblong, to 6.5 mm long;
stamens 4, the anthers narrowly oblong, 2.8-4 mm long, 0.6-0.8 mm wide, obtuse,
cordate at the base, the filaments dorsifixed, to 1 mm long, attached above the
middle of the tube; ovarian disc coroniform, to 0.35 mm long, the stigmas 2,
upright, scarcely thicker than the style, 1-1.5 mm long, the style 1.8-3 mm long,
ca. 0.2 mm wide, the ovules about as long as broad. Fruits not seen.

Faramea talamancarum is known from Costa Rica and Panama.

BOCAS DEL toro: Ri'o Teribe, between Quebrada Huron & Quebrada Schlunyik. Kirkhride &
Duke 496 (MO). Duwebdulup Peak, N Rio Teribe, Kirkhride & Duke 584 (MO). Fish Creek Mts,
Wedel2l9l (F, MO), 2243 (MO, US), colon: Santa Rita Ridge, Dwyer 8555 (MO), darien: Airstrip,
Cana gold mine. 480 m. Crocil 37968 (MO). Paya. Stern et al. 230 (MO, US). Cana, Stern et al. 656
(MO).

16. Faramea terreyeae Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 189.
1940. type: Panama, Terry & Terry 1479 (F, holotype, isotype MO).

Shrubs to 8 ft tall, the branchlets terete, smooth, glabrous, the nodes well
spaced. Leaves lanceolate or elliptic to 13 cm long, to 3.6 cm wide, at least 3
times longer than wide, often falcate, acute at the apex, cuspidate or caudate
acuminate for ca. 1.5 cm, ca. 1.5 mm wide in the middle, prominulous above,
subplane and grooved beneath, the lateral veins ca. 20, strict, joining a delicate
and vaguely undulate vein ca. 0.2 cm from the margin, but without a distinctly
elevated submarginal vein, papyraceous, discolorous; petioles to 0.5 cm long,
slender, ca. 0.8 mm wide, glabrous; stipules connate, cylindrical, to 3 cm long,
ca. 0.5 cm wide, the awn ca. 1 cm long. Inflorescences terminal, to 2.5 cm long.

176 ANNALS OF THK MISSOURI BOTANICA[. GARDI-N [Vol. 67

the peduncle ca. 1 cm long, the lowermost bracts folded, ovate rotund or com-
pressed rotund, to 2 cm long, with a slender awn arising from a slender keel-like
thickening, papyraceous, glabrous. Flowers not clearly seen (cf. text below).
Fruits not known.

Faramca terrcycae is known only from the type collection. It is closely related
to F.jefensis Dwyer which has a very reduced inflorescence subtended by 2 large
leaf-like bracts. F. terrcycae has papyraceous leaves whereas those of F,jefensis
are thin coriaceous; the hypanthium and the calyx of F, terrcycae are about 2
mm long, while those of F. jefensis are about 6 mm long. Unfortunately the
flowers of F. terrcycae are poorly known. Standley described the inflorescence
as umbellate and capituliform; he noted that in addition to the 2 outermost large
bracts there are 2 much shorter inner bracts. His description of the floral parts
is meagre (translated from Latin): ''Flowers short-pedicellate; hypanthium widely
obovoid, 1.5 mm long, the calyx ring shaped, ca. 0.6 mm long, truncate, remotely
and minutely denticulate.''

darien: Cana-Cuasi Trail, Chepigana, 3000 ft, Tcny ct Terry 1479 (F, MO).

17. Faramea woodsonii Standley, Ann. Missouri Bot. Gard. 28: 470. 1921. type:
Panama, V^oodson & Schcry 164 (MO, holotype).

Shrtihs, the branchlets slender, terete, smooth, glabrous, the nodes well
spaced. Leaves oblong, 3-7 cm long, 1-2.6 cm wide, tapering obtusely or oc-
casionally acutely toward the apex, acuminate, the acumen 0.4-1 cm long, obtuse,
cuneate to occasionally obtuse at the base, the costa slender, prominulous above
and beneath, the lateral veins ca. 9, at first strict, then arcuate, forming an un-
dulate vein 1-1.5 mm from the margin, 1-2 smaller, irregular, and soon branching
veins arising between the lateral veins, the other conspicuous submarginal vein
delicate, ca. 0.35 mm from the margin, the margin delicately revolute, the inter-
venal areas spreading reticulate, stiffly chartaceous, concolorous, drying gray or
green, glabrous; petioles to 0.4 cm long; stipules connate, the sheath not deeply
incised laterally, ovate rotund, to 4 mm long, glabrous, the median awn of each
part almost as long as the sheath, extending along the body as a median keel.
Inflorescences terminal, the flowers ca, 4, spread radially, the peduncle obsolete.
Flowers to 2 cm long; hypanthium oblong, ca. 1 mm long, glabrous, the calycine
cup ca. 0.5 mm long, with minute glands within, the teeth minute, disposed as 5
points; corolla blue, the tube narrow cylindrical, to 11.5 mm long, ca. 1.3 mm
wide below, ca. 3 mm wide near the mouth, petaloid, glabrous, the lobes 4, ovate
oblong, to 7 mm long; stamens 4, the anthers subsessile, oblong, 3-4 mm long,
attached at the middle of the tube; style subulate, 1.5-3 mm long, the stigmas 2,
minute, ca. 0.5 mm long, the hypanthium angular in cross section. Fruits on
slender pedicels longer than the pericarp, the latter compressed rotund, to 0.8 cm
long, smooth, glabrous, the persistent calyx ca. 0.5 mm long.

Faramea woodsonii is known only from Panama.

coclf': El Valle, Allen 3544 (MO); Duke 12152 (MO); Duke & Dwyer 13914 (MO); Dwyer 11868
(MO); Woodson ct- Schery 164 (MO). Panama: El Llano-Cartf Road, 16-181^ km N of Panamerican
Highway, 400-450, Nee & Tyson 10940 (MO).

1 98l)J

DWYKR— FLORA Of PANAMA (Family 179. Rithiacciw)

\11

Figure 40. Ferdinandusa panamensis Standley & Williams. Habit (x'/2). [After Wedcl 2250.]

38. FERDINANDUSA

Ferdinandusa Pohl, PI. Bras. Icon. Deer. 2: 8. 1828-1829. Substitute name for
Ferdinandca Pohl, 1827. type: F. speciosa Pohl.

Ferdinandca Pohl, Flora 10: 153. 1827, not Fcrdinanda Lagasca, 1816. type; No type indicated.
AspidaiUheni Benth. in Hook., J. Bot. 3: 217. 1841. type: A. rndiieoides Benth.
Gomphnsia Wedd., Ann. Sci. Nat. Bot. 3: 10: 14. 1848. type: No type indicated.

Trees or shrubs. Leaves with stipules interpetiolar. Inflorescences terminal,
cymose paniculate or corymbose paniculate. Flowers 4-5-merous, the calycine
cup cupuliform, the teeth reduced; corolla variable in color, infundibuliform or
hypocrateriform, the lobes contorted in the bud, obliquely 2-lobed, emarginate;
stamens ca. 4, the anthers exserted or included, often variable in size, the fila-
ments often variable in length; ovary 2-celled, the ovules several to numerous in

178 ANNALS OF THI: MISSOURI BOTANICAL GARDEN [Vol, 67

each locule. Fruits capsular, cylindrical to rotund, loculicidally dehiscent, the
seeds few to numerous, peltately affixed to an axile placenta, winged, the wing
occasionally laciniate,

Ferdinandusa, a genus of about 20 species, is found in Panama and in South
America.

1. Ferdinandusa panamensis Standley & Williams, Ceiba 3: 34. 1952. type: Pan-
ama, Wcdel 2232 (F, holotype; isotype, MO), — Fig. 40.

Trees to 60 ft tall, the branchlets terete, smooth, villose, minutely resinous
pustulate (bases of deciduous hairs), the nodes well spaced. Leaves oblong, oc-
casionally falcate, 10-17 cm long, 6.0-9.5 cm wide, obtuse and short acuminate,
the acumen to 1 cm long, obtuse, occasionally truncate, the costa slender, prom-
inulous above, prominent beneath, the lateral veins 12-15, prominulous, arcuate,
stiffly papyraceous, glabrous above and beneath, the intervenal areas reticulate;
petioles to I cm long, stout, villose, the lamina decurrent on the upper side;
stipules deciduous, deltoid lanceolate, ca, 1 cm long, acuminate, glabrous. Inflo-
rescences terminal, longer than the uppermost leaves, the peduncle shorter than
the uppermost leaves, glabrous with 2 branches arising at the tip, the flowers
several, ascending. Flowers with the hypanthium short, subrotund, ca. 2.5 mm
in diam., glabrous, the calycine teeth reduced to 4-5 fleshy points; corolla white,
the tube cylindrical, expanded at the apex and at the base, 1.5-2.3 cm long, 0.14
cm wide in the middle, fleshy to subcoriaceous, glabrous, the lobes 5, imbricate,
oblong, ca. 0.8 mm long, not cucculate; stamens 5, the anthers dimorphic, usually
2 long and 2 short, dorsifixed, the thecae of some anthers dimorphic, oblong to
rotund, the filaments linear, to 7 times the length of the anthers, attached below
the middle of the tube; ovarian disc prominent, the style subulate to 11 mm long,
the stigmas 2, ca. 1.5 mm long, wider than the style, oblong rotund. Fruits not
seen.

Ferdinandusa panamensis is known only from Panama.

BOCAS DtL TORO: Fish Creek, Chiriqui Lagoon, Wedel 2232 (F, MO), 2250 (MO).

39. GALIUM

Galium L., Sp. PI. 105. 1753. Gen. PI., ed. 5. 46. 1754. type: Galium mollu-
go L.

Erect or decumbent herbs or vines, the stems often 4-angled, slender. Leaves
3-4 or more at a node, verticillate; stipules foliaceous. Inflorescences terminal
or axillary, in dichasial panicles, the flowers not subtended by an involucre.
Flowers bisexual, unisexual or polygamous dioecious, pedicellate; hypanthium
articulate with the pedicel; calycine cup ovoid or globose, the lobes reduced or
absent; corolla white, yellow or green, the lobes 3-4, valvate in the bud; stamens
2-4, exserted, the styles 2, the stigmas capitate; ovary 2-celled, the ovules soH-
tary. Fruits didymous, fleshy or dry, separating finally into closed segments or
only one carpel maturing; seeds concave.

Galium is a genus of about 400 species. It is well distributed throughout the
world, especially in the temperate climates.

1980] DWYER— FLORA OF PANAMA (Family 179. /?///7/rt(cac) 179

Literature:

Dempster, L. T. 1973. The Polygamous Species of the Genus Galium (Rubi-
aceae) Section Lophogalium of Mexico and Southwestern United States. U.

Calif. Publ. Bot. 64: 1-36.
. 1978. The genus Galium (Rubiaceae) in Mexico and Central America.

U. Calif. Publ. Bot. 73: 1-33.
Greenman, J. M. 1898. Revision of the Mexican and Central American Species
of Galium and Relhunium. Contr. Gray Herb. n.s. 14: 455-457.

a. Leaves 6-8(-10) at a node 2. G. mcxicanum

aa. Leaves 4 at a node.

b. Leaves with lateral veins evident; fruit with white, uncinate hairs — 3. G. uncinulatum
bb. Leaves with lateral veins not evident: fruit glabrous — L G. aschcnhornii

1. Galium aschenbornii Schauer, Linnaea 20: 70L 1847. type: Mexico, Aschen-
borne 92 (JE, not seen).

Ruhia acuminata Mart. & Gal., Bull. Acad. Roy. Sci. Bruxelles 11: 127. 1844. type: Mexico, Galeotti

2362 (K, holotype).
Galium gemmiflorum Mart. & Gal., Bull. Acad. Roy. Sci. Bruxelles II: 126. 1844. type: Mexico,

Galeotti 2604 (BR, holotype; isotype, K, not seen).
G. galeottiamtm Walp., Repert. Bot. Syst. 6: 17. 1846. typf: not seen.
G. uropctalum Hemsl., Biol. Centr. Amer., Bot. 2: 66. 1881. type: not seen.

M r

Vines, the stems angular, obtusely winged, glandular shiny, glabrescent, often
with a few regularly arranged, retrorse, stumpy hairs. Leaves sessile, 4 per node,
ovate oblong, to 5 mm long, acute, the lateral veins not evident or scarcely
visible, subcoriaceous, glandular shiny above and presumably beneath, with sev-

m

2-flowered, to 1 cm

long. Flowers not seen. Fruits subrotund, to 4 mm in diam., glabrous, drying
black, obviously pedicellate.

Galium aschenbornii occurs in Mexico and Central America. Dempster pro-
vides an excellent description of the flowers and fruits (1978) based on a study
throughout the range of the species: "flowers few, or sometimes many, 1 to
several in imperfect, perfect, or compound dichasia borne on short leafy deter-
minate branches; flowers staminate, pistillate, or perfect, on the same plant, or
sometimes on different plants; corollas rotate, glabrous, usually red or pink, but
sometimes yellow, white, or greenish, the lobes much prolonged at the apex;
fruits glabrous, fleshy, black and wrinkled when mature, 3-7 mm across when
dry."

CHiRiQUi: Volcan de Chiriqui, ca. 7000 /t, Davidson 882 (MO, left-hand side of sheet). Chiriqui
Volcano, Killip 3582 (US).

2. GaliummexicanumH.B.K.,Nov.Gen. Sp.P1.3:337. 1818. neotype (Demps-
ter, 1978): Mexico, Solhrig & Ornduff4540 (UC). Original Humboldt & Bon-
pland type apparently lost. — Fig. 41.

G. inexicaiuim var. ^hihnitum Greenm., Contr. Gray Herb., 14: 458. 1898. type: Mexico, Nelson

583 (US).
G. mexiccinuni var. plutyphyUiim Greenm., Contr. Gray Herb. 14: 458. 1898. type: Mexico. Nelson

3165 (US).

180

ANNAI.S OF THH MISSOURI BOTANICAL GARDtN

I Vol. 67

^ ^^ P

FiGURH 41. Galium tnexicanum H.B.K. Habit {xV^). [After White 206.]

Vines, the stems weak, stramineous, angular, branching profusely, the hairs
stiff, minute, reflexed, uncinate, occasionally weak and spreading, the nodes well
spaced. Leaves sessile or subsessile, usually 6-8(-10) at a node, occasionally 4,
linear oblong or narrowly obovate oblong, 0.4-2.0 cm long, 1.3-2.0 mm wide,
rounded or acute at the apex, tapering narrowly at the base, the costa prominulous
above and beneath, the hairs well spaced, uncinate, rigid, the margin often re-
volute and with stiff, uncinate hairs, the blade glandular shiny, pubescent be-
neath, the hairs with bulbous bases, often deciduous, the blade then granulose.
Inflorescences cymose paniculate, to 2 cm long (? longer). FUnvers minute, 0.6-
1.5 mm long; hypanthium subrotund, ca. 0.35 in diam., pubescent; calycine cup
and lobes absent; corolla white, greenish white, occasionally purple or red, the
tube scarcely measurable or to 0.2 mm long, the lobes 4, (2) ovate oblong, 0.5-

19801

DWYfiR— FLORA OF PANAMA (Family 179. Riihiaceac) 181

0.7 mm long, the tip acuminate, often deflexed adaxially, the acumen to '/a the
length of the body of the lobe, petaloid, dorsally pubescent; stamens 4, the anthers
oblong, ca. 0.2 mm long, the filaments 0.2-0.5 mm long, attached at the junction
of the corolla lobes; style short, the stigmas terminal, capitate. Fruits compressed
rotund, less than 1 mm long, wider than long at maturity, the hairs dense, thick,
falcate, ca. 0.1 mm long.

Galium mcxicanum ranges from Mexico to Panama.

CHiRiQUi: Volcan Bard, E slope, 1 mi from summit, ca. 3000 m, Croat 34905 (MO). El Baru,
above Boquete, 1200-1800 m, DAn\ 9919 (MO). Near Boquete. DAny & DArcy 6463 (MO). E
slope of Volcan de Chiriqui, WNW of Boquete, Daridsc & DAny 10187 (MO). 11 km WNW of
Boquete. Daridsc & DArcv 10224 (MO). Bajo Mono, Boquete, Davidson 595 (MO). Volcan de
Chiriqui. 8000 ft, Davidson 935 (MO). W side of Volcan Baru, oak forest, 2000-2500 m, Mori &
Bolten 7355 (MO). Valley of upper Rio Chiriqui Viejo, Monte Lirio, 1300-1900 m, Seibert 276 (MO).
3 mi N el El Volcan, ca. 5000 ft, Tyson 5721 (MO). Volcan de Chiriqui, White 206 (MO, US). Finca
Lerida to Pena Blanca, 1750-2000 m, Woodson c6 Schery 334 (MO). Potrero Muleto to summit,
Volcan de Chiriqui, 3500-4000 m, Woodson & Schery 443 (MO).

3. Galium uncinulatum DC, Prodr. 4: 600. 1830. type: not seen.

G. ohstipnm Schlecht., Linnaea 9: 592. 1834. rvPF.: Mexico, Schiede 248 (HAL, not seen, isotypes

MO, NY; P, W, neither seen).
G. nelsonii Greenm., Proc. Amer. Acad. Arts 33: 460. 1898. type: Mexico, Nelson 628 (US).

Herbs, the stems slender, weak, 4-angled, much branched, hispid. Leaves
4-nate, subsessile, elliptic oblong, or ovate oblong, 1.5-2.3 cm long, 0.5-1.3 cm
wide, rounded, obtuse or deltoid at the apex, cuneate or rounded at the base, the
apiculum minute but obvious, the lateral veins 2, basal and arcuately ascending,
usually with 2 other more delicate veins arising along the costa, membranous,
hispid on both sides; stipules not seen. Inflorescences terminal or axillary, di-
chotomously corymbose, to 7 cm long; peduncle to 4 cm long, with 2 or more
frequently foliose bracts to Vi the length of the regular leaves, subtending a pair
of branches, the flowers few. Flowers pedicellate, the pedicels 0.1-1.0 cm long;
hypanthium rotund, ca. 0.7 mm in diam., the hairs abundant, strict, white, stiff,
terminally uncinate, to 0.7 mm long, the calycine cup and teeth not visible; corolla
greenish yellow or yellow, the tube short, the lobes 4, ovate or oblong, ca. 1 mm
long, acute, pubescent inside and out. Fruits rotund or compressed rotund, didy-
mous, the body 1-2 mm wide, the hairs dense, often obscuring the body, radiate,
tawny, usually hooked at the apex.

Galium uncinulatum

Mex

America.

chiriqui: E of Boquete, road to Cerro Azul, 1300-1500 m, Croat 26754 (MO). E slope of Volcan

de Chiriqui, 2200-2300 m Davidsc & D'Arcv 10178 (MO). Bajo Mono, Boquete, 4500 ft, Davidson

585 (MO) Volcan de Chiriqui, Boquete, 7000 ft, Davidson 882 (MO). Valley of Rio Chiriqui Viejo,

1300-1900 m, Seihcrt 205 (MO). Rio Chiriqui Viejo Valley, near El Volcan, White 184 (MO). Finca

Lerida, 1750 m, Woodson & Schery 219 (MO).

40. GENIPA

Genipa L., Gen. PI. ed. 5. 87. 1754. lectotype: G. atnericana L.

Trees. Leaves opposite, coriaceous, petiolate; stipules interpetiolar, often per-
sistent, triangular, connate basally. Inflorescence of a single axillary or terminal

182 ANNALS OF THK MISSOURI BOTANICAL GARDEN [Vol. 67

cyme, the flowers several, occasionally solitary. Flowers large, bisexual, 5-6-
merous; calyx campanulate or turbinate, truncate, the teeth absent or of 5-6 short
lobes; corolla short cylindrical, the lobes 5-6, contorted in bud, often longer than
the tube; stamens 5-6, the anthers usually sessile, inserted at the mouth, dorsi-
fixed, exserted; ovary 2-ceIled, the ovules numerous, usually disposed horizon-
tally. Fruits baccate, usually globose, smooth, leathery, the wall thick, the seeds
numerous, large, usually trapeziform, disposed in several rows in a fleshy pulp.

Mexico

Central America and in South America.

a. Leaves drying black; corolla pubescent outside 1. G. amcrkana

aa. Leaves drying brown; corolla glabrous outside.

b. Leaves lanceolate, 3.7-5.2 cm wide, deltoid at the apex; corolla lobes ca. 4.5 mm

wide; fruits ca. 3 cm in diam. 2. G. curviflora

bb. Leaves obovate rotund, obovate or oblong, 3.5-10.0 cm wide, rounded or obtuse at the

apex; corolla lobes ca. I cm wide; fruits 3.5-5.0(-7.0) cm in diam. 3. G. williamsU

1. Genipa americana L., Syst. Nat., ed. 10. 2: 931. 1759. type: not seen
Fig. 42.

Gardenia genipa Swartz, Prodr. Veg. Ind. Occ. 51. 1788. type: not seen.
Genipa oblon^ifolia Ruiz & Pavon, Fl. Peruv. Chil. 2: 67. 1798. type: not seen.
G, grandifolia Pers., Syn. PI. I: 198. 1805. type: not seen.

G. caruto H.B.K., Nov. Gen. Sp. PI. 3: 407. 1820. type: Venezuela and Colombia, Hamholdt &
Bonpland (P, not seen).

G. humilis Veil., FI. Flum. 94. Ic. 2: pi. 142. 1825. type: not seen.

G. puhesccns DC, Prodr. 4: 379. 1830. type: Cuba, de la Osa not seen.

G. harhata Presl., Symb. Bot. Upsal. 2: 13. 1832. type: Brazil, Blanche!, not seen.

Genipa americana var. caruto (H.B.K.) Schum. in Mart., Fl. Bras. 6(6): 352. 1889.

G. excelsa Krause, Bot. Jahrb. Syst. 40: 327. 1908. type: not seen.

G. americana var. caruto fo. grandifolia Chod, & Hassl., Bull. Herb. Boissier ser. 2, 4: 171. 1904.

type: Paraguay, Hassler 7851 (G, isotype).
G. excelsa Krause, Bot. Jahrb. Syst. 40: 327. 1908. type: not seen.

Trees 20 m tall, the twigs terete, ultimately angular, the bark smooth, glabrous,
the nodes well spaced or crowded terminally. Leaves usually strongly ascending,
elliptic, elliptic oblong or obovate oblong, 11-28 cm long, 6-13 cm wide, deltoid
or somewhat rounded toward the apex, acuminate, the acumen short, attenuate
toward the base, the costa prominulous above, prominent beneath, the lateral
veins 1 1-20, arcuate ascending, thin coriaceous to coriaceous, shiny and glabrous
above, often drying black, glabrous to velutinous pubescent beneath, the hairs
dense but short; petioles stout, to 1 cm long; stipules free, ovate, to 1.5 cm long,
acute, coriaceous to thickly scarious, glabrous to puberulent toward the scarious
margins, the margins thickened, pubescent, drawn into a subulate process. Inflo-
rescences terminal, cymose, 4-10 cm long or much longer; peduncle short, the
bracts ovate, 3-4 mm long, acuminate, the branches to 9, few or many flowered.
Flowers to 3.5 cm long, bisexual on pedicels to 1 .2 cm long; hypanthium narrowly
oblong; calycine cup turgid, ca. 0.6 cm long, stiffly carnose, the margin truncate,
the teeth as mere points, glabrous except often with a few hairs marginally;

1 980]

DWYER— FLORA OF PANAMA (Family 179. Riihiaccae)

183

FiGURF, 42. Genipo americana L.— A. Flowering branch (x .45). [After Blum & Dwyer 2477.]—
B. Flower (x2.7).— C. Ovary in cross section (x2.7). [After Bartlclt & Lasser 16465.]— D. Fruit
(X.45). [After Croat 7913.]

corolla cream white, the tube short cylindrical, ca. 1.5 cm long, 0.8-1.0 cm wide,
densely velutinous hairy or villose outside and in, the hairs to 2 mm long, the
lobes 5-6, oblong, 1.5-2.5(-3.0) cm long, longer than the tube, widely deltoid at
the apex, reflexed at anthesis; anthers 5, subsessile, narrow oblong, 8.0-9.5 mm
long, the connective produced for ca. 1.5 mm at the apex, ultimately acute or
obtuse, the filaments attached near the mouth; stigmatic lobes 2, narrowly lan-
ceolate, erect, ca. 7 mm long, partially exserted. Fruits oblong or subrotund, to
9 cm in diam., glabrous, silvery gray outside, the persistent calyx short, giving
a craterlike effect, ca. 1 cm in diam.

184 ANNALS OF THE MISSOURI BOTANICAL GARDIvN |Vol. 67

Gcnipa amcrkana is found in tropical Florida, Mexico, Central America,
West Indies, and tropical South America, extending to Paraguay.

Duke (1968) states that according to the Cuna: 'Tulp of the fruits is edible

raw; it is used to make sweet and fermented beverages; sometimes pickled with

vinegar and onions. The flowers are attractive to bees and the fruits to cattle.

Indians paint their bodies black with the pulp; some OICS personnel claimed it

does improve the complexion and state that it is insecticidal. It is the Tree of
Life' to the Cuna. 'Apples of Immortality' arise from the Earthmother's placenta

and umbilical cord; the juice turns the skin black, thus hiding the individual from
the devils that cause disease. It appears to be bacteriocidal, germicidal, and insect
repellent. It is said to be the only medicine capable of flushing the parasitic catfish
which attacks man south of the study areas. The Cuna have a special song of the

jagua {Gcnipa americana) known as *quanekigala.' " In Panama the most common
name is ''Jagua''; other common names are "Guaytil Blanco'; "Guaitil Blanco';
"Jagua Amarilla"; "Jagua Blanca"; "Jagua Colorado"; and "Jagua de Mon-
tana." It is used throughout its range for construction of buildings, furniture
and carts.

Julian A. Steyermark (The Botany of the Guayana Highland, Mem. N.Y. Bot.
Garden 23: 349. 1972) discusses in detail the typification of Gcnipa americana.

CANAL zonk: Cerro Viejo, K 16 C, Blum 1265 (MO). Fort Kobbe, Duke 3943, 4189, 4259 (all
MO). Farfan Beach, Duke 11734 (MO). Ri'o Mar, Duke 12427 (MO). K-9 Road, Ehhii^er 516 (MO).
Mandingo River, Hayden 57 (MO). Arrafjan, Hayden 112 (MO). Albrook, U.S. Army Tropic Test
site, Hayden I34A (MO). Survival School, Fort Clayton, Kirkhride & EUas 268 (MO). Howard Air
Force Base, SE of Kobbe Beach, Oliver & MacBryde 1867 (GH, MO, US), 1899 (MO). Ancon Hill,
Seihert 409 (MO). Cerro Luisa, vicinity of Pedro Miguel Locks, Stern & Chambers 8 (MO). Survival
School, Fort Clayton, 7v.s7>/? 1051 (MO), hahro Colorado island, Cro^/r 7913 (MO). Cardenal Creek,
Fort Clayton, Tyson 1279 (MO). Miraflores, White 126 (MO), barro Colorado island: Robyns 65-
33 (MO); Shattuck 415 (MO), chiriqui: 12.4 mi N of David, Lewis et al. 728 (GH, MO), cocle: Rio
Hato Airstrip, Blum tt Dwyer 2477 (MO). Penonome, Dwyer 2002 (MO), darien: Rio Ucurganti,
Bristan 1185 (MO). Isla Casaya, Duke 10390 (MO), herrlra: Guarare, Rodriguez 26 (MO), los
SANTOS: 4 mi S of Limon, Correra & Gale^os 66 (MO). Punta Mala. Croat 9755 (MO). Candelaria,
Duke 12453 (MO). Monagre Beach, Tyson et al. 3021 (MO). Panama: Rio Pacora, Bartleti & Lasser
16465 (MO). Cerro Campana, Castillo 2 (MO). Rio Pita, Duke 4763 (MO). Farfan Beach, Duke 1 1734
(MO). Rt'o Mar, Duke 12427 (MO). La Bayoneta, Perlas Archipelago, Dnver 1733 (MO). Tocumen,
Dnyer2894, 4347 (both MO). San Jose Island, Johnston 1358 (MO). Palo Seco, Mejia 20 (MO). Isla
del Espiritu Santo, Tyson 5558 (MO). Isla Taboga, Woodson et al. 1480 (MO), veraguas: Santiago,
Dnyer A Kirkbride 7427 (COL, DUKE, MO, UC), 7429 (GH, MO).

2. Genipa curviflora Dwyer. '*^ type: Panama, Ncc cfe Warmhrodt 10347 (MO,

holotype).

Shrubs or small trees to 4 m, the branchlets angular, smooth, minutely pu-
berulent, the nodes turgid, often well spaced. Leaves lanceolate, 11.5-20.0 cm
long, 3.7-6.8 cm wide, deltoid toward the apex, the acumen short, to 1 cm long,
ultimately obtuse, cuneate toward the base, the costa prominulous and often

**" Genipa eurviflora Dwyer, spec. nov. Frutices vel arhores ramulis puberulis. Folia lanceolata,
M. 5-20.0 cm longa, 3.7-6.8 cm lata, venis lateralibus 7-12, tenui-coriacea supra glabra subtus pu-
berula; petiolis ad 2.5 cm longis; stipulis ovato-trapeziformibus. Infloreseentiae terminales ad 4 cm
longac, floribus ca. 7. Flores pedicellati cupula calycis ad 0.5 mm longa, coriacea glabra dentibus
nullis; corolla tubo angusto-cylindrico, ca. 1.5 cm longo, falcato, glabro lobis 4 carnosis basi unila-
teraliter auriculatis; antheris sessilibus lineari-oblongis, ca. 8.5 mm longis. Fruetus subsessiles rotun-
di, ca. 3 cm in diam., laeves glabri.

I9H()J DWY1:R— FLORA OF PANAMA (Family \79. Rnhiaicac) 185

sulcate above, subprominent beneath, the lateral veins 7-12, at first stiff, then
arcuate toward the margins, uniting to form an undulate submarginal vein ca. 5
mm distant from the margin, prominulous beneath, the intervenal areas spreading,
conspicuously thin coriaceous, concolorous, glabrous above, minutely puberulent
beneath; petioles to 2.5 cm long, ca. 0.2 cm wide, lignose; stipules free, often
deciduous, ovate trapeziform, to 1.7 mm long, acute, at first scarious then some-
what ligneous, puberulent. Inflorescences terminal, to 4 cm long; peduncle absent
or to 0.5 cm long, the flowers ca. 7, perhaps flabellately disposed. FUnvers (in
bud) on pedicels to 1 cm long, rigid, shiny, the calycine cup acetabuliform, ca.
0.5 mm long, coriaceous, glabrous, the margin undulate with lobes absent or 4,
widely triangular, ca. 0.2 mm long; corolla white, the tube narrow cylindrical, ca.
1.5 cm long, falcate, glabrous, carnose, the lobes 4, the buds conspicuously ob-
tuse at the apex, to 1 cm long, ca. 0.45 cm wide, carnose, concave, auriculate
unilaterally basally; stamens 4, the anthers sessile, linear oblong, ca. 8.5 mm
long, ca. 0.8 mm wide, obtuse toward the apex, attached above the middle of the
tube; ovarian disc coroniform, ca. 1 mm long, black when dry; style slender, to
1.2 cm long, plane, the stigmas 2, narrowly triangular, ca. 1 mm long, acute,
scarcely wider than the style, pubescent, the ovary sometimes lacking, the ovar-
ian wall thin, the locule ample, the ovules numerous. Fruits subsessile, rotund,
ca. 3 cm in diam., smooth, glabrous, purple black when dry, the calycine scar
ringlike, ca. 0.6 cm in diam.

Genipa curviflora is known only from Panama. Few flowers were available
for dissection and these are in the bud stage. On dissecting two buds of Duke
10678 I found that the locule of the ovary is located at the proximal end of what
appears to be the pedicel. Two buds of Nee 10347 showed no ovary in the
presumed pseudopedicel, although the style and stigmas were evident. Thus, it
seems advisable to call the structure in the latter a true pedicel and its counterpart
{Duke 10678) a pseudopedicel.

ifl

wiUiamsii Standley, known only

from Panama, being distinguished by its narrower leaf blades which taper some-
what at the apex and which are only thin coriaceous. The fruits are smaller than
those of G. nilliamsii.

darien: Camp Summit between Morti and Sasardi, 1800 ft, Duke 10678 (MO). Panama: El
Llano to Carti Road, ca. 14 km N of Panamerican Highway, 350-500 m, Folsom & Kaiikc 1403 (MO);
Folsoin et al. 1475 (MO). El Llano-Carti Road, 8 km N of Panamerican Highway, 450 m. Nee &
Warmhrodt 10347 (MO).

Wash

Willia

Trees 4-20 m tall, the twiglets stout, ultimately 4-angled, the bark rough,
smooth, glabrous, the nodes moderately well spaced, the leaf scars triangular
rotund, ca. 5 mm in diam. Leaves oblong rotund to obovate oblong, to 33 cm
long, to 16 cm wide, rounded or obtuse at the apex, not acuminate, cuneate,
rounded or obtuse at the base, the costa slender, prominulous above, prominent
beneath proximally, the lateral veins 9-12, arcuate, with an undulate, submarginal
vein well set in from the margin, coriaceous, scarcely discolorous, drying a deep
brown, glabrous; petioles 0.5-6.0 cm long, stout, to 0.4 cm wide, rigid, lignose.

186 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

often curved basally; stipules deciduous, presumably rhomboid oblong, to 3.2 cm
long, to 1.8 cm wide, striate, coriaceous, glabrous except ciliolate, thickened
basally. Inflorescences terminal, the flowers 3-4(-7) on a suppressed peduncle,
to 1 cm long, the bracteoles triangular, to 3 mm long. Flowers perhaps monoe-
cious, subsessile or pedicellate, the pedicels to 1 cm long, ca. 0.2 cm wide, drying
black, glabrous; buds ca. 5 cm long, clavate, the mass of corolla lobes oblong,
to 2 cm long, ca. 1.3 cm wide; hypanthium often blending in with pedicel, tur-
binate, ca. 5 mm long, drying black, glabrous, the calycine cup scarcely measur-
able, platelike, with 5 vague lobes, compressed rotund, ca. 1 mm long, wider
than long, glabrous, coriaceous; corolla white, the tube 1.8-3 cm long, ca. 0.5
cm wide in the middle in bud, to 1 cm wide in open flowers, coriaceous, glabrous
outside, the lobes 5, imbricate, triangular oblong, to 2.1 cm long, to 1.5 cm wide,
glabrous outside, often with a few scattered white hairs within; male flowers with
anthers sessile, 1.8-2.0 cm long, ca. 1 mm wide, attached below the mouth;
rudimentary style linear, ca. 2.5 cm long, ca. 1 mm wide; stigmas 4, unequal (at
least one of 4), oblong, to 4 mm long, acute with a few hairs adaxially. Fruits
rotund, to 7 cm in diam., smooth, glabrous, drying black, the wall to 1 cm thick,
the seeds numerous, piano compressed, oblong or subrotund, to 1 cm long, yel-
low, embedded in a white pulp.

Genipa williamsii is known only from Panama and Colombia. This species is
easily recognized by a combination of characters: leaves rounded and obtuse at
the apex, drying brown; buds elongate and clavate at the apex; fruits very large
and drying black. I have not seen female flowers. One of the striking floral char-
acters is the fact that the flower buds are elongate and clavate while the open
flowers appear much shorter than the buds. According to Duke (8026) the ''fruit
(is) very bitter.''

Panama: coci.e: Furlongs Finca, Cerro Pilon, 880 m, Croat 37571 (MO). 3 km NE of El Valle,
Mori ct KalUinki 2959 (MO). La Mesa, 4 km N of El Valle, 875 m, Nee & Dwyer 9152 (MO), darihn:
Cana, Williams 814 (US). Panama: Cerro Jefe, Duke 8026 (MO). Cerro Azul to Cerro Jefe, Dwyer
504 1 A (MO). 9.4 km N of Goofy Lake, 900 m, FoLsom et al, 1987 (MO). 4.8 km N of highway, W of
El Llano, Cerro Jefe, Gentry 5517, 6138 (both MO). El Llano-Carti Road, 12.7 km from Panamerican
Highway, Mori et al. 4688 (MO). El Llano-Carti' Road, 8-11 km from Panamerican Highway, Mori
7712, 7717 (both MO). E slope of Cerro Jefe, Tyson 3397 (MO). Cerro Jefe, Tyson et al. 4441 (MO).
COLOMBIA: CHOCo: Municipio de Novita. Vereda Llanadas. Ladera norte del Cerro Torra. Filo al
Oeste del Rio Surama. Camino al Alto del Oso, 600-900 m, Forero et al. 3082 (MO).

41. GEOPHILA

Geophila Don, Prodr. Fl. Nepal. 136. 1825. Nomen conserv, type: G. reniformis
D. Don, nom. illeg. = G. herhacea (Jacq.) Schumann.

Carinta E. F. Wight, Contr. U.S. Natl. Herb. 9: 216. 1905. type: Psydiotria herhacea Jacq. =

Geophila herhacea (Jacq.) Schumann.
Geocardia Standley, Contr. U.S. Natl. Herb. 17: 444. 1914. type: not indicated.

Creeping herbs. Leaves often cordate, membranous; petioles elongate. Inflo-
rescences axillary, usually pedunculate, the flowers few in a terminal cluster.
Flowers with the calycine tube dentate or lobate; corolla funnelform, pilose in
the throat, the lobes valvate; stamens 4-7, the filaments slender, inserted in the
throat; ovary 2-celled. Fruits fleshy, drupaceous, the 2 nutlets piano convex,
compressed dorsally, usually costate.

19801 DWYER— FLORA OF PANAMA (Family 179. Ruhicueae) 187

Geopfula, a genus of about 30 species, occurs throughout the tropical areas
of the world.

Literature:

Steyermark, J. A. 1977. A new member of the Rubiaceae from Panama. Phy-
tologia 35: 401-402.

a. Leaf blades with auricles not overlapping at the base; bracts lanceolate or lance oblong,
b. Leaves with abundant cystoliths beneath, the blades 2.5-6.0 cm long, 2.0-4.5 cm

wide; peduncles 0.1-0.2 cm long at anthesis, glabrous 1. G. croatii

bb. Leaves lacking cystoliths beneath, the blades 2-9 cm long, 1.0-8.5 cm wide; pedun-
cles 0.8-5.0(-8.0) cm long at anthesis, puberulous 2. G. macropoJa

aa. Leaf blades with auricles overlapping at the base; bracts subulate 3. G, rcpens

1. Geophila croatii Steyermark, Phytologia 35(6): 401-402. 1977. type: Panama,
Croat 6647 (MO, holotype).

Creeping herbs, glabrous, conspicuously rhizophorus at the nodes, the roots
fibrous. Leaves widely ovate or lance oblong, 2.5-6.0 cm long, 1.5-4.5 cm wide,
acute or acuminate at the apex, cordate at the base, the lateral nerves 4-5, prom-
inulous beneath, uniting to form a vein 1-5 mm from the margin, membranous,
glabrescent or the costa and the veins with a few hairs with abundant cystoliths
beneath; petioles 0.5-9.0 cm long, glabrous to puberulent; stipules ovate, ca. 3
mm long, obtuse, glabrescent. Inflorescences capitate umbellate, the peduncle at
anthesis 1-2 mm long, elongating in fruit, to 20 mm long, the flowers usually 5-
7 in a head; bracts lance oblong or lanceolate, 3.0-4.5 mm long, 1.0-1.5 mm
wide, glabrous; pedicels less than 1 mm long at anthesis, to 3 mm long in fruit,
glabrous. Flowers with hypanthium and calyx 2.0-4.5 mm long, the calycine
lobes 5, narrowly oblanceolate or lanceolate, 2.5-3.5 mm long, much longer in
fruit, glabrous; corolla white, ca. 5 mm long, with hairs clustered below filament
attachment, the lobes slender; stamens with filaments connate below. Fruits
ovoid or globose, 3-4 mm in diam., red, often in a cluster of 3; seeds oblong, to
4 mm long, plane on one side, somewhat twisted, the ribs 3, faint.

Geophila croatii is known only from Panama.

CANAi zone: Barro Colorado Island, Croat 6647, 10989 (both MO). Gaillard Highway between
Summit Garden and Gamboa, Sullivan 27 (MO).

2. Geophila macropoda (Ruiz & Pavon) DC, Prodr. 4: 537. 1830.

Psychotria luavropoda Ruiz & Pavon, FI. Peruv. Chil. 2: 63. 1799. type: Peru, Ruiz. & Pavon, not

seen.
Coccocypsclum macropodum (Ruiz & Pavon) Britton, Bull. Torrey Bot. Club 18: 107. 1891.
Cephaclis violacfolia H.B.K., Nov. Gen. Sp. PI. 3: 379. 1818. typh: Colombia, between Barthomome

and Nares, Humboldt l^ Bonpland, not seen.
Geophila violacfolia (H.B.K.) DC, Prodr. 4: 537. 1830.
Geophila herhacea violacfolia (H.B.K.) Chod. & Hassl., Bull. Herb. Boissier II. 4: 180. 1904.

Gcocardia violacf>lia (H.B.K.) Standley. Contr. U.S. Natl. Herb. 17: 445. 1914.

Creeping herbs, the stems weak, glabrous, often farinose. Leaves 1-4 at a
node, cordate or ovate cordate, 2-9 cm long, 1 .0-8.5 cm wide, occasionally wider
than long, acute at the apex, cordate at the base, the auricles not overlapping,
occasionally inequilateral, the costa plane above, prominulous beneath, the lateral

188 ANNALS OF THE MISSOURI BOTANICAL GARDLN [Vol. 67

veins 4-6, the intervenal areas reticulate, papyraceous, glabrous above, glabres-

cent beneath, the cystoliths not evident; petioles weak, flexuous, 0.8-14.0 cm
long, rubescent, pubescent; stipules not seen. Inflorescences with peduncles 0.3-
3.5 cm long, much longer in fruit, puberulous, the flowers few, aggregated ter-
minally in clusters to 0.8 cm wide; bracts lanceolate, to 3 mm long. Flowers with
the hypanthium rectangular, ca. 1.5 mm long, 1 mm wide, the calycine cup ca.
1 mm long, stiffly petaloid, glabrous outside and inside, with a few glands proxi-
mally within, these separate or clustered in 2's or 3's, oblong, ca. 0.1 mm long,
mostly obtuse, the teeth 5, linear lanceolate, slightly unequal, 2.2-3 mm long, to
0.5 mm wide; corolla white, not seen; stamens not seen; ovarian disc hemispher-
ical, to 0.8 mm long, smooth and glandular shiny, the style slender, to 3 mm long,
glabrous, the stigmas 2, digitiform, ca. 0.8 mm long. Fruits (usually 2 maturing),
ovoid, to 1 cm long, glabrous, the calyx persistent, 1-2 mm long.

Geophila macropoda occurs in Costa Rica, Panama and Colombia. ''Nomas
Tuwat^^ (Cuna, fide Duke 14317).

BOCAS DhL TORo: Banana farm, Armuelles, Dickson 2292 (MO). Vicinity of Nievecita, Woodson
et id. 1851 (F, MO), darien: Rio Pirre, trail from EI Real to Tucutf, Duke 5201 (MO). Panama: Piria-
Canasas Trail near Piria, Duke 14317 (MO), san blas: Headwaters, Rio Mulatupo, Elias 1739 (GH,
MO). 3-4 hours up Rio Mulatupo, Kirkhridc 217 (MO).

3. Geophila repens (L.) Johnston, Sargentia 8; 281, 1949, — Fig. 43.

Rondidctlo repens L., Syst. Nat. ed. 10. 928. 1759. type: Jamaica, not seen.
Psychotria repens (L.) L., Amoen. Acad. 5: 377. 1759.

P. herhaeea Jacq., Enum. PI. Carib. 16. 1760. type: Martinique, not seen.

Cephaelis renlforinis H.B.K., Nov. Gen. Sp. PI. 3: 377. 1819. type: not seen.

GeophUa cordata Miq., Linnaea 17: 72. 1843. type: Surinam, Focke 471.

Mapour'ui cordata (Miq.) Mull.-Arg., Mart. Fl. Bras. 6(5): 426. 1881.

A/, herhaeea (Jacq.) Mull.-Arg., Mart. Fl. Bras. 6(5): 427. 1881.

Geophila herhaeea (Jacq.) Schum., Engl. & Prantl. Nat. Pflanzenfam. 4. 4: 119. 1891.

Geocardia herhaeea (Jacq.) Standley, Contr. U.S. Natl. Herb. 17: 444. I9I4.

Carinta repens (L.) Smith & Downs, Sellowia 7: 88. 1956.

C. repens (L.) Brem., Verb. Kon. Nederl. Akad. Wetens 54: 118. 1963. Redundant combination.

Carinta repens var. amerieana Brem., Verb. Kon. Nederl. Akad. Wetens, 119. 1963. type: not seen.

Prostrate herbs, rooted at the nodes, the stems weak, the nodes well spaced.
Leaves cordate, occasionally ovate rotund, (1.2-)1.7-4.5(-5.0) cm long, 1.3-4. 2(-

5.0) cm wide, obtuse, widely deltoid or rounded at the apex, basally cordate,
auriculate, the auricles overlapping, the lateral veins 5, the 3 proximal veins
arising from a common point, prominulous beneath, the veinlets inconspicuous,
glabrous to scabridulous above, glabrous beneath except on the costa, usually
with cystoliths on the lower surface; petioles 0.6-4.5(-6.0) cm long; stipules not
seen. Inflorescences capitate with the peduncle 0.8-5.0(-8.0) cm long, glabrous
to papillate at anthesis, the flowers few; bracts subulate, to 5 mm long, pilose,
the outermost often with 2 short lateral lobes at the base. Flowers (in bud) with
the hypanthium oblong or rectangular, ca. 1 mm long, glabrous, the calycine cup
funnel shaped, ca. 0.8 mm long, glabrous outside and inside, eglandular within,
the calycine lobes 5, subulate or lanceolate, ca. 3 mm long, slightly unequal, to
1 mm wide at the base, pubescent marginally; corolla white or orange, clavate in
bud, to 7 mm long (10 mm long, fide Johnston), fleshy, petaloid, pubescent es-
pecially at the apex outside, glabrous and with numerous raphides within; stamens

1980]

DWYER— FLORA OF PANAMA (Family 179. Riihlaccac)

189

Figure 43. Gcopliila rcpcns (L.) Johnston. Habit (x 1). [After Tyson & Blum 1658.]

5, the anthers oblong, ca. 1 mm long, obtuse; style slender, the stigmatic lobes
2, short. Fruits red, ovate oblong, to 0.5 cm long; calyx cup and lobes persistent,
to 3.5 mm long; seeds twisted and rugulose.

Geophila rcpens extends from Mexico throughout Central America, and in
the Guianas, Venezuela, Colombia, south to Argentina. It also occurs in the West

190 ANNALS 01- THE MISSOURI BOTANICAL GARDLN [Vol. 67

Indies. Duke 14345 notes that the herb is used medicinally by the Bayano Cuna
Indians, who call it, ''Nomas Puruiwat/'

BOCAS DEL TORo: Changuinola, Dunlap 565 (F). Chiriquicito to 5 mi S along Rio Guarumo, Lewis
et al. 2097 (MO). Pumpkin River, Wcdcl 2577 (MO), canal zonl: 2 mi from Gatiin Locks, Blum
1491 (MO). Summit Garden, Mori c£ Kallunki 1885 (MO), Balboa, StanJley 25503 (MO). Old Fort
Lorenzo, Tyson 2207 (MO). Gamboa, Tyson 4570 (MO). Fort Sherman, Tyson & Blum 1658 (MO).
Miraflores Lake, White 252 (F). Salamanca Hydrographic Station, Rio Pequeni, Woodson et al. 1584
(F, MO). Vicinity of Summit Garden, Witherspoon & Witherspoon 8700 (MO), barro Colorado
island: Bailey & Bailey 3 (F); Bangham 558 (F); Bartlett & Lasser 16738 (MO); Croat 4347, 5834.
6464. 7976, 10I7L 10382, 15121 (all MO); Ehin^er 230 (F, MO); Fosher^ 48163 (MO); Hayden 153,
1030 (both MO, UC, VEN); Salvoza 967 (F); Shattnck 130 (F, MO); Starry 286 (F, MO); Woodson
& Schery 986 (MO). CHiRiQuf: San Bartolome, Burica Peninsula, Woodson & Schery 919 (F, MO).
darien: Rio Chico, from junction with Rio Chucunaque to ca. 1 hour by outboard, Burcli et al. 1080
(MO). Pinogana near El Real, Croat & Porter 15477 (MO). 10 mi S of El Real, Ri'o Pirre, Duke 5447
(MO). Rio Pucro to Quebra Maskia, Duke 13055 (MO). Quebrada ^Tomachimuricate'^ Correa Casa
de Bartolo, Kennedy 2854 (MO). Rio Tuquesa, middle of Tuquesa Mining Co., camp called Charco
Peje, Mori 7054 (MO). Paya, Rio Paya, Stern et al. 431 A (MO), los santos: Headwaters Rio Ped-
regal, 25 mi SW of Tonosf, Lewis et al. 2903 (MO). Panama: EI Llano and Rio Mamonf, Duke 5553
(MO). Piria Canasas Trail near Piria, Duke 14345 (MO). Arraijan, Woodsim et al. 769 (MO).

42. GONZALAGUNIA

Gonzalagunia Ruiz & Pavon, Fl. Peruv. Prodr. 12. 1794. type: G. dcpendens
Ruiz & Pavon.

Dug^ena West, Bidar. Ste. Croix 269. 1793. Nomen nudum, type: Z>. richardii, based on Barleria

hirsuta Jacq.
Buena Cav., Anales Hist. Nat. 2: 278. 1800, not Pohl 1827. type: B. panamensis Cav.
Gonzalea Pers., Syn. PI. 2: 132. 1805. New name for Gonzalagunia Ruiz & Pavon.

Shrubs or small trees, the branchlets slender. Leaves petiolate or subsessile;

r

Stipules free, usually triangular subulate. Inflorescences terminal, elongate, slen-
der, spiciform or thrysiform. Flowers with the hypanthium rotund, the calyx with
4-5 small, persistent lobes; corolla funnelform or salverform, the tube villous,
the lobes 4-5, valvate or imbricate; stamens 4-5, the filaments slender, attached
to the tube; anthers slender, included; ovarian disc crenulate, the style slender,
the stigmas small, 2-4-lobed, the ovules numerous, the placentas small. Fruits
baccate, globose, 2-4-suIcate, the pyrenes 2 or 4, chartaceous to bony; seeds
numerous, minute, subglobose, deeply foveolate.

Gonzalagunia. a genus of about 25 species, is common throughout the Amer-
ican tropics and the West Indies. It is among the easiest genera of the Rubiaceae
to recognize because of its pendent spiciform inflorescences.

Literature:

Donnell-Smith, J. 1899. Undescribed plants from Guatemala and other Central

American Republics. Bot. Gaz. 27: 336.
Standley, P. 1921. New Rubiaceae from Colombia and Costa Rica. Contr. U.S.

Natl. Herb. 17: 446.

. 1929. Studies of American plants II: Rubiaceae. Field Mus. Nat. Hist.,

Bot. Sen 4: 278-281

a. Leaves 10-13 cm long; lateral veins of larger leaves 6-12.

b. Largest bracts 5-8(-12) mm long: flowers blue; fruits dicoccus 1. G. bracteosa

19801 DWYFR— FLORA OF PANAMA (Family 179. Ruhiaccae) 191

bb. Largest bracts 3-5 mm long: flowers white or red; fruits 4-coccus.

c. Leaves subsessile or with petioles to 0.3 cm long.

d. Stipules subulate. 8-9 mm long: calycine lobes ca. 2 mm long, obtuse —

3. G. ovatifolia

dd. Stipules ovate or deltoid, ca. 6 mm long; calycine lobes ca. 0.7 mm long,

acute 6. G. riidls

cc. Leaves with petioles 0.5-2.0 cm long.

e. Leaves 7-22 cm long: stipules 8-10 mm long, corolla pink, the tube ca. 8

mm long, densely appressed villose within 5. G. rosea

ee. Leaves 5-13 cm long: stipules ca. 3 mm long; corolla white, the tube 10. 0-

13.5 mm long, glabrous within 4. G. panaincnsis

aa. Leaves 15-24 cm long; lateral veins of larger leaves 12-16 (except G. rosea).
f. Lateral veins of larger leaves 12-16; flowers white.

g. Flowers pedicellate 2. G. kalliinkii

gg. Flowers sessile or subsessile 7. G. vera^uensis

ff. Lateral veins of larger leaves 6-12; flowers pink 5. G. rosea

1. Gonzalagunia bracteosa (Donnell Smith) Robinson, Proc. Amer. Acad. Arts
45: 405. 1910.

Gonzalea bracteosa Donnell Smith, Bot. Gaz. 33: 252. 1902. tyim.: Gunlemala, Donnell Smith 6583 (F.

NY).
Diiggena bracteosa (Donnell Smith) Standley, Contr. U.S. Natl. Herb. 18: 125. 1916.

Shrubs 1.5-2.0 m tall, the lowermost branches occasionally prostrate, the
branches smooth, glabrous to villose, wiry, to 0.15 cm in diam., the nodes often
swollen, well spaced. Leaves oblong, obovate oblong or narrowly elliptic, 8-
18(-22) cm long, 2.5-8.5 cm wide, widely deltoid or scarcely obtuse at the apex,
short acuminate, the acumen to 1.5 cm long, acute, subcuneate or obtuse at the
base, the costa prominulous above, prominent beneath, the secondary veins 6-
9(-l 1), strongly arcuate ascending, the intervenal areas spreading reticulate, char-
taceous, pubescent, the hairs 1.0-1.5 mm long, appressed, white, more dense
beneath; petioles short or absent, to 0.5 cm long; stipules connate, cylindrical,
the sheath closely appressed to stem, thin, densely golden villose or hirsute,
occasionally carinate, the hairs more dense along the keel; each division tapering
into an acute awn to 5 mm long. Inflorescences axillary, pendent, (5-) 15-25 cm
long, of ca. 20 flower clusters well spaced along an often flexuous rachis; pedun-
cles 4-9 cm long; bracts several subtending the mass of flowers, deflexed, nar-
rowly ovate lanceolate, 5-8(-12) mm long. Floners sessile, the hypanthium and
calycine cup subrotund, ca. 1.2 mm long, carnose, densely villose, the calycine
lobes 5, to 0.5 mm long, densely villose outside, glabrous within, with a minute
gland at the base of adjacent lobes; corolla blue (in bud), to 4 mm long, the tube
apparently glabrous outside, villose within, the lobes 5, villose outside and within;
stamens 5, the anthers oblong, to 1 mm long, minutely acuminate, affixed near
the orifice; style and stigmas to 2.8 mm long, piano compressed, the style slender
along most of its length, expanding above to 0.7 mm wide, the style and 2 stig-
matic lobes sagittate, the lobes to 0.4 mm long; ovarian disc compressed cylin-
drical, to 0.2 mm long.

colon: Rio Guanche. ca. 3-5 km above bridge on Portobelo Road, Croat 37012 (MO). Rio
Guanche, ca. 2.5 km upriver from bridge, road to Portobelo, Mori et al. 6470 (MO). dariiIn: Cocalito,
Dnyer4S60 (MO). Cana-Cuasi Trail (Camp 1). Chepigana. Terry & Terry 1621 (F. MO). Panama:
Campo Tres, 3 mi NE of Altos de Pacora, 500-800 m, Croat 22701 (MO, US). El Llano-Carti Road,
Croat 33763 (MO); DArcy 10613 (MO); Liesner 1201 (MO, US); Mori et al. 4588 (MO); Mori &
KalliinU 5587 (MO).

192 ANNALS OF THI- MISSOURI BOTANICAL GAKDLN [Vol. 67

2. Gonzalagunia kallunkii Dwyer.''" type: Panama, Mori & Kallunki 3141 (MO,
holotype).

Shrubs to 2 m tall, the twiglets terete, smooth, the hairs short, dense, golden,
the nodes well spaced, the pith occasionally porate. Leaves elliptic oblong, 17-
24 cm long, 5.0-1 1.5 cm wide, acute, short acuminate, the acumen short, widely
triangular, to 1.5 cm long, often slightly falcate, basally acute, cuneate to atten-
uate acute, the blade slightly decurrent on the petioles, the costa prominulous
above, prominent beneath, the lateral veins 12-14, arcuate, the undulate sub-
marginal vein to 3 mm from the margin, membranous, discolorous, dark red
brown above when dry, dull gray green beneath, the hairs numerous, often ap-
pressed strigose, sometimes glabrescent; petioles to 2 cm long, rigid, lignose;
stipules free, triangular, to 8 mm long, acute, with a median and densely bearded
keel, the hairs golden. Inflorescences densely pubescent, to 35 cm long, to 2 cm
wide: peduncle to 6 cm long, to 0.12 cm wide at the base. Flowers with pedicels
to 0.5 cm long, the hypanthium and calycine cup compressed rotund, to 3 mm
long, truncate, densely hairy, the calycine lobes 4, triangular, 1-1.5 mm long,
hirsute outside, glabrous within; corolla white (in bud), the tube slender, 5.5-7.0
mm long, densely villose outside, glabrous within; stamens 4, subsessile, the
anthers to 1.3 mm long, attached at base of tube; style 3 mm long, the stigmas
ca. 0.7 mm long. Fruits white, rotund, ca. 0.5 cm in diam., densely appressed
villose.

GonzciUigunia kallunkii is known only from Panama. It is named in honor of
Jacquelyn Kallunki, who has contributed much to the collection of valuable her-
barium material in Panama. It is closely related to G. rosea Standley which has
pink flowers, fewer lateral veins, and glabrate fruit.

VLRACiUAs: Valley of Dos Bocas, between Alto Piedra and Calovebora, 350-400 m, Croat 27462.
27733 (both MO). NW of Santa Fe, 1 1 km from Escuela Agricola Alto de Piedra, Mori & Kallunki
314! (MO).

3. Gonzalagunia ovatifolia (Donnell Smith) Robinson, Proc. Am. Acad. Arts 45:
405. 1910.

Gonzalea ovatifolia Donnell Smith, Bot. Gaz. 27: 336. 1899. syntyphs: Costa Rica: Donnell Smith

6599, 6600 (both F); Pilticr A Tondtiz 9171: Panama: Hart 113.
Dii^gcna ovatifolia (Donnell Smith) Standley, Contr. U.S. Natl. Herb. 18: 126. 1916.

S/iruhs to 3 m tall, the branchlets pubescent. Leaves subsessile, ovate oblong,
to 10 cm long, to 5 cm wide, acuminate, the lateral veins (6-)8-12, discolorous,
dark chocolate color above, lighter beneath, glabrescent above, sericeous be-
neath; stipules subulate, 8-9 mm long, triangular at the base, 2-3 mm wide.

^^^ Gonzalai^ania kallunkii Dwyer» spec, nov. Fruticcs ad 2 m aiti, ramulis laevibus aureo-pu-
berulis. Folia elliptico-oblonga, 17-24 cm longa, 5.0-11.5 cm lata, acuta, acumine ad 1.5 cm longo,
venis lateralibus 12-14, membranacea glabrescentia vel ciliis crebris appresso-strigosis; petiolis ad 2
cm longis; stipulis ad 8 mm longis carina media dense aureo-barbata ornatis. Inflorcsccntiae ad 35
cm longac floribus rare solitariae plerumque in fasciculis florium paucorum sessiles vel subsessiles
bene distantes separatis. Florcs hypanthio dense puberulo. rotundi. ca. 3 mm diam.; corolla alba tubo
ad 7 mm longo extus ciliis appressis elongatis albis praedito, lobis ad 2.5 mm longis intus glabris.
Fructas rotundi ad 0.5 cm diam., dense hirsuti.

19801 DWYER— Fl.ORA ()l- PANAMA (Family 179. Ruhiaccac) 193

Inflorescences terminal, (8-) 1 7-27 cm long with peduncles 2-3 cm long or seem-
ingly sessile and subtended by regular-sized leaves, the cymules 1-5-flowered,
sessile, the bracts subulate, 3-5 mm long. Flowers with the hypanthium com-
pressed rotund, ca. 1.5 mm long, the calycine lobes 4, oblong, ca. 2 mm long,
usually curved, ciliate, with 1 to several rotund glands at the junction of the lobes;
corolla white, the tube to 2 cm long, strigose, the lobes 4, obtuse, often as long
as the tube; stamens 4, not exserted; ovarian disc pulvinate and bearing a few
hairs, the style with the 4 stigmas included. Fruits tetracoccus, to 1 cm in diam.
with white flesh persisting, hirsute, drying white ca. 3 mm in diam.; seeds angular,
honeycomb reticulate.

This species is known only from Costa Rica, Colombia and Panama. Donnell
Smith (1899) cited Hart 113, collected in Laguna de Chiriqui (Bocas del Toro),
Panama in 1895.

BOCAS DEL TORo: Changuinola Valley, Diinlap 102 (F), 496 (F, BM). Without other locality,
Wecii'l 841 (MO), cocle: La Mesa above El Valle. Folsom 3653 (MO), san blas: Mainland opposite
Achituppu, Lewis et at. 100 (F, GH, MO). Colombia: choco: Trail between Alto Coriche and Camp
Curiche, E of Boca Curiche, 10-400 ft, Diikc & Idroho 11306 (NY).

4. Gonzalagunia panamensis (Cav.) Schum. in Mart., Fl. Bras. 6(6): 292. 1889.
Fig. 44.

Biicmt panamensis Cav., Anales Hist. Nat. 2: 279. 1800. type: Panama, Nee (?MA).

Gonzalea cornifolia H.B.K., Nov. Gen. Sp. PI. 3: 416. 1820. type: not seen.

G. panamensis Spreng., Syst. Veg. 1: 417. 1825. type: Panama, Seemann 351 (BM, not seen: photo,

MO).
G. leptantha A. Rich, in Sagra. Hist. Cuba 11: 16. 1850. type: Cuba, Ramon cle la Sai^ra. not

seen.
Gonzalagunia leptantha Maza, Anales Soc. Esp. Hist. Nat. 23: 289. 1894. type: not seen.

Gonzalea liavesii Wernham, J. Bot. 51: 219. 1913. type: Panama, Hayes (BM).

Duggena liavesii (Wernham) Standley, Contr. U.S. Natl. Herb. 18: 125. 1916.

D. leptantha (A. Rich.) Standley, Contr. U.S. Natl. Herb. 18: 125. 1916.

Shrubs to 10 ft tall, the branchlets terete, smooth, viliosulose, the nodes well
spaced, mostly 3-6 cm apart. Leaves elliptic or ovate elliptic, 5-13 cm long, 1-
4(-6) cm wide, acute at the apex, usually acuminate, the acumen to 2 cm long,
cuneate to subobtuse at the base, often markedly inequilateral, the costa prom-
inulous above and beneath; petioles slender, 1.0-1.5(-2.0) cm long, ca. 0.8 mm
wide, pubescent; stipules ovate oblong, widely triangular to subulate, to 3 mm
long, the awn usually evident, weak, as long as or longer than the body. Inflo-
rescences terminal, solitary, to 31 cm long, the peduncles flagelliform, 1.5-4.0
cm long, to 0.2 cm wide, pubescent, the cymules sessile or subsessile, 0.5-1.0
cm apart; bracts linear subulate or triangular subulate, shorter than the hypan-
thium or much longer, to 7 mm long, pubescent; pedicels short or absent. Flowers
with the hypanthium subrotund, ca. 0.8 mm long, puberulent, the calyx ca. 1 mm
long, the teeth 4, triangular, ca. 0.4 mm long, the cup stiffly petaloid, glabrous
and eglandular within; corolla white, the tube narrowly cylindrical, 10.0-13.5 mm
long (or longer), ca. 1 mm wide, petaloid, sparsely hairy outside, sometimes
glabrescent, glabrous within, the lobes 4, oblong rotund, 3.2 mm long; stamens
4, the anthers narrowly oblong, 2.0-2.2 mm long, the filaments short, ca. 0.7 mm
long, attached 3-4 mm below the mouth; style slender, 7-10 mm long, the stigmas

194

ANNALS OF THE MISSOURI BOTANICAL GARDHN

[Vol. 67

FiGURfc 44. Gonzxihii^unia pananwnsis (Cav.) Schum. — A. Habit (x'/z). — B. Flower, opened to
show stamens (x5'/2). — C. Ovary, cross section (x5'/2). [After Tyson 1422. \

1980] DWYER— FLORA OF PANAMA (Family 179. Riihiaceac) 195

4, ca. 0.7 mm long. Fruits 4-lobed, rotund, 3-4 mm in diam., glabrous to pubes-
cent, white or purple black at maturity.

Gonzcilogunia panamensis ranges from Mexico to Colombia, and it also occurs

in the West Indies. A common name in Panama is ''Niguita." Stier 200 reports

the common name "Ina Kapuriwiwa'' by the San Bias Cuna Indians. Steyermark

(1972) pointed out that Gonzxilagiinia comifoUa (H.B.K.) Standley (1929) is the

''Gonzcilagunia panamensis of Standley and other authors, not G. panamensis
(Cav.) Schum,'' but he was referring to Standley's use of his name for plants

of Venezuela, not his use of the name for plants of Panama.

Literature:

Steyermark, J. A. 1972. The Botany of the Guayana Highlands — Part IX. Mem.
New York Bot. Card. 23: 312.

BOCAS DHL TORo: Changuinola Valley, Dufilap 496 (BM, US), canal zone:: Ancon, Heriherto
(US). Cerro Luisa, Road C-2, Croat 10773 (MO). Ft. Sherman, Duke & MusscU 6639 (MO). Quarry
Heights, Dnyer2608 (MO). Fort Sherman, Lhvyer 7183 (MO). Coco Solo, Dnycr <^ Duke 7883 (GH,
MO, UC, US). K-9 Military Road, Ehini^wr 543 (MO). Balboa Heights, Grcenman & Greemmui 5032
(MO). Ancon, Killip 12052 (US). Gamboa, PittUr 3711, 3785 (both US). Ancon, Rimlee & Ronlec
396 (US). Ancon Hill, Scihcrt 404 (MO). Balboa, Standley 26977 (US). Corozal, Standley 27358
(US). Fort Sherman, mouth of Chagres, Tyson 1536 (MO). Fort San Lorenzo, Tyson 2214 (MO).
Miraflores, White <t White 57 (MO), chiriqui': El Hato (Volcan), Ehini^er 816 (MO). Frances Arriba
School, 14 mi N of David, Lewis et al. 675 (GH, K, MO, NY), cocle: El Valle, Bhim et aL 2381
(MO); Duke 13271 (MO). Penonomc, Dnver 2010 (MO). El Valle, D\v\er238I (MO). Near El Cope,
Garner 38 (MO). El Valle, Seihert428 (MO), colon: Portobelo, Croat 1 1412 (MO); Elias ct Kirkhride
1642 (MO, UC, VEN); Folsom 3753 (MO); Gentry 1738 (MO). N side Rio Guanche, V2 km upstream
from Puerto Pilon-Portobelo Highway, Nee 7088 (MO), hhrrhra: 10 mi S of Ocii, Tyson 2844 (MO).
Panama: Campana, Allen & Alston 1865 (MO). Pacora and Chepo, Dod^e et al. 16667 (BM, MO).
San Jose Island, Erianson 544 (US). Chagres, Fendler 288 (MO). San Jose Island, Johnston 76 (MO).
Las Cumbres, Rivera 18 (MO). Chepo, Tyson 1422 (MO). Veracruz, Wendehake 9 (MO), san bi as:
Between Ri'o Diablo and Rfo Acuatf, near Nargana, Duke 14859 (MO). Mainland opposite Achituppu,
Lewis et al. 100 (GH, MO). Niatuppu, Stier 200 (MO), vhraguas: 5 mi NE of La Mesa, Blum &
Tvson 668 (MO). Santiago, Dwxer 1253 (MO). Ponuga, Dwyer & Kirkhride 7441 (GH, MO, US).
Santiago, Dwyer et al 7557 (COL, MO, VEN, UC).

5. Gonzalagunia rosea Standley, Ann. Missouri Bot. Gard. 25: 836. 1938. type:
Panama, White & White 7 (F, holotype; isotypes, MO, NY).

Trees, small, to 5 m tall, or shrubs, the branchlets terete, stiff or flexuous,
densely pubescent. Leaves oblong, ovate oblong or elliptic, 7-22 cm long, 2-8
cm wide, deltoid at the apex, often with an acumen to 1.5 cm long, basally cuneate
or rounded, the costa prominulous above and beneath, the lateral veins 8-13,
prominulous, arcuate, papyraceous, the hairs moderately dense, strigose or
spreading, often farinose above (bases of deciduous hairs), usually densely golden
villose beneath; petioles 0.5-2.0 cm long, 0.1-0.2 cm wide, pubescent; stipules
oblong triangular or triangular, 8-10 mm long, acuminate, the acumen often 4
times the length of the body, keeled, golden pubescent. Inflorescences terminal,
spiciform, 15-35 cm long, often nodding, to 5 cm wide, pubescent; peduncles to
4 cm long, to 0.18 cm wide, the cymules well spaced; bracts small. Flowers with
the hypanthium obovoid, ca. 3 mm long, ca. 1.4 mm wide, villosulose, the ca-

lycine cup ca. 1 mm long, petaloid, the teeth 4-5, triangular, ca. 0.6 mm long,
with 2-4 subrotund glands at the junction of the lobes; corolla pink, the tube
slender, ca. 8 mm long, ca. 1.6 mm wide in the middle, densely pubescent outside,
densely appressed and white villose within, carnose, the lobes 5, 2 mm long.

196 ANNALS OFTHF MISSOURI BOTANICAL GARDKN |VoI. 67

thick, the margins infolded, densely villose adaxially; stamens 4, the anthers
oblong, ca. 2.5 mm long, ca. 0.25 mm wide, the filaments short, inserted -A way
up the tube; stigmas 4, unequal, subfalcately digitiform, 0.6-1.0 mm long, the
style filamentous, ca. 9 mm long, the ovary 4-celled, thick-walled, the ovules
numerous, rubescent. Fruits 4-coccous, 2-3 mm in diam., glabrate.

Gonzxilagunia rosea is known from Costa Rica and Panama. Mori & Dressier
7875 from Cerro Colorado, Chiriqui, is noteworthy in having glabrous leaves.

BOCAS DLL TORo: Between Quebrada Gutierrez and E slope I. a Zorra, Kirkhride S: Duke 737
(MO), chiriqui': Cerro Horqiieta. Blum & D\\yer26J8 (MO). Banibilo and Cerro Punta, Croai 10676
(MO). Top of peak between Barii and Respinga, ca. 3000 m, D'Any 10125 (MO). La Popa, above
Boqiiete. D'Arcy 10859, 10946 (both MO). Bajo Grande, D'Arcy & D'Arcy 6561 (MO). Bajo Chorro,
Davidson 152 (MO, US), 259 (MO). Cerro Horqueta, Duke et aL 13615 (MO); Dwycr et al. 490 (MO,
SCZ, UC, VEN). Boquete, Dwycr 6992. 6995 (both MO). Cerro Horqueta, Dwycr & Harden 7702
(MO): Dwyer & Lallathin 8774 (MO). El Volcan on road to Costa Rica, Ehini^er 816 (MO. US).
Quebrada Hondo and divide on Caldera-Chiriquicito Trail, Kirkhride & Duke 927 (MO). 27 km NW
of El Hato del Volcan, 5000-5300 ft, Mori <& Bolten 7199 (MO). Cerro Colorado, 50 km N of San
Felix, 1200-1500 m, Mori c<- Dressier 7875 (MO). Cerro Punta, Rid^way & Solis 2409 (MO, UC).
Boquete, Stern et al. 1028 (MO. US). 1 m SW of Cerro Punta, Tyson 5646 (MO). Bambito, Tyson
5861 (MO). Rio Chiriqui Viejo Valley, White ct White 7 (F, MO), 80 (MO). Quebrada Velo, Woodson
& Schery 262 (MO), darien: Upper Rio Membrillo and Camp 7, Duke 10908 (MO). HtRRHRA: 10 mi
S of Ocu, Tyson et al. 2844 (MO), vlraguas: NW of Santa Fe. 1 1 km from Escuela Agricola Alto
de Piedra, Valley Ri'o Dos Bocas, Mori et aL 3841 (MO).

6. Gonzalagunia rudis (Standley) Standley, J. Wash. Acad. Sci. 17: 170. 1927.

Du^iyenia rudis Standley, Contr. U.S. Natl. Herb. 18: 125. 1916. type: Panama, Pittier 2378 (US).
Gonzahii^unia aeutifolia Rusby, Descr. S. Amer. PI. 131. 1920.

Small trees, to 3 m or shrubs, the branchlets terete, slender, 0.1-0.5 cm wide,
puberulent, the nodes well spaced. Leaves ovate oblong, trapeziform oblong,
often falcate and inequilateral, 3-11 cm long, 1-4 cm wide, deltoid to acute at
the apex, acuminate, the acumen vague or to 1.5 cm long, acute, rounded, obtuse
or occasionally truncate at the base, the costa prominulous above and beneath,
the lateral veins 8-10, strongly arcuate, papyraceous, glabrescent or pubescent
above, the hairs white, strigose, villose beneath, especially on the veins; petioles
to 0.3 cm long, the blades sometimes sessile; stipules ovate or deltoid, ca. 6 mm
long, the awn subulate, as long as the body, the body sometimes much reduced
and the awn then to 5 times longer. Inflorescences terminal, occasionally ap-
pearing axillary as on reduced side branches, solitary, exceeding the terminal
leaves, to 20 cm long, flagelliform, the peduncles 0.5-3,0 cm long, slender, to 0.1
cm wide, the flowers occasionally solitary or 3-4 in sessile or subsessile cymules,
these 0.5-1.0 cm apart: bracteoles linear subulate, to 5 mm long. Flowers with
the hypanthium rotund, ca. 1 mm long, truncate, villose, the calycine tube ca.
0.3 mm long, the teeth 4, unequal, triangular, ca. 0.7 mm long, obtuse to acute,
with a minute gland in the sinuses; corolla white, the tube cylindrical, ca. 3 mm
long, expanded at the apex, white villose on the outside and within, the lobes 4,
oblong rotund, ca. 1.5 mm long, with papillae and hairs on upper surface; stamens
4, the anthers narrowly oblong, 1.0-1.3 mm long, the filaments slender, ca. 0.5
mm long, inserted near base of tube; style linear, 2.5-3.0 mm long, the stigmas
3-4, ca. 0.3 mm long. Fruits white, coccoid, rotund, to 0.5 cm in diam., fleshy,
pubescent, the calycine lobes persistent, often radiating in starlike fashion.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaccac) 197

Gonzalagunia rudis has a limited range, being known only from Panama and
Colombia. In Panama it is known as '^Niguita/' Among the Bayano Cuna of
Panama it is called ^^Usiburnu/^ ^Turuiwat^' and ^4nna Ulu'^ {Duke. 14460),
Duke 14548 notes that it is used by the Choco Indians for snakebite. Stier 177
lists the common names ''Ipar kae Tummat'' (San Bias) and ^'Ipa Kae'' (San
Bias) and notes that it is used in '^Ibia ina'' (apparently as a medicinal).

CHiRiQuf: 21 km WNW of Puerto Armuelles, Buscy 459 (MO). 4 mi S of Puerto Armuelles,
Liesncr 379 (MO), canal zone: Mohinga Swamp, Bartlctr & Lasser 16856 (MO). Gamboa, Clewcll
& Tyson 3217 (MO). Pipeline Road, Croat 9383, 12341 (both MO). Pipeline Road, within 5 mi of
Gamboa Gate, DAny 9285 (MO). Rio Indio, Dodge & Allen 17298 (MO). Gamboa. Ehinger 904
(MO). Pipeline Road, G(v;/rv & Dressier 1967 (MO). 25 km NE of Cerro Azul on Rio Piedras, Mori
& KallunU 3367 (MO). Chagres at El Vigia, Pittier 2378 (US). Gatiin, Standley 27180 (US). Frijoles,
SumJley 27560 (US). Gamboa, Tyson 1521 (MO), colon: Tumba Vieja, Dodge et cd. 16935 (MO).
darien: Rio Tuquesa below Quebrada Venado, Bristan 1033 (MO), Rio Pirre, Bristan 1314 (MO).
Rio Uruceca, Bristan 1441 (MO), Rio Chico, Dnke & Bristan 450 (MO). Ri'o Tuira and Rio Paca,
Duke 5029 (MO). Rio Pirre, 10 mi S of El Real, Dnke 5450 (MO). Rfo Balsa, Duke 8715 (MO). Rio
Sambu, Duke 9306 (MO). Cerro Pirre, Duke & Elias 13813 (MO, UC, VEN). Rio Morti, Duke 14134
(MO). Trail near Piria, Duke 14328 (MO). Canasas and Sabalo, Duke 14460 (MO). Rio Tuira, Duke
14548 (MO). Puerto Sta. Dorothea, Dwyer 2275 (MO). Rio Cuasi, Kirkhride & Bristan 1439 (MO).
Yaviza. Stern et al. 147 (MO). Rio Paya, Stern et al. 365 (MO, US). LOS Santos: Rio Pedregal, 25
mi SW of Tonosi, Lenis et al. 2922 (COL, MO, UC, VEN). Panama: Rio La Maestra, Allen 22
(MO). Near Jenene, Duke 3851, 3889 (both MO). El Llano to Ri'o Mamoni, Duke 5596 (MO). Piria-
Caiiasas trail near Piria, Duke 14328 (MO). Fort Lorenzo, Ehinger 458 (MO). Rio Chagres, 5 mi SW
of Cerro Brewster, Lewis et al. 3443 (MO). 25 km NE of Cerro Azul on Ri'o Piedras, 550 m, Mori
& Kallunki 3367 (MO). Rio Chagres at El Vigia, Pittier 2378 (MO), san blas: Puerto Obaldia, Croat
16713 (MO). Rio Cuadi, Duke et al 3658 (MO, UC, VEN). Cangandi-Mandinga Airport Road, Duke
14789 (MO). Rio Diablo and Rio Acuati. Duke 14859 (MO). Isia Pino, near Mulatupo, Elias 1702
(MO, UC). Playon Chico, Mainland, Gentry 6413 (MO). Mainland opposite Achituppu, Lewis et al.
132 (GH, K, MO, NY, UC, US). Alligandi, Lewis et al. 143 (GH, K, MO, UC, US). Trail to inland
village of Armila, 3-8 km SW of Puerto Obaldia, Mori et al. 6800 (MO). Playon Chico, Stier 157, 177
(both MO).

7. Gonzalagunia veraguensis Dwyer. ''' type: Panama, Mori 6673 (MO, holotype).

Shrubs to 2 m tall, the twigs terete, golden velutinous, the nodes well spaced.
Leaves elliptic or elliptic falcate, 5-25 cm long, 3-9 cm wide, deltoid or acute at
the apex, acuminate, the acumen triangular, ultimately acute, cuneate to acute
at the base, the costa prominulous above, prominent beneath, the lateral veins
ca. 14, arcuate, not forming a distinct marginal vein, prominulous, usually with
1 short, soon branching or evanescent smaller vein between a pair of lateral veins,
the numerous smaller veins arising from the veins often in a subperpendicular
fashion, the intervenal areas spreading reticulate, rigidly papyraceous or subcori-
aceous, discolorous, somewhat lustrous above, with a few scattered hairs on the
costa and the veins above or in young leaves scattered over the upper surface,
densely shaggy villose beneath, the hairs slender, appressed, to 2 mm long, more

^^ Gonzalagunia veraguensis Dwyer, spec. nov. Frutices ad 2 m alti ramulis angulatis ciliis
appresso-ferrugineis. Folia lanceolala, 15-28 cm longa. 4.5-8.5 cm lata, venis lateralibus 10-16 pa-
pyracea supra glabra praeter in costa venisque subtus dense pubescentia ciliis ad 2.5 mm longis.
Infloreseentiae terminales, ad 50 cm longae. dense pubescentes, pedunculo ad 6 cm longo, cymis
sessilibus, ad 4 cm latis, 1.0-2.5 cm distantibus; bracteis ad 0.5 cm longis. Flores sessiles; calyce
lobis subulatis, 1.5-2.0 mm longis; corolla tubo cylindrico, ad 7.5 mm longo, extus ciliis crebris
appressis, lobis 4, ca. 3 mm longis; anlheris ca. 1.8 mm longis; stigmatibus 4 inaequalibus, ad 0.5 mm
longis. Fruetus non visi.

198 ANNALS OF THE MISSOURI BOTANICAL GARDHN [Vol. 67

abundant on the costa and the veins; petioles 0.3-3.2 cm long, lignose, stout,
pubescent; stipules often persistent, ovate or triangular, to 1.5 cm long, to 0.6
cm wide, acute, glabrous outside except for a densely pubescent median keel

which arises from a densely pubescent broad basal band ca. 1.5 mm deep. Inflo-
rescences terminal, solitary, spikelike, 15-60 cm long, to 2 cm wide, the flowers
in well spaced aggregates; bracts linear, usually as long as the pedicels. Flowers
sessile or subsessile; hypanthium rotund, ca. 2 mm long, sericeous, the calycine
cup to 0.7 mm long, glabrous, often with a few irregular glands in the sinuses,
the latter triangular or subulate, to 1.8 mm long, acute; corolla white, the tube
narrowly cylindrical, to 7 mm long, thickly petaloid, the hairs outside dense,
white, appressed, puberulent to sparsely ciliate within, the lobes 4, oblong, ca.
3 mm long, obtuse; stamens 4, the anthers ca. 1.8 mm long, obtuse, the filaments
slender, almost equaling the anthers, inserted 1.5-3.0 mm above the base of the
tube; ovarian disc annular, ca. 0.2 mm deep, the style ca. 3 mm long, the stigmas
4, unequal, digitiform, to 0.5 mm long. Fruits not seen.

Gonzxihigunia veraguensis is known only from Panama. It is readily recog-
nized by its many veined shaggy villose (beneath) leaf blades and its very short
corolla.

VERAGUAs: Valley of Ri'o Dos Bocas between Alto Piedra above Santa Fe and Calovebora, 350-
400 m. Croat 27462, 27733 (both MO). Near Escuela Agricola Alto de Piedra, Mori 6673 (MO).

43. GUETTARDA
Guettarda L., Sp. PI. 991. 1753; Gen. PI., ed. 5: 428. 1754. type: G. speciosa L.

Matthiohi L., Sp. PI. 1192. 1753, not R. Brown, (1812. Nomen conserv., Cruciferae). type: A/.

srahra L.
Halesia P. Br., Civ, Nat. Hist. Jam. 205. 1756, nomen rejic. not J. Ellis ex L. 1759 (Nomen conserv.,

Styracaceae).
Laui^eria Jacq., Enum. PI. Carib. 2. 1760. type: L. odorata Jacq.

Trees or shrubs. Leaves occasionally ternate, usually petiolate; stipules de-
ciduous. Inflorescences axillary, cymose, the flowers occasionally solitary, usu-
ally secundly disposed or congested; bracteoles present or absent. Flowers with
the hypanthium ovoid or globose, the calycine cup tubular, cupular or campan-
ulate, truncate, edentate or 2-9-toothed; corolla funnelform or salverform, the
tube elongate, the lobes 4-9, imbricate, 2 of them exterior; stamens 4-9, inserted
in the tube, the anthers slender, dorsifixed, included, the filaments frequently
wanting; style slender, the stigmas capitate or 2-lobed, the ovary 2-9-celled, the
ovules 1 per cell, pendulous. Fruits drupaceous, globose, ovoid or oblong, thinly
carnose, usually with a fine velvety indumentum, often acutely angular, the pu-
tamen ligneous or bony, the testa membranous.

Guettarda, a genus of about 80 species, occurs throughout tropical America.
It also occurs in New Caledonia (fide Willis 1966).

a. Inflorescences with cymes 3-flo\vered; peduncles stout 5. G. runiuliftora

aa. Inflorescences with cymes more than 3-flowered; peduncles slender.
b. Leaves with lateral veins 6-10.

c. Fruits obovate oblong, less than 2.2 cm long 6. G. sanhhisensis

cc. Fruits rotund, 7.0-1.5 cm in diani.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaccae) 199

d. Leaves 10-16 cm long; petioles to 2 cm long 2. G.foliacca

dd. Leaves 2.0-5.5 cm long; petioles to 0.7 cm long 4. G. odorata

bb. Leaves with lateral veins 10-12.

e. Leaves glabrate above; fruits compressed rotund, ca. 2 cm long, not sharply

angled 3. G. macrosperma

ee. Leaves villose or viljosulose above: fruits oblong, to 0.8 cm long, sharply angled

L C chiriqiiicnsis

1. Guettarda chiriquiensis Standley, Ann. Missouri Bot. Gard. 25: 838. 1938.
type: Panama, White & White 22 (F, holotype; MO, isotype).— Fig. 45.

Trees to 30 ft tall, the branches terete or angular, the twiglets angular, stout,
densely puberulent, the nodes usually well spaced. Leaves ovate rotund to lan-
ceolate, 6.5-18.0 cm long, 2.5-12.5 cm wide, acute or somewhat rounded apically,
frequently cuspidate, cuneate to occasionally obtuse or rounded at the base,
rarely truncate, the costa prominulous above, prominent beneath, the lateral veins
10-11, prominulous, especially beneath, those in the lower V^ of the blade strict
but marginally arcuate, those of upper VS markedly arcuate, stiffly papyraceous
to subcoriaceous, usually discolorous, villose or villosulose or both; petioles to
4.5(-10.0) cm long, usually rigid, densely pubescent; stipules often persistent at
the ends of the branchlets, ovate or oblong, 12-13 mm long. Inflorescences ax-
illary, 1 per axil, about as long as the petiole of the adjacent leaf, occasionally
longer, the peduncles 1-2 cm long, densely golden pubescent, bifurcating, the
branches 2.5-4.0 cm long, the flowers unilaterally disposed on the branch, usually
8-12 per branch. Flowers sessile; hypanthium oblong, ca. 2.5 mm long, the ca-
lycine cup ca, 1 mm long, the 4 teeth deltoid, ca. 0.5 mm long, densely pubescent
outside, corolla white or purplish white, the tube narrowly cylindrical, ca. 12 mm
long, ca. 1.5 mm wide in the middle, densely golden hirsute outside, the hairs
deflexed, glabrous within, carnose, the lobes 4, oblong, the margin conspicuously
lobed and scalloped, the whole ca. 6 mm long, and 5 mm wide at the base,
glabrous within, puberulent outside; stamens 4, the anthers oblong, ca. 2.8 mm
long, ca. 0.7 mm wide, obtuse, dorsifixed, the filaments plane, ca. 0.35 mm wide,
attached just below the orifice; stigmas 2, oblong, 1 mm long, subconcave adax-
ially, the style 15-18 mm long. Fruits sessile, oblong, to 0.8 cm long, ca. 0.5 cm
wide, the remnants of calycine cup persistent as an apiculum, sharply 4-angled
and 4-sulcate, drying grey-green, the pubescence dense.

Guettarda chiriquiensis is known from Panama and Colombia but may be
expected in adjacent Costa Rica. The solitary Colombian collection cited below
occurs at a much lower elevation than the Panama collections.

CHiRiQUi: Between Cerro Punta and Las Nubes, 1800-2100 m, Croat 26394 (MO). Valley of Ri'o
Chiriqui Viejo, N of Volcan City, 5200-5600 ft, Duke 9037 (MO). El Hato, 5000-6000 ft, Ehin^er 822
(MO). Boquete, 1400 m. Maasola 28 (MO). 27 km NW of El Hato del Volcan, 5000-5300 ft, Mori
& Boltcn 7189 (MO). Cerro Punta, 6000 ft, Tyson 5820 (MO). Above Jaramillo Arriba, N slopes of
Cerro Palo Alto, 4700 ft, Webster 16684 (MO). Volcan de Chiriqui, 1700 m, Woodson S^ Schery 508
(MO), cocle: El Valle, Correa3l6 (MO). Cerro Pilon, Dnke 121 1 1 (MO); Dnyer 10566 (MO): Dnyer
^ Correa 7946. 7975 (both MO); Dwyer & Lallathin 8618 (MO). La Mesa, Dwyer & Nee 11936 (MO);
Gentry 5637, 6837 (both MO). El Potroso, Folsom & Collins 1554 (MO). Cerro Pilon, Mori 6555
(MO)." La Mesa, Nee & Dwyer 9180 (MO). Cerro Pilon, Porter et al. 4416 (COL, DUKE, MICH,
MO, NY, SCZ, UC, VEN). Panama: Cerro Campana, Correa & Dressier 368 (MO); Croat 12158
(MO); Dwyer S: Kirkbride 7835 (MO); Gentry 4917 (MO); McDaniel 6857 (MO); M<n-i & Kallunki
3572 (MO). COLOMBIA: cuoco: Rio San Juan just above Istmina, 100 m, Gentry & Fallen 17650 (MO).

200

ANNALS Of THE MISSOL Rl BOTANICAL GAKDLN

[VoL 67

Figure 45. Gucttarda cliiricjniensis Standley. — A. Flowering branch (x'/2). — B. Flower
(xL5).— C. Ovary sections (x3). [After Dirv^'r & Nee 11936.].— D. Fruiting branch (x'/2). [After
Gentry 5637. ]

2. Guettarda foliacea Standley, Contr. U.S. Natl. Herb. 18: 139. 1916. type:
Panama, Pittier4031 (US, holotype; isotype, F).

Trees, to 15 m tall, or shrubs, the twiglets terete, smooth, glabrous, ultimately
pubescent, often with spreading lateral branches. Leaves often crowded at the
apex of the twiglets, obovate oblong to elliptic, 3-17 cm long, 1.2-6.5 cm wide,

19801 DWYER—FI.ORA or- PANAMA (Family \19. Riihiavcac) 201

widely deltoid or rounded at the apex, the acumen short or absent, less than 1
cm long, basally obtuse or truncate, rarely acute or subcordate, usually inequi-
lateral, the costaprominulous beneath, the lateral veins 8-10, arcuate, the veinlets
pinnatiform, stiffly membranous or chartaceous, concolorous or discolorous, the
hairs above white or scattered, the hairs beneath short or long, appressed or
spreading, often strigose, appearing shaggy, the axils often barbate; petioles 0.8-
2.0 cm long, slender, puberulent; stipules not seen. Inflorescences axillary, usu-
ally spreading, the peduncle slender, to 3 cm long, ca. 0.5 mm wide, pubescent,
the few branches short and often secundly disposed; bracts deciduous, oblong,
to 6 mm long, ca. 1 mm wide, sericeous, the calycine cup cupuliform, scarious,
usually slightly longer than the hypanthium, sericeous, truncate, the teeth absent;
corolla white, the tube narrowly cylindrical, to 1.5 cm long, ca. 1 mm wide in the
middle, sericeous outside, glabrous within, the lobes 4-5, oblong, ca. 3 mm long,
obtuse, thinly carnose, sericeous outside, the anthers subsessile, narrowly ob-
long, 3 mm long, attached ca. 2 mm below the mouth; style slender, the stigmas
compressed, capitate rotund, ca. 0.25 mm long. Fruits sessile, rotund, 0.8-
1.5(-2.0) cm in diam., finely velutinous, red at maturity.

Gnettardafoliacea occurs in Panama and Colombia. "Guyababa''; "Espino."

BARRO COLORADO ISLAND! Bailey & Bciilcy 202. 611 (both F); Croat 6029, 6061, 6114. 6240. 6540.
7405, 7422. 7670 (all MO), 1145 (F). \\764 (MO). 15094 (F), \5\60 (MO): Dwycr 1416 (MO); Ehiniii.-r
267 (MO), 390 (MO. US); Foster 951. 1002 (both F); Knif^'ht 69-1, 69-40 (both MO); Shattuck 8 (F);
Starrv 222 (F); Zetek 3580 (MO), canal zonf: Road S-11, NW of Escobal, Croat 12471 (MO).
Pipeline Road, Croat 15094 (MO). Coco Solo, Gentry 6191 (MO). Barbacoas Railroad Station, Hayes
167 (F). Pipeline Road, Mori & Kallunki 2051 (MO). Trinidad River, Pittier403l (F, US). cHiRiQUi:
Progreso, Cooper & Slater 227 (US), 236 (F, US), darien: Rio Ucurganti, Bristaii 1175, 1178 (both
MO). S of El Real, Duke 5036 (MO). Rio Chucunaque between Rio Membrillo and Ri'o Subcuti, Duke
8602 (MO). Rio Sabana, above Santa Fe, Duke 14085 (MO). Piria-Canasas trail near Piria, 100 m,
Dake 14310 (MO). Between Canasas and Sabalo, Duke 14478 (MO). El Real Qiiebrada, Trapiche,
Duke & Bristaii 322 (MO). Campamento Buena Vista, Stern et al. 957 (US). 2 mi E of Santa Fe,
Tyson et al. 4845 (MO). Panama: Rio Chepo, Rio Bayano, Duke 3987 (MO). San Jose Island between
Bodega Bay and Rio Mala Puerco, Duke 12552 (MO). Llanos of Rio Mar, Dwycr 1799 (MO). Rio
Pasiga, Gentry 2287 (MO). Between Cariita and dam site, Gentry & Tyson 1704 (MO). El Llano, road
to Jesus Maria, Tyson 6873 (MO), vf.raguas: Rio Trinidad, below Rio de Jesiis, Dwyer 1321 (MO).
Santiago, Stern et al. 9% (MO, US). Colombia: choco: Behind beach. El Valle, sea level. Gentry
& Fallen 17231 (MO).

3. Guettarda macrosperma Donnell Smith, Bot. Gaz. 18: 204. 1893. type: Gua-
temala, Heyde c5 Lux 3160, not seen.

Trees to 6 m tall, much branched, the twiglets terete, glabrous to pubescent,
the nodes moderately well spaced, the lenticels minute, often inconspicuous.
Leaves often crowded toward the apex of the twiglets, elliptic oblong, oblong
rotund or occasionally pyriform, 12-20 cm long, 6-13 cm wide, deltoid to obtuse
at the apex, acuminate, the acumen short, the apex of the blade often eroded,
basally obtuse to widely cuneate, occasionally acute at the base, the costa prom-
inent beneath, the lateral veins ca. 12, prominent, rigid, often light in color and
contrasting with the rest of the blade, at first angular ascending then arcuate
toward the margin, the ultimate veinlets vaguely lineolate, stiffly chartaceous,
often brittle when dry, concolorous, glabrous or pubescent on the costa and the
veins above, beneath with the hairs short, white, strigose, the vein axils often
shortly barbate; petioles to 4 cm long, 0.10-0.25 cm wide, stiff, often curved,

202 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol.67

glabrous to pubescent; stipules deciduous, not seen. Inflorescences axillary, the
peduncles to 1.5 cm long, to 1.5 mm wide, stiff, the pubescence minute, green,
velutinous. Flowers not seen. Fruits sessile, compressed rotund, ca. 2 cm wide,
the pubescence minute, velutinous, yellow green or green when dry, delicately
ribbed, the ribs disposed as faint and scarcely elevated lines, fruit wall thin; seeds
5-7, in a hard, white, pinkish or molasses-colored pulp.

Guettarda macrosperma extends from Guatemala to Panama. The Panama-
nian collections, while relatively abundant, lack flowers, making critical identi-
fication difficult. Donnell Smith in his original diagnosis describes the mature
leaves of G. macrosperma as "2^)4 x VA in." with "6-7 costal nerves." The
fruits are described as "obovoid oblong (6-7 x 3-4 inches)." The leaves of Pan-
amanian material are considerably larger, the lateral nerves are about 12, about
double the number listed for the type which was collected at about 3,000 ft at
Santa Rosa, Guatemala. The fruits of the Panamanian material do not exceed an
inch in diameter while Donnell Smith describes the fruits as 4 inches in diameter.
Unfortunately, I did not see the type to verify this incredibly large fruit. Paul
Standley identified Wedel 266 as G. macrosperma. It and CarrasqidUa et al. 387
have much smaller leaves even though the fruit is like that of the exsiccatae
below. About V2 of the Panamanian material was collected near sea level in the
limestone outcrops at Madden Dam.

CANAL zone: Madden Lake, Croat 38326 (MO). Fort Sherman, Duke & MusseU 6638 (MO).
Madden Dam, Boy Scout Road, Dwycr & Elias 7504 (GH, MO); D\s\er & Havden 20 (MO): Ebinger
(MO). Area W of Limon Bay, Gatiin Locks, Johnston 1704 (MO). Madden Dam, Boy Scout Road,
KirkhriJc 42 (MO): Kirkhridc & EUas 260 (MO): Lewis el al. 15 (GH, K, MO, NY, UC, US). Pipeline
Road, 4 km NW of Gamboa, Nee 7225 (MO). Sotto Caballo, Smith et al. 3318 (F, MO), colon: Rio
Boqueron, 3 km from La Pelona, CarrasqidUa et al. 387 (MO). Between Garotte and La Guaira,
DArcy 9349 (MO). Portobelo, Dwyer 5149 (MO). Maria Chiquita, Dwver & Kirkhride 7772 (Gh!
MO. UC, US), darifn: Santa Fe, Duke 14085 (F). san blas: Ustupo, D'Arex 9534 (MO). Ailigandf
River. Duke & Bristan 349 (MO). Rio Guadi. Camp Diablo, drill site 22, Duke et al. 3646 (MO) Isla
Pino near Muiatupo, 200 ft, Elias 1705 (MO, UC).

4. Guettarda odorata (Jacq.) Lam., Tab. Encycl. Meth. 2: 219. 1819.

Laugeria odorata Jacq., Enum. PI. Carib. 16. 1760. rvpi : Cuba, ,^ky////>i. not seen.

Shrnhs or trees, to 3.5 m tall, the branchlets terete, distichous, well spaced,
the bark drying dull grey, often mottled, occasionally conspicuously lenticellate.
Leaves oblong, 2.0-5.5 cm long, 1.0-2.5 cm wide, cuneate to rounded at the
apex, acuminate, the acumen vague, basally cuneate, obtuse or vaguely rounded,
the costa plane, immersed or prominulous above, prominent beneath, the lateral
veins (6-)8-9, arcuate, the veinlets lineolate, irregularly perpendicular to lateral
veins, the intervenal areas often mosaic-like, papyraceous, glabrate above, villose
or merely puberulent on the costa and the veins as well as the intervenal areas,
below the hairs of the veinlets occasionally shaggy; petioles 0.2-0.7 cm long,
slender, puberulent or villose, often curved; stipules ovate deltoid, ca. 2 mm long,
acute, pubescent. Inflorescences axillary, 1-2 per axil, 3-5 cm long, the pedun-
cles 1.5-3.0 cm long, capillaceous or wiry, to 0.5 mm wide, pubescent, producing
1 cymule at the apex, this usually 5-flowered, or bifurcating and producing a pair
of 5-8-flowered cymules. Flowers sessile or subsessile; hypanthium subrotund.

1980] DWYFR— FLORA OF PANAMA (Family 179. RuhuHcae) 203

ca. 1 mm long, carnose, the calycine cup ca. 2.5 mm long, carnose, eglandular
within, the teeth absent; corolla white or reddish, the tube narrowly cylindrical,
to 16.5 mm long, ca. 1 mm wide, puberulent outside, glabrous within, the lobes
(4_)5(_6), oblong, 2.5-4.2 mm long, thinly carnose; stamens (4-)5, the anthers
linear oblong, 1.5-3.0 mm long, ca. 0.2 mm wide, dorsifixed, subsessile or sessile,
occasionally the filaments as long as the anthers, attached immediately below the
mouth or at the upper % of the tube, the ovarian disc annular, the style linear,
8-14 mm long with appressed ascending hairs, the stigmas 2(-3?), capitate, ca.
0.2 mm long, slightly wider than the style. Fruits rotund, to 0.7 cm in diam., the
persistent calycine cup to 1.3 mm long, the pericarp smooth, wrinkled when dry,
minutely grey green, puberulent, the hairs eventually deciduous^

Guettarda odorata occurs in Panama and in northern South America. It has

West

''Huesio Negro."

Steyermark (1972) discusses in detail his designation of a Jacquin collection from
Cuba as the lectotype.

CANAL zone: Fort Kobbe, Allen 1407 (F); Duke 3957, 4241 (both MO). Farfan Beach, D\yyer
& Hayden 7532 (GH, MO, US). Barro Colorado Island, Ehin^i^cr 267 (MO). Survival School, Curundu,
Twson 1059 (MO). Fort Clayton, Tyson 6779 (MO). MiraHores Lake (?), Wliiie 139 (MO). Ancon Hill,
Woodson et al. 1331 (F, MO). Fort Kobbe, Woodson ct (d. 1407 (MO), chiriqui': 6 mi S of Puerto
Armuelles, 0-150 m, Licsner 428 (MO), cocie: Penonome, Dwyer 2033 (MO), darien: Mouth of
Rio Yape, Allen 371 (F, MO). Rio Pirre, Duke 4962 (MO). Rio Cucunati at Puente Quemado, Duke
8806 (MO), herrhra: Ocu, Ehinger 1062 (MO). La Cabuya, Las Minas, Lao 139 (MO). Panama:
Pedro Gonzales, Allen 2597 (F). Calzada Larga, Croat 12385 (MO). Ri'o Mar, Savanna, Duke 12426
(MO). Isla Taboga, Hayes 637 (F). Las Sabanas, Pittier 6702 (F). San Jose Island, Sfimson 5326
(MO). Isla Taboga, Woodson et a!. 1482 (F, MO). E of Pacora, Woodson et al. 723 (F, MO), ver-
AGUAs: 5 mi N of Santiago, Rfo Santa Maria, Blum ct Tyson 616 (MO), 5 mi NE of La Mesa, Blum
& Tvson 667 (MO). 2 mi W of Santiago, Transisthmian Highway, Dwyer et al. 7544 (GH, MO).
Quebrada S of Santa Fe, 450 m, .V^^^^ 8040 (MO). San Jose, Santiago-Santa Fe Road, 400 m. Nee

8170 (MO).

5. Guettarda ramuliflora Beurl., Kongl. Vetensk. Acad. Handl. 132. 1854. type:
Panama, Billbcrg (S).

Trees, to 8 m tall or shrubs, the twigs terete, puberulent, ultimately finely
velutinous, the nodes well spaced. Leaves ovate oblong or sagittate, 6-9 cm long,
3-5 cm wide, acute to widely deltoid at the apex, acuminate, the acumen short,
somewhat vague, obtuse to widely deltoid at the apex, acuminate, the acumen
short, somewhat vague, obtuse to widely cuneate at the base, the costa promi-
nulous above, prominulous beneath, the lateral veins 6-8, arcuate, with short,
white, appressed, striate, densely appressed hairs on the costa and the veins
beneath, the intervenal areas lineolate but not conspicuously so, concolorous,
membranous; petioles slender, 1-3 cm long, ca. 0.1 cm wide, appressed villosu-
lose; stipules not seen. Inflorescences axillary, to 3.5 cm long, the peduncle stout,
to 1 cm long, velutinous, the flowers few, the cymes 3-flowered (fide Standley
1928) crowded at the end of the peduncle. Flowers sessile, villose; hypanthium
broader than long, 1 mm long, the calycine cup turgid, cupuliform, ca. 3 mm
long, truncate, edentate; corolla white, the tube narrowly funnelform, to 2.2 cm
long, to 0.2 cm wide, carnose, villose outside, glabrous within, the lobes 5, ob-
long, to 5 mm long, to 3.5 mm wide, obtuse, the margin with occasional shallow

204 ANNALS OF THE MISSOURI BOTANICAL GARDLN |Vol. 67

notches; stamens 5, sessile, the anthers narrowly oblong, obtuse, acutely sagittate

at base, attached ca. 3 mm below the mouth, the ovarian disc cylindrical, 1.3 mm

long, villosulose at the base, truncate, the style linear, ca. 7,5 mm long. Fruits
not seen.

Guettarda ramidiflora is known only from Panama.

PROViNcr: unknown: Billheri^ (S). canai zone: DeviLs Beach, woods back of beach, NW part of
Canal Zone, Johnston 1717 (MO).

6. Guettarda sanblasensis Dwyer.-^'- type: Panama, Duke ct ai 3646 (MO, holo-
type).

Trees 15-20 m tall, the branchlets terete, ultimately puberulent, the nodes
moderately well spaced. Leaves elliptic or elliptic rotund, 5-14 cm long, 4-10 cm
wide, toward the apex, rounded, deltoid, subacute or acuminate, the acumen
inconspicuous or to 1.5 cm long, ultimately acute, at the base cuneate or widely
cuneate, the costa plane above, prominulous beneath, the lateral veins 8-10,
arcuate, the venules pinnatiform, rigidly chartaceous, concolorous, green, shin-
ing, the hairs on both sides minutely strigose, those on the veins appressed, short,
velvety; petioles to 3 cm long, ca. 0.12 cm wide, stiff; stipules not seen. InJIo-
rescences axillary, the peduncle to 3 cm long, pubescent, the branches to 3 cm
long. Flowers not seen. Fruits obovate oblong, to 2.2 cm long, to 1.2 cm wide,
subtruncate at the apex, minutely velutinous, yellow green when dry, attenuate
toward the apex, the calycine cup ca. 2 mm long, ca. 1.5 mm wide.

Guettarda sanblasensis is known only from Panama. It is distinguished by its
apically subtruncate fruit.

SAN BLAs: Headwaters of Rfo Cuadi, Camp Diablo, drill site 22, N 82.2, E 87.8, 273 ft. f^uke et
(iL 3646 (MO, holotype).

44. HAMELIA

Hamelia Jacq., Enum. PI. Carib. 16. 1760. lectotype: //. erecta Jacq. = H.
patens Jacq.

HamcUia L., Sp. PI., ed. 2, 246, 1762, orth. mut. = Ihvnclia Jacq.

Tan^araca Adans., Fam. PI. II: 147. 1763. tvpel: not indicated.

Loniccm Adans., Fam. PI. II: 157. 1763. Not Lonicera L. (1753). typf.: not indicated.

DuhamcUu Pers., Syn. PI. 2: 203. 1805. typh: not indicated.

Tepesla Gaertn., Fruct. Sem. PI. 3: 72. 1805. iype: T, duhia Gaertner.

Shrubs or small trees. Leaves opposite or in whorls of 3-5, petiolate; stipules
free, entire or 3-lobed, small. Inflorescenees terminal, of thryses, dichasia or
cymes, the floral axes often spreading, the flowers usually secund; bracts decid-
uous. Flowers with the hypanthium 5-Iobed, the lobes often reduced, persistent

^- Guettarda sanblasensis Dwyer, spec. nov. Arhores 15-20 m altae. Folia elliptica vel elliptico-
rotunda, 5-14 cm longa, 4-10 cm lata, venis lateralibus 8-10 arcuatis, rigido-chartacca; petiolis ad 3
cm longis. Inflorescentiae in fructu axillares pedunculo ad 3 cm longo. Flares haud suppetunt. Fructus
obovato-oblongi, ad 2.2 cm longi, ad 1.2 cm lati, minute velutini in sicco luleo-virides cupula calycis
2 mm longa, 1.5 mm lata coronati.

1980] DWYI-R— FLORA OF PANAMA (Family 179. Ruhiuceae) 205

in fruit; corolla tubular or inflated, orange, red or yellow, the lobes 5, imbricate;
stamens 5, the connectives produced apically, the filaments attached at the base
of the corolla; ovary (4-)5-celled, the style slender, the stigmas 1-5. Fruits fleshy,
the ovarian disc often persistent and forming a truncated cone; seeds numerous,
piano compressed, foveolate.

Hamclia ranges from southern Florida through Mexico, Central America, the
West Indies and South America. There are about 16 species in the genus.

Literature:

Elias, T. 1976. A monograph of the genus Hamclia (Rubiaceae), Mem. New

York Bot. Card. 26(4): 81-144.
Wernham, H, F. 1911. A Revision of the Genus Hamclia. Jour, Bot. London

49: 206-216.

a. Corolla funnel shaped, yellow (sect. Amphituba).

b. Corolla tube 1.0-1.3 cm long; fruits 0.5-0.7 cm long 1. H. axillaris

bb. Corolla tube 1.8-5.0 cm long; fruits 7-16 mm long.

c. Calycine lobes 3-5 mm long; stipules subulate 2. H. calycosa

cc. Calycine lobes 0.2-2.5 mm long: stipules triangular to ovate.

d. Corolla tube 4-7 mm broad at the apex; inflorescence 10-25-flowered ..__

8. //. xerocarpa

dd. Corolla 10-20 mm broad at the apex: inflorescence 4-15-flowered

3. H. macrantha

aa. Corolla tubular, yellow, orange or red (Sect. Hamelia).

e. Lateral veins 3-5 pairs.

f. Inflorescence a much reduced dichasium, flowers usually 2-5; leaves 0.8-2.8

(-4.8) cm broad 7. H. san^uinca

ff. Inflorescence a spreading dichasium or cyme, flowers 3-8; leaves 2-7 cm broad

6. H. rovirosae

ee. Lateral veins 5-11 pairs.

g. Floral parts and leaves densely villous.

h. Leaves with lateral veins 3-5 pairs; seeds 1.0-1.2 mm long

6. H . rovirosae

hh. Leaves with lateral veins 7-10 pairs; seeds 0.6-0.9 mm long ..__ 5. H. patens

Floral parts and leaf undersides glabrous to puberulent.

i. Flowers 12-15 mm long; leaves with 11-16 pairs of lateral veins; corolla

yellow 4. //. magnifolia

ii. Flowers 15-25 mm long; leaves with 7-10 pairs of lateral veins: corolla

yellow to red 5. I!, patens

1. Hamelia axillaris->'* Swartz, Nov. Gen. Sp. PI. Prodr. 46. 1788, lectotype:
(Elias, 1976): Jamaica, Swartz (S).

Shrubs to 4 m tall, the branchlets terete to angular, glabrous to puberulent.
Leaves opposite or rarely ternate, elliptic, narrowly elliptic or obovate, 5-17 cm
long, 2-7 cm wide, acuminate at the apex, often conspicuously so, attenuate at
the base, glabrous above, often puberulent beneath and barbate in the vein axils,
the lateral veins 8-11, arcuate, chartaceous to membranous, glabrous or sparsely

l-4(

M

flowered compound dichasia, often disposed as scorpioid cymes; peduncles to

'^'^ For a list of synonyms see Elias (1976). Only this name has been used for Panamanian material.

206 ANNAl.S OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

1.5 cm long, the branches 1.5-4.5 cm long, spreading; bracts ovate, ca. 0.5 mm
long, puberulent. Flowers sessile or on pedicels to 1 mm long, sparsely puberu-
lent; calycine lobes to 1.5 mm long, puberulent, often densely so; corolla yellow,
the tube infundibular, expanding at the apex, to 11.5 mm long, the lobes ovate,
to 2 mm long, puberulent to glabrous; anthers 5-7 mm long, the apiculum ca. 0.5
mm long, the filaments to 5 mm long, the style slightly shorter than the tube, the
stigmas ca. 1.2 mm long. F/7//7.V globose or subglobose, 5-7 mm long, ultimately
black; seeds ca. 1 mm long.

Hamelia axillaris is a weedy shrub extending from Mexico through Central
America to northern South America. It also occurs in the West Indies.

CANAi. zone: Road S-IO, N of Escobal near junction of Road SI, Croat 12484 (MO). Ri'o
Mcndosa, Navy Reservation, 8 km N of Gamboa, Dressier 4087 (MO). Camp Piiia, Fort Sherman,
Dnkc 4398 (MO). Madden Dam, Boy Scout Road, Dwyer & Ellas 7491 (GH, MO, UC, US). N base
of Mindi Hills, Johnsion 1637 (MO), barro Colorado island: Croat (5299, W940, 11862. 11901 (all
MO); Haydcn 150 (MO); Starry 284 (MO), colon: Portobelo, Ehinger 118 (MO), darien: Rio Tu-
queza, below Quebrada Venado, Bristan 1034 (MO). Rio Aruza, Bristan 1367 (MO). Rio Balsa be-
tween Quebrada Chusomocatre and Rio Areti, Duke 8718 (MO). Quebrada Madkia off Rio Pucro
above Pucro, Duke 13013 (MO). Tumaganti, Duke 14160 (MO). Rio Tuira between Rio Punusa and
Rio Mangle, Duke 14635 (MO). Between Rio Perrecenega and Rancho Frio, Duke & Ellas 13860,
13867 (both GH, MO, US). Between Paya and Palo de Los Letras, Duke & Klrkhricle 14034 (MO).
Ri'o Tuquesa, Middle Tuquesa Mining Co. Camp called Charco Peje, Mori 7090 (MO). Between Rio
Escucha Ruido and Rio Tuira, Stern et al. 578 (MO). Cana, 1750 ft, Stern el al. 670 (MO). Campa-
mento Buena Vista, Rio Chucunaque, Stern et al. 828 (MO). 3 mi E of Santa Fe, Tyson et al. 4715
(MO). LOS Santos: Los Toretos, Dwyer 2424 (MO). Panama: Piria-Canasas Trail near Piria, ca. 100
m, Duke 14331 (MO). Cerro Azul, ca. 2000 ft, Dwyer 2072 (MO). Within 2 mi of Achiote, Eolsom
3871 (MO). Dam site near Rio Bayano beyond Chepo, Gentry ct Tyson 1676 (MO). Cerro Campana,
Mass et al. 1731 (MO); Mori & Kallunki 1937 (MO); Webster & Breckon 16510 (MO). Gorgas
Memorial Laboratory, ca. 25 km NE of Cerro Azul, Rio Piedras, 550 m, Mori c^- Kallunki 3459 (MO).

2. Hamelia calycosa J. D. Smith, Bot. Gaz. 12: 132. 1887. type: Guatemala,
Tiicrckhciiu 454 (US, holotype; NY, isotype).

//. chiapensis Brandegee, Univ. Cal. Publ. Bot. 6: 71. 1914. type: Mexico, Purnus 7030 (F, MO

NY).

H. hoyaeana Standley, Field Mus. Nat. Hist., Bot. Ser. series 11(5): 213. 1936. type: Colombia,
Lawranre 503 (NY, holotype, MO, US).

//. klu},'li Standley, Field Mus. Nat. Hist.. Bot. Ser. 13. No. 364: 96. 1932. type: Peru, Klu;^ 3664
(F, holotype).

Shrubs or (rccs to 10 m tall, the twiglets terete, smooth, glabrous, ultimately
puberulent, branching freely, the nodes well spaced. Leaves oblong, often 3-4-
nate at the ends of twigs, lanceolate or trapeziform lanceolate, 1.5-11.0 cm long,
0.9-3.5 cm wide, tapering acutely at the apex, acuminate, the acumen ca. 0.7 cm
long, acute to cuneate at the base, the costa grooved, plane or prominulous above,
the lateral veins 4-7(-9), arcuate, membranous or papyraceous, glabrous above
and beneath except occasionally pubescent on the costa beneath, often barbate
in the vein axils beneath; petioles slender, 0.8-2.0 cm long, wiry, glabrate; stip-
ules subulate, to 2.5 mm long, puberulent. Inflorescences terminal, rarely axil-
lary, the scorpioid cymes lax and spreading, the flowers many in 3-4 fascicles;
peduncles to 2 cm long, often minutely puberulent, the floral axes with up to 4
flowers, 0.6-4.0 cm long; bracts triangular to subulate, to 2 mm long, glabrous.
Flowers secund, the pedicels to 5 mm long; hypanthium oblong, puberulent, the

1980] DWYF.R— FLORA OF PANAMA (Family 179. Riihiaccae) 207

calycine cup scarcely measurable, the lobes oblong, to 1.5 mm long, fleshy, pu-
berulent; corolla pale orange or yellow, striped maroon, the tube somewhat in-
fundibuliform, 30 mm long, constricted narrowly at the base, the base ca. 1.5 mm
wide, ca. 7.5 mm wide in the middle of the tube, ca. 1 cm wide at the mouth,
puberulent outside, glabrous within, the lobes ovate, to 6 mm long; stamens 5,
the anthers linear oblong, to 2 cm long, ca. 1 mm wide, the connective extended
apically as a rotund petaloid appendage, ca. 0.75 mm long, sagittate at the base,
the filaments plane, ca. 4 mm long, attached ca. 6 mm from the base of the tube;
style to 15 mm long, the stigmas 5, connate into a mass, to 1.5 mm long. Fruits
cylindrical, 7-15 mm long, 4-6 mm wide, the ovarian disc conical and truncate;
seeds to 1.2 mm long, minutely foveolate.

Hamelia calycosa extends from Mexico to Peru.

coci.e: N rim of El Valle, Allen 1785 (GH. MO. NY). Cerro Pilon, 700-900 m, Duke 12 151
(MO). Cerro Pilon, Duke & Dwyer 13898 (COL. MO, SCZ, UC). N of El Valle, Lewis el al. 1797
(COL, DUKE, K, MO, UC). La Mesa, 850 m, Mori 6599 (MO), panama: Rio Cascada, 300-400 m,
Correa & Dressier 1652 (MO), veraguas: Alto Piedra, Santa Fe, 1200 m, Lao 515 (MO).

3. Hamelia macrantha Little, Caribbean Forest. 9: 274. 1948. type: Ecuador,
Spruce 6193 (K, holotype, not seen).

H. grandiflora Spruce, in Wernham, J. Bot. 49: 209. 1911. lectotype: Brazil, Spruce 6193 (K,
holotype; photo, MO); syntvpe: Colombia, Triana 1759 (F, frag.). Not LHeritier, 1788.

Shrubs or trees to 5 m tall, the branchlets terete, ultimately angular, minutely
puberulent, the nodes well spaced. Leaves ternate toward the apex, broadly
oblong to ovate, 13-23 cm long, 4-15 cm wide, rounded, obtuse or widely deltoid
at the apex, rounded, subtruncate to cuneate basally, the costa plane to promi-
nulous above and beneath, the lateral veins 11-15, membranous to chartaceous,
glabrous above, glabrous to rufous puberulent beneath; petioles 6.0-9.5 cm long,
glabrous or puberulent; stipules ovate, 3.1-3.5 mm long, acute, glabrous or rarely
pubescent. Inflorescences terminal or axillary, the dichasia many flowered; pe-
duncle terete, to 8 cm long, the floral axes to 2 cm long; bracts ovate, to 0.7 mm

long, acute, glabrate to villosulose. Flowers sessile or on pedicels to 2 mm long;
hypanthium 3-5 mm long, the calycine cup scarcely measurable, the lobes tri-
angular to ovate, to 1.3 mm long, acute, glabrous to tomentose; corolla yellow,
the tube infundibular, 2-3 cm long, constricted basally, glabrous, the lobes ovate,
to 4 mm long, glabrous; stamens included, the anthers to 20 mm long, the con-
nective produced at the apex, ovate, ca. 1 mm long, sagittate basally; style to 20
mm long, the stigmatic lobes linear, 5-8 mm long. Fruits ellipsoid, to 1.5 cm
long, red to finally lustrous black at maturity; persistent ovarian disc conical and
truncate; seeds to 1 mm long, delicately foveolate.

Hamelia macrantha ranges from Costa Rica south to Ecuador.

CANAL zone: Pipeline Road, Agua Salud. Kennedy & Andrews 1894 (MO), colon: Rio Guanche,
Correa et al. 1898 (MO), cocle: N of El Valle, road to La Mesa, 1000 m, Allen 3724 (MO). El Valle
near La Mesa. Correa & Dressier WOO (MO). Between Cerro Pilon and El Valle, Duke & Dwyer
13939 (MO). El Valle, ca. 2200 ft, Dwver 1913 (MO). Cerro Pilon, 900 m, Dwyer & Lallathin 8625.
8632 (both MO). El Valle, ca. 1000 m, Gentry 5694 (MO). Cerro Pilon, Lallathin 10-4. 28-C (both
MO). PANAMA: El Llano Carti-Tupile Road, 12 mi above Panamerican Highway, Liesner 1236 (MO).

208 ANNAI.S OF 1 HK MISSOURI BOTANICAL CiARDHN [Vol. 67

4. Hamelia magnifolia Wernham, J. Bot. 49: 210. 1911. type: Guatemala, Stand-
ley 64569 (F, holotype: NY, US, isotypes).

Shrubs or small trees, to 6 m tall, the branchlets angular to tetragonal, gla-
brous, the nodes well spaced. Leaves rarely ternate at the tips of the branchlets,
ovate to ovate elliptic, occasionally puberulent; petioles 1.0-5.5 cm long; stipules
deltoid, 4-7 mm long, to 2 mm wide at the base, glabrate. Inflorescences terminal,
of compound dichasia, often appearing cymose, the floral clusters numerous and
many flowered; peduncle to 3 cm long. Flowers with the calycine cup to 2.5 mm
long, the lobes deltoid, ca. 0.6 mm long or shorter, puberulent marginally; corolla
yellow, the tube 7.5-12.0 mm long, glabrous except often minutely puberulent at
the base, the lobes to 1.5 mm long, about as broad at the base; anthers narrowly
oblong, 6-8 mm long, the connective produced apically as an ovate or deltoid
appendage, ca. 1 mm long, falcate and ultimately acuminate, basally sagittate.
Fruits oblong, to 1 cm long or longer; persistent ovarian disc ca. 1 mm long.

Hamelia magnifolia is known only from Costa Rica and Panama. Kirkhride
& Duke 757 list the common name ''Ojo movado."

Bot AS DEI TORo: Gualaco, Cerro Galera Chorea, Allen 5055 (F). chiriqui: San Bartolo Arriba,
W of Puerto Armiielles, Cmaf 26705 (MO). Chiriquicito-Calderas Trail between Quebrada Higueron
and Gutierrez, Kirkhride & Duke 757 (MO). Puerto Armuelles. Woodson & Schery 845 (MO). San
Bartolome Peninsula, Woodson & Schery 868 (MO), darii'n: Tres Bocas on Ri'o Cuasi, Kirkhride
1361 (MO). VERAGUAs: Isla de Coiba, Dwyer 1565 (MO).

5. Hamelia patens^* Jacq., Enum. PI. Carib. 16. 1760. type: Dominican Republic,
Jacqiiin, not seen.

Shrubs or small trees, the branchlets glabrous to sparsely villous. Leaves
opposite or usually 3-nate or 4-nate (rarely 5-nate), elliptic, ovately elliptic or
obovate elliptic, 4-18 cm long, 2.2-7.8 cm wide, deltoid to acuminate at the apex,
rounded to acute at the base, the costa slender, the lateral veins 7-10, usually
prominulous beneath, chartaceous to membranous, glabrous to densely villous
beneath; petioles 0.5-4.0 cm long; stipules triangular to awl shaped, 2-6 mm long,
villous. Inflorescences terminal, solitary or of a few fascicles, the flowers in
dichasia, often helicoid, the flowers few to numerous, peduncle glabrate to vil-
lous, 0.4-4.0 cm long, the floral axes to 4.5 cm long; bracts triangular, less than
1 mm long, villous, deciduous. Flowers secund, to 2.5 cm long, sessile or sub-
sessile, the hypanthium and calycine tube bell shaped, to 4 mm long, glabrate to
villous, the lobes ovate, ca. 1 mm long, glabrate or villous, persistent; corolla
orange red to red, 12-20 mm long, 2-3 mm wide, glabrous to villous outside, the
lobes ovate, to 2 mm long, glabrous to villous; stamens often exserted, the anthers
8-12 mm long, the terminal appendage ca. 0.5 mm long, the filaments shorter
than the anthers, attached in the upper V3 of the tube; style slender, about equaling
the corolla tube, the stigmas oblong, about V5 the length of the style; ovary
5-celled. Fruits ellipsoid or rotund, to 1 cm long, black at maturity, the ovarian
disc often slightly rostrate; seeds 0.5-1.0 mm long.

54

Foralist of synonyms see FJias(l976). Only this name has been used for Panamanian material.

19801 DWYHR— FLORA OF- PANAMA (Family 179. Ruhiaccac) 209

Hamelki patens occurs in tropical Florida, throughout the West Indies, Mex-
ico, Central America, and in South America south to Bolivia.

Key to varieties (adapted from Elias, 1976):

a. Leaves villous: hypanthium and calyx villous 5a. var. patens

aa. Leaves glabrous above; hypanthium and calyx usually glabrous 5b. var. glabra

5a. Hamelia patens Jacq. var. patens.

CANAi zone: Gaillard Highway, near Summit Garden, Croat 9777 (MO). Road 2C2, Cerro Luisa,
Croat 10774 (MO). Albrook, Dwyer 6744 (MO): Dwyer & Hayden 4 (MO); Dwycr & Rohyns II
(MO): Ehingcr 847 (MO). Road K2 off road to Cocoli, Kirkhridc 80 (US). Farfan Beach, Kirkhridc
& Elias 72 (MO). Balboa, Mori & Kallunki 1711 (MO). Cocoli, Riley 133 (MO). Road K-9, Stern et
al. 308, 347 (both MO). Albrook. Stimson 5146 (MO). Contractors Hill, Stimson 5242 (MO). Madden
Dam, Stimson et al. 5085 (MO). Curundii. Tyson 1073 (MO). Chiva-Chiva Trail near Miraflores Lake,
Tyson 1404 (MO). Cocolf, Tyson 1635 (MO). Barro Colorado Island, White 122 (MO). Miraflores
Lake. White tC White 188 (MO). cttlRiQUi: Cerro Horqueta, ca. 7000 ft, Blnm & Dnyer 2608 (MO).
Bajo Mono, Boquete, Davidson 469 (MO). Finca Collins, Boquete, 5000 ft. Dnyer 7037 (MO). Bo-
quete, Cerro Horqueta, Dwyer & Hayden 7686 (GH, MO, NY, UC, US). N of San Felix, 36.2 km
by road from Interamerican Highway, Mori & Kallunki 6012 (MO). Finca Lerida to Boquete, ca.
1300-1700 m, Woodson et al. 1106 (MO), cocle: Forest behind Club Campestre, ca. 700 m. Duke
13263 (MO). Las Margaritas and El Valle, Woodson etal. 1726 (MO). DARtEw: Without other locality,
Bristan 96 (MO), herrera: Road between Las Minas and Pese, ca. 600 ft, Duke 12335 (MO). Ocu,
Ebinger 2002 (MO). 12.5 mi S of Ocii, 1200 ft. Lenis el al. 1668 (GH. MO, UC, US), los santos:
Upper Rio Pedregal. 350-400 m, Croat 34490 (MO). Los Toretos, Duyer 2424 (MO). Pcdasi, Dnyer
2483 (MO). Guaybo, Stern et al. 1903 (MO). Tonosi, Tyson et al. 3108 (MO). Panama: Isla Pedro
Gonzales. Allen 2594 (MO). Rio Congo, Carrasquilla et al. 208 (MO). 2 mi from U. of Panama.
Transisthmian Highway, D'Arcy & D'Arcy 6108 (MO). Isla Chepillo, Duke 10317 (MO). Tocumen,
Dnver 1850 (MO). Lidice, Jaramillo 1 (MO). Cerro Silvestre, Reece 29 (MO). 5 mi N of Panama
City, Stimson 5045 (MO). Arraijan, Woodson el al. 1367 (MO). Isla Taboga, Woodson et al. 1436
(MO), veraguas: 5 mi N of Santiago, Blum ct Tyson 607 (MO). 2-4 mi E of Santiago, Duke 12357
(MO). Santiago, Dnyer & Kirkhride 7419 (G, MO, US). Montijo, 8 mi S of Santiago, Tyson 6034
(MO).

5b. Hamelia patens var. glabra"^'^ Oerst., Vidensk. Meddel. Dansk Naturhist. For-
en. Kj0benhavn 1852: 42. 1853. type: Nicaragua, Oersted s.n. (C, holotype,

not seen). — Fig. 46.

bocas del toro: RR track near Milla 5, Croat & Porter 16462 (MO). Isla Bocas del Toro,
Durkee 71-150 (MO). Rio Changuinola, Dnyer 42. 2758 (both MO). Quebrada Huron, 300-400 ft,
Kirkhride <^ Duke 445 (MO). Changuinola to 5 mi S at junction of Ri'os Changuinola and Teribe,
Lenis et al. 821 (COL, MO). Changuinola. Lenis et al. 974 (MO, SCZ, UC). Shepherd Island,
McDaniel 5160 (MO). Isla Colon. Wedel 96. 2483. 2930 (all MO). Water Valley, Wedel 664, 2752
(both MO). Chiriquf Lagoon, Wedel 1236. 2598. 2901 (all MO). Nievecita, Woodson et al. 1850 (MO).
CANAL zone: Cruces Trail and Madden Dam Road, Rarllctt & Lasser 16346 (MO). Juan Mina, Bartlell
& Lasser 16514. 16531 (both MO). Fort Sherman, Blum & Dnyer 2131 (MO). Pipeline Road near
Gamboa, Clenell ct- Tvs,m 3252 (MO). Road S-10 N of Escobal, Croat 12436. 12461 (both MO).
Farfan Beach. DAre\'& DArcx 6072 (MO). Coco Solo Weather Station, Duke 4282 (MO). Camp
Piiia Duke4418A (MO). Madden Dam, Boy Scout Camp Road, Dnyer 6928, 9163 (both MO); Dnyer
& Elias 7476 (GH, MO, US), 7483 (GH, MO), 7498 (COL. MO, UC), 7499 (MO). Madden Dam,
Dnyer t6 Lallathin 8835 (MO). Fort San Lorenzo, Ehinger 459 (MO). Frijoles, Greenman & Green-
man 5156 (MO). Mandingo Road, Haxden 61 (MO). Gatun, Hladik 525, 538 (both MO). Below
Madden Dam lookout station, Kirkhride s.n. (MO). Boy Scout Camp Road, Madden Dam, Kirkhride
40. 41. 51 (all MO). Farfan Beach Road. Kirkhride & Elias 72 (MO). Rfo Chagres Bridge, SE of
Gamboa, l.asseigne 4268 (MO). Pipeline Road, 16 mi N of Gamboa, Lenis et al. 5461 (MO, SCZ).

^■^ For a list of synonyms see Elias (1976). Only this name has been applied to Panamanian

material.

210

ANNALS OF THR MISSOURI BOTANICAL GARDEN

[Vol. 67

Figure 46. Hainclia patens var. glabra Oerst.
ing branch (x 1). | After FMas 7498.]

A. Habit (xVi). [After Elias 1699.]— B. Fruit-

RR V/2 mi W of Gamboa, Ncc & Mori 3613 (MO). Rodman Marine Base, Nouickc et al. 3579 (COL,
MO, UC), Chiva-Chiva Trail, 2 mi N of Miraflores Locks, Tyson 1382. 1400 (both MO). Old Fort
Lorenzo, Tyson 1539, I55I (both MO). 5 mi NW of Cocoli, Tyson 1622 (MO). Galiin RR Station,
Tyson 22H0 (MO). 1 mi N of Summit Garden, Tyson <& Blum 1941 (MO). Farfan Beach, Tyson ct ai
3177 (MO). Miraflores, White 115 (MO). Pipeline Road, Wirlierspoon <i Witherspoon 8293 (MO).
Salamanca Hydrographic Station, Rio Pequcni, Woodson el al. 1613 (MO), barro Colorado island:
Aviles 11 A (MO): Bartlett & Lasser 16746 (MO): Croat 4053, 4273, 4392, 6362, 6370, 6814, 6962 (all
MO); Dnyer et al. 8471 (MO); Ehini^er 36 (MO); Hayden 39. 1039, 1040, 1041 (all MO); Shattuck
167, 924 (MO); Starry 76 (MO); Tyson 4211 (MO); Wetnwre & Abbe 41, 173 (both MO): Woodworth
& Vestal 622 (MO); Zetek 4270 (MO). tiiiRiQUi: Puerto Armuelles, LYArey 10071 (MO). Chiriqui-
Veraguas boundary line, old gravel main highway, Dwyer 2930 (MO). Boquete, 3300-4200 ft, Lewis
et al 642 (GH, MO), cocle: El Valle, road to La Mesa, Croat 13312 (MO). Club Campestre, ca. 700
m, Duke 13263 (MO). Cerro Pilon and El Valle, 700-900 m, Duke ct D\v\er 13898 A (MO). El Valle,
Dwyer 1833, 10498 (both MO). El Valle, vicinity of Cerro Pilon, Duxer & Correa 7969 (MO, NY,
UC). La Mesa, Dnyer & Duke 8294 (MO). Cerro Pilon, Kirkbride 1084 (MO). El Valle, Lao 280
(MO); Lewis et al. 1788 (COL, MO, UC, VEN). Rio Grande, Rosario 5 (MO). El Valle, Seibert 425

19801 DWYER— FLORA OF PANAMA (Fiimily 179. Ruhiaccac) 21 1

(MO). Llano Bonito, S of Las Margaritas, Seihcrt 529 (MO). El Valle, Tyson ct al. 3967 (MO), Las
Margaritas and El Vallc, Woodson et al. 1244 (MO), colon: Rio Gatuncillo, vicinity of Santa Rosa,
Allen 4160 (MO). Sardinilla, Blum & Tyson 507, 525 (both MO). Sabanita, Croat 11068 (MO). Por-
tobelo, DAny & D'Arcy 6684 (MO): Duycr 5008 (MO). Chagres, Gutierrez 23 (MO). 2.4 mi SW of
Pina, Lewis et al. 1852 (COL, MO, UC). 1 mi S of Gatuncillo, Stimson 5260 (MO). Achiote, Tyson
et al. 4510 (MO), darihn: Rio Tuqueza below Quebrada Venado, Bristan 1073 (MO). Rio Pirre,
Croat cC Porter 15522, 15453 (both MO). 2-3 mi S of El Real, Duke 4841 (MO). Rio Pirre, 2-5 mi
above El Real, Duke 5082 (MO). Cerro Pirre, S of El Real, 100-600 m, Duke 5292 (MO). 10 mi S of
El Real, Rio Pirre, Duke 5449 (MO). Between Rio Membrillo and Rio Yaviza, Duke 8620 (MO). Rio
Pinas, Duke 10564 (MO). Cerro Chucula, drainage Rio Pavarando, Gentry 4253 (MO). Upper Ri'o
Tuqueza, Le Clezio 132 (MO). Rio Tuquesa, Charco Chiva, Mori 6972 (MO). Without locality, Sexton
218 (MO). Yaviza, along Rio Chucunaque. Quebrada Barbua, Stern et al, 85 (MO). Trail El Real to
Pinogana, Stern et al. 274 (MO). Trail between Paya and Payita, Stern et al. 354 (MO). Tucuti,
Chepigana, Terry & Terry 1406 (MO). 3 mi N of Santa Fe, Tyson et al. 4622 (MO). Los Santos: Las
Tablas, Dwver II57A (MO). 17.8 mi S of Macaracas, 1 100 ft, Lewis et al. 1607 (MO, UC). Panama:
Panama City, Lefevre Park, Bartlett c?- Lasser 16335 (MO). Road to Cerro Campana, Croat 12047
(MO). Rio Caiiita, Jenene, Duke 3818 (MO). Rio Bayano above confluence with Rio Chepo, Duke
3985 (MO). 6 mi E of Chepo, Duke 4063, 4082 (both MO). Rio Mamonica, 4 mi beyond Chepo, Duke
5575 (MO). San Jose Island, between Bodega Bay and Ri'o Mala Puerco, Duke 12543 (MO). Piria-
Canasas Trail, Duke 14312 (MO). Rio Bayano. 1-4 mi above Piria, 100 m, Duke 14398 (MO). Alha-
juela, D\v\er 1042, 1045 (both MO). Tocumen, Dwyer 4065 (MO). I km E of El Llano, Gentry 6467
(MO). San Jose Island, Johnston 147 (MO). Transisthmian Highway and Madden Dam, Lewis et al.
5303 (MO, SCZ, UC). Panama, Moran 12 (MO). 25 km NE of Cerro Azul, Rio Piedras, 550 m, Mori
& Kallunki 3286 (MO). La Chorrera, Rodriguez 14 (MO). El Llano, Tyson 1739 (MO), san blas:
Puerto Obaldia, Croat 16959 (MO). Sasardi, Duke 10135 (MO). Rio Diablo and Rio Acuati, Duke
14877 (MO). Ailigandi, Dwyer 6848 (MO). Soskatupu Island, Elias 1662 (COL, MO, UC), 1699 (MO).
Headwaters of Rio Mulatupo, Elias 1755 (GH, MO, UC, US). Mainland opposite Playon Chico,
Gentrx 6371 (MO). 3-4 hrs up Rio Mulutupu on foot, Kirkhride 219 (MO), veraguas: Isla de Coiba,
Dwyer 1565 (MO). Tute, Dwyer 4283 (MO). 3.9-5.0 mi N of Santa Fe, 500-1000 m. Gentry 2952,
2999 (both MO).

6. Hamelia rovirosae Wernham, J. Bot. 49: 211. 1911. type: Mexico, Rovirosa
499 (NY, US, isotypes).

//. patens var. coronata J. D. Smith, Bot. Gaz. 4: 4. 1905. type: Guatemala, Tuerckheim 8532 (US,

holotype; F, NY, isotypes).
H. purpurascens Blake, Contr. Gray Herb. 52: 105. 1917. typh: British Honduras, Peck 104 (G,

holotype).

Shrubs, the branchlets villous, the hairs septate. Leaves ternate, occasionally

opposite, elliptic to ovate, 5-16 cm long, 2-7 cm wide, acuminate, attenuate at

the base, the lateral veins 3-5, arcuate, glabrous to sparsely villose above, villous
beneath; petioles 0.3-2.0 cm long; stipules linear, often widely so, 5-8 mm long.
Inflorescences terminal, dichasial or often helicoid; peduncles to 3 cm long, vil-
lous, the floral axes 1.5-3.5 cm long, each axis 3-8-flowered, the flowers secund,
sessile or on pedicels 0.1-0.2 cm long. Flowers with the hypanthium to 4.5 mm
long, villous, the calycine lobes oblong or triangular, 2.0-5.4 mm long, villous,
glabrous within; corolla reddish orange to scarlet, the tube 1.8-2.2 cm long, 2-3
mm wide, villous, the lobes ovate, 1-2 mm long, acute, villous outside; anthers
10-12 mm long, the terminal appendage ovate, ca. 0.5 mm long, the filaments 7-
9 mm long; ovary 5-celled. Frnits cylindrical, to 1.2 cm long, 0.5-0.8 cm wide,
red, finally black; seeds ca. 1 mm long.

Hamelia rovirosae is found in Mexico and throughout Central America.

Boc AS DHL TORo: Changuinola Valley, Dunlap 192 (F).

212 ANNALS OF THE MISSOURI BOTANICAL GARDHN [VoL 67

7. Hamelia sanguinea Elias, Mem. New York Bot. Gard. 26(4): 116. 1976. New
name for//, paiiciflora Standley.

//. ptiiniflom Standley, Contr. U.S. NatL Herb. 18: 139. 1916. type: Panama, Piriier 3469 (US,
holotype). not H. painiflora Willd. (1825).

Shrubs or small trees, 4-7 m tall, the branchlets terete to angular, glabrous
or puberulent, these usually forming numerous short branchlets. Leaves mostly
ternately disposed, ovate, trapeziform oblong or ovate rotund, 2.0-8.5 cm long,
1.0-3.5 cm wide, attenuate at the apex, acuminate, the acumen triangular, to 1
cm long, acute, attenuate acute to cuneate at the base, the costa prominulous or
grooved above, the lateral veins (3-)4-5, arcuate, membranous to subcharta-
ceous, glabrate above, usually minutely papillate and with numerous raphides,
sparsely short ciliate beneath, often only on the costa and the veins, ciliolate;
petioles slender, wiry, scarcely measurable or to 3 cm long, pubescent; stipules

Infl'

5(-9)-flowered, terminal on short lat-

eral branches; peduncles 1.0-2.5 cm long, the floral axes to 5 cm long, each with
up to 4 flowers, the flowers secund, those in the axils of dichasia sessile or
subsessile, the others on pedicels. Flowers with the hypanthium oblong, to 4.5
mm long, glabrous, the raphides numerous; calycine cup obsolete, the lobes
ovate, to 1.2 mm long, acute; corolla orange red to red, glabrous, the tube cylin-
drical to 2 cm long, ca. 0.3 cm wide, the lobes ovate, ca. 1.5 mm long or longer,
pubescent outside; anthers linear, 8-10 mm long, the terminal appendage ca. 0.5
mm long. Fruits suborbicular, about twice as long as wide at maturity, the per-
sistent ovarian disc rostrate; seeds to 1.2 mm long,

Hamelia sanguinea is known only from Panama.

CANAL zonl: 1 mi above Madden Lake. Duke 4479 (MO). Madden Dam, Boy Scout Camp Road,
Dum- c5 Elicis 7488 (GH, MO, UC. US), 7494 (GH. MO, US), 7514 (MO. UC), 7515 (COL, MO,
UC); Dwycr & Haydcn 21 (MO); Dwycr Sl Lallathin 8834 (MO). Madden Dam, Ehiii^er 875 (MO).
Boy Scout Road, Madden Dam, Haydcn 77 (MO). Forest of Gatiin, Hladik 512 (MO). Madden Dam
Road, Boy Scout Camp Road, Kirkhridc 50 (MO). Madden Dam, Lewis et al. 22 (MO, SCZ, UC).
Trail, Rio Petitpie, road Ft. Sherman from Gatun Locks (S2 or 82), Mori & Kcdiunki 2679 (MO). Boy
Scout Road, Madden Dam, PorWr et al. 4017 (MO, SCZ). Boy Scout Camp, Madden Lake, Tyson
5438 (MO). Boy Scout Camp Road, Madden Dam, Tyson & Dwyer 1201 (MO). Salamanca Hydro-
graphic Station, Rio Pequeni, Woodson et al. 1595 (MCi). darien: Quebrada Venadoand Peje Swamp,
headwaters of Rio Tuquesa, Bristan 1021 (MO). Near Refugio, 15-21 mi N of Santa Fe, Duke 10265
(MO). Morti Hydro, 260 m, Duke 14236 (MO).

8. Hamelia xerocarpa Kuntze, Rev. Gen. PI. 1: 284. 1891. type: Costa Rica,

Kuntze 20281 (NY, holotype).

//. mai^niloha Wernham, J. Bot. 49: 209. 1911. type: Nicaragua, Tate 200 (K, holotype, not seen).
//. rowleci Standley, J. Wash. Acad. Sci. 15: 7. 1925. type: Costa Rica, Rowlce & Stork 75 (US,
holotype).

//. haaghtii Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 116. 1940. type: Colombia, Haat/lit
2176 (F).

Shrubs or small trees, the branchlets ultimately angular, scarcely lenticellate,
villous. Leaves ovate to elliptic, 8-37 cm long, 4-14 cm wide, acuminate, atten-
uate to rounded at the base, the lateral veins 11-18, chartaceous, pilose above,
sericeous along the veins, especially in the axils; petioles 0.8-7.0(-8.5) cm long,
villous; stipules triangular, 7-17 mm long, cuspidate, often with 2 reduced lateral

198{)1

DVVYER— FLORA OF PANAMA (Family 179. Riihicucae) 213

teeth, glabrate. Inflorescences terminal, of dichasia or scorpioid cymes; pedun-
cles 1.5-5.5 cm long, villous, the floral axes to 15 cm long, ascending, often
ferrugineous pilose, the flowers secund, ca. 25 on each axis, pedicels to 0.2 cm
long. Fhrn-ers with the hypanthium 3-4 mm long, glabrous to spreading pilose;
calycine lobes ovate, 1-2 mm long, glabrous to pilose; corolla yellow salverform,
to 3.5 cm long, the tube 0.4-0.7 cm wide, glabrous to densely puberulent, the
lobes ovate, 4-8 mm long, acute: anthers 10-16 mm long, acute, the connective
extended for ca. 0.1 mm at the apex, acute, sagittate at the base, the filaments
8-15 mm long; ovary 5-loculate, the style ca. 20 mm long, the stigmas 5, to 5 mm
long.

Hamelia xerocarpa is known from Costa Rica, Panama, and Colombia.

BOCAS DEL TORo: Changuinola Valley, Lincoln Creek, Dunlap 572 (F). Fish Creek lowlands,
Wcclel hi (F MO). Carretera, Wedel 648 A (MO). Water Valley, Wedel 742 (F, MO). Chiriqui
Lagoon, Wcdel 1171 (F, MO). Old Bank Island, Wcdcl 2024 (MO, US). Fish Creek lowlands, Wedcl
2381 (F, MO), dariizn: Rio Tuquesa, Middle Tuquesa Mining Co. Camp called Charco Peje, Mori
7086 (MO). Punta Guayabo Grande and Piinta Guayabo Chiquita, Stern c5 Chambers 177 (MO, NY).
VERAGUAS: NW of Santa Fe, 1 1 km from Escuela Agricola Alto de Piedra, 450-550 m, Mori et at.

3828 (MO).

45. HEMIDIODIA

Hemidiodia Schum. in Mart., Fl. Bras. 6(6): 29. 1888. type: Spernmcoce ocim-
ifolia Willd. = Hemidiodia ocimifolia (Willd.) Schum.

Perennial herbs, often woody and profusely branching from the base. Leaves
narrow, the veins conspicuous; stipules adnate to the petioles, forming a setif-
erous sheath. Inflorescences axillary, epedunculate, the flowers fasciculate.
Flowers with the calycine lobes 4; corolla funnelform, the lobes 4, valvate; sta-
mens attached near the mouth, exserted. Fruits of 2 mericarps separating at
maturity from a median septum, opening basally.

Hemidiodia consists of a single species which extends from Mexico to South
America. It also occurs in the West Indies and in Malaya.

Literature:

Bremekamp, C. 1934. Diodia (Gronov.) L. in Notes on the Rubiaceae of Suri-

nam. Rec. Bot. Neerl. 31: 305

(Willd.) Schum. in Mart

Fig

47.

Spcnnacoce ocimifolia Willd. ex R. & S. Syst. Veg. 3: 530. 1818. type: not seen.

Suhshruhs to 1 m tall, often branching from the base, the stems usually an-
gular, smooth, puberulent or glabrous, the bark thin, often exfoliating, the nodes
well spaced. Leaves narrowly lanceolate, ovate lanceolate to trapeziform lan-
ceolate, to 6.5(-7.5) cm long, to 1.3 cm wide, acute at the apex, usually attenuate
acute at the base, the costa prominulous to plane, the lateral veins ca. 7, strongly
arcuate, chartaceous, puberulent or farinose puberulent above and beneath, the
petioles almost absent or 0.3-1.0 cm long, ca. 0.25 mm wide; stipules adnate to

214

ANNALS OF THE MISSOURI BOTANICAL GARDLN

[Vol. 67

Figure 47. Hcmidiodin ocimifotia (Willd.) Schum.— A. Habit (xVi).— B Flower (x6)— C
Fruits (x5). [After Mi Daniel 6H75.]

the petioles, the body fan shaped, 2-5 mm long, membranous, turgid, puberulent,

with 8

Infl

the leaf axils, to 0.6 cm long. Flowers with the hypanthium oblong, ca. 1.5 mm
long, petaloid, glabrous, the calycine teeth 4, scarcely visible or as triangular

1980] DWYHR— FLORA OF PANAMA (Family 179. Riihiaccac) 215

points, to 0.15 mm long, glabrescent; corolla with the tube short cylindrical, ca.
1.3 mm long, petaloid, glabrous, the lobes 3-4, ca. equal to the tube or much
shorter, oblong, obtuse, glabrous outside; stamens 3-4, the anthers oblong ro-
tund, ca. 0.2 mm long, the filaments shorter than or as long as the corolla lobes,
attached at the mouth; stigmas compressed rotund, ca. 0.15 mm wide, the style
ca'. 1.3 mm long, the septum of the ovary thin. Fruits oblong, 3-4 mm long,
bicarpellate, each carpel 1-seeded, separating from a median septum, dehiscing
at the base, densely pubescent, the calycine lobes persistent, to 0.2 mm long.

Mexico

Meo

leaves are "used for kidney and urinary disorders, boiled as a tea."

BOCAS DFi TORo: Station Milla 7.5 on Changuinola-Almirante Railroad, Croat 38095 (MO). Chan-
guinola River. 10-15 mi inland. Lewis et id. 887 (G, H, K, MO, NY, UC, US). Chiriquicito to 5 mi
S along Rio Guarumo. Lewis et al. 2075 (MO, SCZ, UC). canal zone: S-5 Road, Caribbean side,
Blum 1505 (MO). Barro Colorado Island, Croat 12549. 12840 (both MO). Pipeline Road within 5 mi
of Gamboa Gate, DArcy 9274 (MO). Hill C6, Fort Sherman, Duke 4354. 4374 (both MO). Pipeline
Road, 16 mi N of Gamboa, Lewis et al. 5464 (MO). Navy reservation, N of Gamboa, Rohyns 69-50
(MO), cocle: El Cope, Burch et al. 1374 (COL, MO, UC). Cerro Pilon, Duke 10275 (MO); Dwyer
et al. 4538 (MO, SCZ, UC), 4563 (MO). Panama: Cerro Azul, Croat 11568 (MO). Cerro Campana,
Croat 12048 (MO). Cerro Jefe, D'Arcy 3967 (MO). Cerro Campana, Duke 5996 (MO). Cerro Azul,
Duke 9352 (MO). Rio Pacora, Duke 12016 (MO). Cerro Campana, Dwyer & Kirkhride 7822 (MO).
Chagres, Fendler 145 (MO). Cerro Jefe, Folsom et al. 2533 (MO). Rio Pasiga, Gentry 2362 (MO).
Cerro Jefe, Gentry 6755 (MO); Kirkhride & Crehbs 8 (MO). Cerro Azul, Lewis et al. 228 (F, GH, K,
MO, NY, UC, US). Cerro Campana, McDaniel6875 (MO). Road to Altos de Pacora, Mori & Kallunki
2351 (MO). TV tower, Cerro Penon, 3 km S of Alcalde Diaz, Nee 8591 (MO). Cerro Campana, Porter
et al. 4183, 4289 (both MO), 4890 (MO, SCZ). Cerro Azul, Tyson 2082 (MO). Cerro Jefe, Tyson et
al. 4309 (MO). Macapale Island, Madden Lake, Tyson 5484 (MO). W of EI Llano, Tyson 6876 (MO).
Arraijan, Woodson et al. 1396 (MO), san blas: Ailigandi, mainland, Lewis et al. 67, 83 (both MO).
veraguas: Ustupo, D'Arcy 9518 (MO). Road from Santa Fe past Agricultural School to base of
Cerro Tute, Folsom 1576 (MO). 3.9-5.0 mi N of Santa Fe, Gentry 2975A (MO).

46. HILLIA
Hillia Jacq., Enum. PI. Carib. 3, 18. 1760. type: H. parasitica Jacq.

Fereira Veil, ex Vand., Fl. Lusit. Bras. Sp. 21. 1788. type: F. vellozoana J. A. & J. H. Schultes.
Saldanha Veil., Fl. Flum. 141. 1825, Not Bureau 1868. lectotype: S. illustris Veil.

Shrubs, mostly epiphytic, glabrous. Leaves succulent, drying coriaceous, pet-
iolate; stipules thin, caducous. Flowers often large, solitary, terminal; hypan-
thium obovoid or cylindrical, the calycine cup absent, the lobes 2-5 or absent,
caducous or persistent in fruit; corolla salverform or funnel shaped, the tube
elongate, expanded toward the mouth, glabrous, the lobes 3-7, contorted in bud;
stamens included; ovary 2-celled, the ovules numerous. Fruits capsular, narrowly
oblong, or cylindrical, the 2 valves dehiscing septicidaljy; seeds imbricate, basally
appendaged, bearing a distal tuft of hairs.

I

HiUia. a genus of about 20 species, occurs in the West Indies, Mexico, Central
America, and South America.

a. Leaves small, 0.6-1.3 cm long.

b. Leaves ca. 0.7 cm long, rounded at the apex; corolla tube ca. 1.5 cm long

1 . H, chiapensis

bb. Leaves 0.6-1.3 cm long, tapering toward the apex; corolla tube to 2.8 cm long

5. H. panamensis

216 ANNAl.S OF'l'Hr, MISSOURI BOTANICAL GARDI.N [Vol. 67

aa. Leaves large, 2-8 cm long.

c. Leaves 2-3 times longer than wide: calycine lobes ca. 1.7 cm long 3. //. ligulifolia

cc. Leaves less than twice as long as wide; calycine lobes less than 1 cm long or absent.

d. Corolla tube 2-3 cm long 4. H. pal

ffiana

dd. Corolla tube 4-7 cm long.

e. Trees to 5 m tall; leaves 0.6-2.8 cm wide, rounded at the apex, drying tan

beneath; corolla tube 6-7 cm long 2. H. chinquiensis

ee. Epiphytic shrubs to 2 m tall: leaves to 4 cm wide, usually deltoid at the

apex, drying black brown beneath; corolla tube 4-5 cm long

6. H. tetrandra

1. Hillia chiapensis Standley, J. Wash. Acad. Sci. 16: 16. 1926. type: Mexico,
Purpus 262 (US, holotype).

la. Hillia chiapensis Standley var. chiapensis.

Epiphytic shrubs, much branched, heathlike in habit, the branches rough,
4-angled, the angles delicately alate, glabrous, the nodes constricted, the point
of petiole insertion depressed and with an annular orifice. Leaves cricoid, elliptic,
oblong elliptic or oval, to 0.7(-0.9) cm long, rounded at the apex, acute to obtuse
at the base, the costa and lateral veins not visible in dried material, coriaceous,
glabrous, marcescent; petioles to 0.2 cm long; stipules deciduous, occasionally
persistent at tip of branchlets, obovate to oblong, 3-4 mm long, rounded, gla-
brous, venose. Flowers solitary, sessile; corolla tube ca. 15 mm long, the lobes
ca. 7 mm long. Fruits capsular, to 3 cm long.

Hillia chiapensis ranges from Mexico to Panama.

Panama: Finca El Indio, Cerro Jefe, Corrca ct til. 1770 (MO). Summit of Cerro Jefe and forests
along road beyond summit, to 2700 ft, Ilaydcn 1021 (COL, DS, MO, UC).

lb. Hillia chiapensis subsp. grandifolia Dwyer,^'' type: Panama, Nee 9698 (MO,
holotype).

Leaves to 3(-4.5) cm long, to 1.2(-1.8) cm wide; fruits to 4 cm long.

The new subspecies is known only from Panama. It differs from //. chiapensis
subsp. chiapensis in having much wider leaves and more elongate fruit.

VERAGUAs: Lower montane wet forest 6-7 km W of Santa Fe on new road past agricultural
school, 2900 ft, Ncc 9698 (MO).

2. Hillia chiriquiensis Dwyer.-^^ type: Panama, Mori & Kallunki 5810 (MO, ho-
lotype). — Fig. 48.

Trees to 5 m tall, branching freely, cricoid in appearance, ultimately black
when dry, the branchlets thick, the nodes well spaced or crowded, ultimately

^'' Hillia chiapensis subsp. grandifolia Dwyer, subsp. nov. Folia ad 3(-4.5) cm longa, ad 1.2(-1,8)
cm lata; fructus ad 4 cm longi.

" Hillia chiriquiensis Dwyer, spec. nov. Arborcs ad 5 m altae ramulis ericoideis. Folia angusto-
obovata vel oblonga, 2.0-6.5 cm longa, 0.6-2.8 cm lata, apice rotundata venis lateralibus ca. 4 arcuatis
plerumque invisibilibus. coriacea glabra; petiolis ad 0.8 cm longis; stipulis terminalibus, ad 1.5 cm
longis. Florcs terminales solitarii, lobis calycis 4 oblongis, ca. 0.4 cm longis, ca. 0.8 cm latis; corolla
viridi, tubo infundibuliformi, ad 7 cm longo, 0.3 cm lato, lobis 4-5, antheris subsessilibus, ca. 1.2 cm
longis, apice connectivo ad I mm producto. Fructus non visi.

19801

DWYER— FLORA OF PANAMA (Family 179. Riihlaicac)

217

FiGURF 48 Hillia. A-C. H. chiriquiensis Dwyer.— A. Habit (xi/2).— B. Flower, opened to
show interior (X 3).— C. Ovary cross section (x 11. 5). [After Mori & Kalliinki58IO.]—D. H. tetrcwJra
Swartz. Fruit (x.6). [After Dwyer <& Ganger 7377.]

angular, nodose in appearance. Leaves crowded at the apex of the twigs, nar-
rowly obovate oblong, 2.0-6.5 cm long, 0.6-2.8 cm wide, rounded at the apex,
acute or cuneate toward the base, the costa scarcely prominulous above, only
visible on the proximal side of the blade, the lateral veins ca. 4, inconspicuous,
at times invisible above, arcuate, the margin lightly revolute, coriaceous, discol-
orous, smooth, drying tan beneath, glabrous; petioles to 0.8 cm long, thick, to
0.25 cm wide; stipules deciduous, persistent only at the tips of the twigs, oblong,

218 ANNALS OF- THE MISSOURI BOTANICAL GARDHN [Vol.67

to 1.5 cm long, obtuse, coriaceous, glabrous. Flowers terminal, solitary, calycine
lobes 4, oblong, 4-6 mm long, 2-4 mm wide, obtuse, coriaceous, glabrous;
corolla green, the tube infundibuliform, 6-7 cm long, to 0.3 cm wide, dilated
toward the apex, the lobes 4-5, white with a pink tinge, ovate oblong or oblong,
to 2.5 cm long, to 1.3 cm wide, obtuse, glabrous; stamens 4-5, the anthers sub-
sessile, oblong, ca. 1.2 cm long, the acumen to 1 mm long, sagittate at the base,
attached about 1.5 cm below the mouth; style linear, to 5.5 cm long, ca. 0.1 cm
wide, lignose when dry, glabrous. Fruits not seen.

Hillia chiriqitiensis is known only from Panama. It is remarkable in being a
fair-sized tree and in having the longest corolla among the Panamanian Hillia.

CHiRiQUi': N of San Felix, Chiriqui-Bocas del Toro border, on Cerro Colorado Copper Mine
Road, lower montane rain forest, 5000-5500 ft, Mori & Kalluiiki 5810 (MO).

3. Hillia ligulifolia Dwyer.^^ type: Panama, Ncc 9812 (MO, holotype).

Epiphytic trees, 3.5-5 m tall, the branchlets numerous, glabrous, drying dark,
tending to be opposite and arcuate, ultimately quadrangular, the nodes well
spaced below, crowded at the tips of the twigs. Leaves narrowly oblong or vague-
ly obovate oblong, to 8 cm long, 1.5-2.5 cm long, the blades usually 2'/2 times
longer than wide, conspicuously rounded or truncate at the apex, basally cuneate
to attenuate acute, the costa thick and prominulous proximally, evanescent dis-
tally, the lateral veins 4-5, inconspicuous, scarcely prominulous, strongly as-
cending, coriaceous, concolorous, drying dark brown, glabrous; petioles variable
in length and thickness, 0.1-1.5 cm long, the thicker petioles short, the blades
decurrent; stipules deciduous, often persisting at the tips of the twigs, of the same
shape and texture as the leaves, to 2 cm long. Flowers solitary, terminal, sessile,
the hypanthium enclosed by 2 bracts (stipules?) and by 4 bracteoles of the same
texture as the leaf blades, to 2 cm long; bracts connate, each part with 2 subrotund
lobes, to 0.6 cm long; bracteoles 4; calycine lobes 4, narrowly oblong, 1.0-1.4
cm long, 0.2-0.3 cm wide, unequal, coriaceous, glabrous; corolla tube ca. 4 cm
long, the lobes 1 cm long, white, the lateral margins tinged with red; stamens 5,
the anthers narrowly oblong, ca. 12 mm long, the connective produced at the
apex; style slender, to 5.5 cm long, the stigmas oblong, ca. 5.5 mm long, ca. 2.5
mm wide. Fruits sessile, narrowly oblong, cigar shaped, often somewhat falcate,
(3.5-)5-6 cm long, to 1 cm wide before dehiscence, rounded at the apex, obtuse
at the base, drying deep red, glabrous, the calyx often persistent, the lobes to 1.8
cm long, to 0.6 cm wide, rounded at the apex, venose.

Hillia lii^ulifolia is known only from Panama. It is distinguished from other
Hillia of Panama by its very large calycine lobes and its leaves, many of which

^^ Hillia ligulifolia Dwyer, spec. nov. Arhores. Folia angusto-obionga vel vix obovato-oblonga,
saepe liguliformia, ad 8 cm longa, i.5-2.5 cm lata, apice rotunda vel truncata, basim versus cuneata
ad attenuato-acuta, venis lateralibus 4-5 inconspicuis. coriacea; petiolis 0.I-L5 cm longis; stipulis
foliis similibus, ad 2 cm longis. Florcs solitarii terminales sessiles; lobis calycis angusto-oblongis,
1.0-1.4 cm longis, 0.2-0.3 cm latis; corolla tubo 4 cm longo, lobis I cm longis: antheris angusto-
oblongis: stylo ad 5.5 cm longo, stigmatibus ca. 5.5 mm longis, ca. 2.5 mm latis. Fnictiis sessiles,
angusto-oblongi, saepe falcati, (3.5-)5-6 cm longi, ad 1 cm lati, obtusi basi, apice rotundati; lobis
calycis persistentibus ad 1.8 cm longis, ad 0.6 cm latis.

19801 DWYER—FLORA OF PANAMA (Family 179. Riihiaceae) 219

are truncate at the apex. Many of the leaf blades have a tonguelike or straplike
shape. Noteworthy is the fact that the petioles are quite variable in length and
thickness. The persistent calycine lobes give the large fruits a striking appearance.
Only one other Panamanian large-leafed Hillia is known in the fruiting stage: H.

tetrandra.

CHiRiQUi: Cerro Colorado, 34.1 km beyond bridge over Rio San Felix, 1390 m, Croat 37199,
37256 (both MO). Cerro Colorado, 35.6 km from Rio San Felix bridge, 1390 m, Sullivan 388 (MO).
PANAMA: Cerro Jefe, ca. 1000 m, Mori & Joly 7926 (MO), veraguas: Lower montane wet forest 6-
7 km W of Santa Fe on new road past agriculture school, 2900 ft, Ncc 9812 (MO).

4. Hillia palmana Standley, J. Wash. Acad. Sci. 18(6): 164. 1928. type: Costa
Rica, Maxon & Harvey 8045 (US, holotype).

Epiphytic shrubs, the branchlets terete, finally compressed, black when dry
and covered with gray spots, glabrous. Leaves persisting toward the apex of the
branchlets, obovate or obovate rotund, 1.5-4.5 cm long, 1.0-2.2 cm wide, round
or truncate at the apex, cuneate basally, the costa scarcely distinguishable, the
lateral veins not visible, thickly coriaceous, concolorous, glabrous, the margin
slenderly callose; petioles thick, to 1 cm long, to 0.3 cm wide; stipules deciduous,
persistent toward the apex, obovate rotund, to 1.5 cm long, rounded, coriaceous,
glabrous, the veins conspicuous or evanescent. Flowers terminal, solitary, to 4
cm long, enclosed partially by 2 large bracteoles, these compressed rotund, to 2
cm long, scarious, glabrous, venose; hypanthium rectangular, to 1.8 mm long,
glabrous, the calycine cup none, the lobes obviously absent; corolla white, the
tube infundibuliform, 2-3 cm long, ca. 0.3 cm wide, rigid, glabrous, the lobes 4,
compressed rotund, to 1 cm long, wider than long, glabrous; stamens subsessile,
the anthers linear oblong, ca. 3 mm long, affixed near the mouth; style ca. 6 mm
long, the stigmas 2, subulate, ca. 3 mm long. Fruits not seen.

Hillia palmana is known only from Costa Rica and Panama. The leaves are
strikingly ligulate. As few flowers are available for dissection I elected to dissect
one whose corolla measured only 2 cm in length. No ovules were located; this
coupled with the absence of fruit in the two known collections is cause for spec-
ulating that the flowers are unisexual. In this dissection the style and stigmas
measured only about Vi the length of the corolla tube.

veraguas: N W of Santa Fe, 1 km from Escuela Agricola Alto de Piedras, Mori & Kallunki 4873
(MO).

5. Hillia panamensis Standley, N. Am. Fl. 32(2): 117. 1921. type: Panama, Pittier
3190 (US, holotype).

H. hatlicwayi Fosberg, Sida 2: 387. 1966. type: Costa Rica, Hathcsvay 1371 (US, holotype).

Epiphytic trees, to 4 m tall, the branchlets cricoid in habit, ultimately bearing
very short and numerous branchlets, the nodes proximate, the leaves tending to
persist only at the tips of the twiglets. Leaves elliptic oblong, 0.6-1.2 cm long,
0.4-0.5 cm wide, tapering slightly toward the apex, subacute to obtuse at the
base, frequently cuneate, the costa and lateral veins invisible above, scarcely
visible beneath, the lateral veins ca. 2, strongly arcuate ascending, subcoriaceous.

220 ANNAl.S OF- THl- MISSOURI BOTANICAL GARDEN [Vol. 67

concolorous, drying dark, glabrous; petioles absent or to 0.2 cm long; stipules
(Standley) narrowly oblong, to 4 mm long, obtuse, brownish. Flowers terminal,
solitary; hypanthium narrowly oblong or turbinate, ca. 2 mm long, glabrous, the
calycine lobes 4, linear oblong, to 9 mm long, ca. 0.8 mm wide, obtuse, gla-
brous, drying red, venose; corolla white, the tube narrowly cylindrical, to 2.8 cm
long, ca. 0.8 cm wide in the middle, glabrous inside and out, the lobes 4, ovate
triangular, to 1.2 cm long, obtuse, glabrous, petaloid, venose; stamens 4, the
anthers subsessile, affixed immediately below the mouth, oblong, ca. 1.8 mm
long, obtuse, the style linear, ca. 12 mm long, the stigmas 2, ca. 3 mm long,
scarcely wider than the style. Frnils linear oblong, 1.0-2.5 cm long; seeds oblong
with terminal hairs to 6 mm long.

CHiRiQuf: Cerro Horqueta, 1650 m, Croal 26957 (MO); Pitticr 3190 (US).

6. Hillia tetrandra Swartz, Prodr. 58. 1788. type: Jamaica, Swarlz (not seen).

H. ma.xoiiii Standley. J. Wash. Acad. Sci. 18: 163. 1928. type: Nicaragua. Maxon 7501 (US. holo-
type).

Small shrubs, to 2 m tall, often epiphytic, the twiglets terete or angular,
smooth, glabrous, the nodes well spaced. Leaves oblong rotund or ovate oblong,
mostly equilateral, to 7 cm long, to 4 cm wide, usually deltoid, occasionally
obtuse at the apex, obtuse, rounded or cuneate at the base, the costa plane
proximally above, subprominulous proximally beneath, evanescent distally, the
lateral veins ca. 5, arcuate, evanescent above and beneath, coriaceous, drying
dark brown; petioles to 1 cm long, to 0.25 cm wide, shiny, puberulent to gla-
brescent, the lamina decurrent on the upper side; stipules oblong, to 1.5 cm long,
tapering, obtuse, coriaceous, glabrous. Floners terminal, solitary, subsessile;
hypanthium oblong, ca. 10 mm long, ca. 3.5 mm wide, glabrous, the lobes 5,
often deciduous, 0.5-1.0 cm long or perhaps scarcely measurable; corolla white
or yellow with the tube narrowly cylindrical, 4-5 cm long, 0.2-0.4 cm wide, often
subfalcate, glabrous inside and out, the lobes 5, oblong, ca. 1.5 cm long, ca. 1.2
cm wide, glabrous; stamens 5, the anthers oblong, ca. 1 1 mm long, 2 mm wide,
obtuse, the filaments short, ca. 1 mm long; style linear, ca. 5.5 cm long, ca. 0.9
mm wide, the stigmas about as wide as the style, to 12 mm long, villose. Fruits
sessile, narrowly oblong, 5(-7) cm long, 0.4-0.9 cm wide, truncate or rounded,
smooth, glabrous, lignose; seed ca. 2 mm long with a tuft of tawny hairs to 12
mm long.

Hillia tetrandra occurs in the West Indies and throughout most of Central

America.

Ma

these are obtuse at the apex. In other respects it resembles the Panamanian

M

when collected at 5:00 p.m. but opened in bag by 7:00 p.m. Moth pollinated?"

CHiRioui': San Felix, Ma.xon d Harvey 8092 (US). N of San Felix at Chiriqui-Bocas del Toro
border. Cerro Colorado Copper Mine Road, 5000-5500 ft, Mori c'i Kallunki 5815 (MO), cocle: Cerro
Valle Chiquito, Scibcrt 495 (MO), colon: Santa Rita Ridge, Dnxer & Gentry 9394 (MO). Panama:
Cerro Campana, Alien 1872 (F); Croa! 14801 (MO). Goofy Lake, road toCerro Jefe. Correa &
Dressier 941 (MO). Cerro Campana, Dressier 3522 (MO). Cerro Jefe, /)iivt^/- & Gauf>er 7377 (MO);
Mori & Kallunki 6095 (MO), veraguas: 5 mi W of Santa Fe, Croal 23079 (MO).

1980) DWYF.R— FLORA OF PANAMA (Family 179. Riihicucac) 221

47. HIPPOTIS

Hippotis Ruiz & Pavon, Prodr. Fl. Peruv. 33. 1794. type: H. trifolia Ruiz &
Pavon.

Shrubs or trees. Leaves with the blades membranous or thinly chartaceous,
the intervenal areas lineolate, petiolate, the stipules large, caducous. Inflores-
cences cymose (or flowers occasionally solitary). Flowers large, 5-merous, tend-
ing to become zygomorphic, the calyx large, spathaceous or bilabiate; corolla
tubular or funnelform, usually curved, the lobes short, plicate valvate; stamens
often unequal in length, inserted near the middle of the tube; style slender, elon-
gate, the ovary 2-celled, the ovules numerous, the placentas adnate peltately to
the septum. Fruits baccate, round; seeds numerous, foveolate.

Hippotis, a genus of about a dozen species, is known from Panama, Colombia,
Venezuela, Ecuador, and Peru.

1. Hippotis albiflora Karst., Fl. Colomb. 1: 33. 1858. typi:: not seen.— Fig. 49.

Diiroia panamcnsis Dwycr, Ann. Missouri Bot. Card. 55: 38. 1968. type: Panama, Kiikhridc &
Duke 571 (MO, holotype).

Trees to 8 m tall, the branchlets subterete, drying lightly rimose, densely
pilose terminally, nodose. Leaves obovate elliptic, to 26 cm long, to 15 cm wide,
widely deltoid at the apex, acuminate, the acumen to 1.5 cm long, the costa
prominulous above, prominent and convex beneath, the lateral veins ca. 12, at
first strict, then slightly arcuate toward the margin, 2.0-2.5 cm apart in larger
blades, prominent beneath, the secondary veins irregularly pinnatiform, the ul-
timate veins enclosing minute, parallel striae, the hairs appressed, white, ca. 1-
2 mm apart, somewhat scabrous, pilose beneath, with some hairs more rigid and
longer, ciliolate; petioles 1.0-2.5 cm long, lignose, pilose; stipules free, decidu-
ous, ovate elliptic, here the younger ones to 1.5 cm long, densely pilose. Flowers
1(7-3), terminal or axillary; hypanthium oblong, 8-10 mm long, densely appressed
pilose, the hairs yellowish white, ascending, 4-5 mm long, the calycine cup tu-
bular for 10 mm then spathaceous for ca. 15 mm, thickly petaloid, densely villose
outside, the hairs stiff, white, to 2.5 mm long, glabrous and eglandular within;
corolla white or greenish white, curved, trumpet shaped, to 45 mm long, ca. 6
mm wide in the middle, densely white pilose outside, glabrous within, stiffly
petaloid or subcoriaceous, the lobes 5, elliptic or ovate, ca. 1 cm long, ca. 0.8 cm
wide, densely white pilose outside; stamens 5, included, the filaments slightly
unequal, to 11.5 mm long and to 0.8 mm wide, plane, densely pilose, attached
below or near middle of the tube, the anthers oblong, ca. 3 mm long; ovarian disc
oblong rotund, to 3 mm long, the style elongate, subplane, ca. 1 mm wide, the
stigmas not seen. Fruits nutlike, elliptic, to 3 cm long, to 1.5 cm wide, often 5-
7-ribbed, densely pilose, the hairs yellowish white, the calyx persistent, about as
long as the fruit; seeds elliptic oblong, ca. 2 mm long, 3-angled, foveolate under
magnification.

Hippotis albiflora occurs in Costa Rica, Panama and Colombia. Kirkbride &
Duke 571 notes the: "corolla tube greenish white, anthers yellow." McDaniel
5079 notes that the plant is a "vine" but this is highly unUkely.

222

ANNALS OF THP: MISSOURI BOTANICAL GARDEN

[Vol. 67

FiGURF 49. flippotis alhiflora Karst. Habit (xVi). [After Kirkhridc & Duke 57/. J

cosiA rica: HrRi-DiA: Puerto Viejo, Sarapiqui, 10^26'N, 84°0rW, Finca La Selva, Harkhorn
1044 (MO). Panama: bocas del toro: Duwebdulup Peak, N of Rio Teribe, across from West Huron,
Kirkhriile & Duke 571 (MO), 4 mi N of Almirante, McDaniel 5079 (MO). Colombia: c hoco: Rain
forest NW of Alto Curiche, ca. 300 m, Duke ct IJroho 1 1258 (MO), santandlr: Magdaiena Valley,
Campo Capote, 30 km E of Carare, 300 m, Gentrx et al. 20083 (MO).

48. HOFFMANNIA

H

O'Hi^^insia Ruiz & Pavon, PL Peru L 55. 1798. type: O. obovata R. & P.

lUg^insla Pers., Syn. PI. L 133. 1805. i.ectotvpiz: //. obovata Pers., "Higginsia'' orth. mut. Steudel
Nom. ed. 2, I: 767. 1840.

Evosniia caripensis H. & B., PL Aeq. 2: 165. 1817. typf: E, caripensis H. & B.

Cainpylohotrys Lem.. PL Serres Jard. Eur. 3: 269. 1847. typi:: T. discolor Lem.

Shrubs, subshrubs or small trees, rarely herbs. Leaves membranous to thickly
chartaceous, usually replete with cystoliths, petiolate or sessile, the stipules de-
ciduous. Inflorescences axillary, occasionally subtended by roots (rhizomatous),
cymose, usually pedunculate, the flowers spreading or conglomerate. Flowers
usually pedicellate, the hypanthium with numerous cystoliths, the calycine tube
short, the teeth often 4, usually shorter than the hypanthium; corolla tube usually
reduced, glabrous within, the lobes imbricate; stamens 3-5, the anther connective
occasionally produced apically; style elongate, slender, glabrous, the stigmas

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaccae) 223

usually 2, if connate then clavate, linear or lanceolate. Fruits baccate, usually
2-celled; seeds numerous, small, foveolate.

Hoffmannla ranges from Mexico to Argentina. There are more than 150
species in the genus, Vi of which may be reduced to synonomy when a monograph
of the genus is written. This is the most challenging among the larger genera of
Rubiaceae in Middle America. The fact that the stipules drop off when the leaf
buds emerge contributes to the taxonomic difficulty of the genus. The flowers
and fruits exhibit little variability for the size of the genus.

Literature:

Dwyer, J. D. 1969. The Genus Hoffmannia (Rubiaceae) in Panama. Annals of

the Missouri Botanical Garden 56: 269-286.
Standley, P. C. 1934a. North American Flora. Rubiales— (Pars) 32: 190-208.

Williams, L. 1973. Hoffmannias from Mexico and Central America. Fieldiana
(Botany) 36(6): 51-60.

1. Leaves bullate.

2. Leaves with lateral veins 22-25 ^2. H. pustidata

T. Leaves with lateral veins 8-14 4. //. bidlata

r. Leaves smooth, not bullate.

3. Petioles with conspicuous lateral inflations (vescicles).

4. Inflorescences axillary and occasionally terminal; inflations of petioles, if present,

poorly developed; lateral veins of leaves ca. 10 18. H. niorii

4'. Inflorescences strictly axillary; inflations of petioles well defined; lateral veins of

leaves 1 1-25.

5. Leaf blades to 25 cm long; petioles to 4 cm long; fruits red ____ 26. H. vescicniifera

5'. Leaf blades to 40 cm long; petioles to 13 cm long; fruits white ____ 12. //. kirkhridei
3'. Petioles without conspicuous lateral inflations.

6. Leaves petiolate.

7. Hypanthium with hairs to 1.7 mm long; inflorescence usually contracted and pe-
duncle short.

8. Inflorescences densely capitate and fasciculate, the peduncle much reduced
or absent, flowers not unilaterally or helicoidally disposed.

9. Small trees; petioles 0.5-0.8 cm long; stigmas clavate 1. //. aeruginosa

9'. Herbs; petioles 1-4 cm long; stigmas linear oblong 3. H. areolata

8'. Inflorescences patulous, the peduncle to 1 .5 cm long, flowers unilaterally and

helicoidally disposed '0- H. gentry!

7'. Hypanthium with hairs less than 1 mm long or absent; inflorescences usually

pedunculate and spreading.

10. Fruits (immature) at least 2 times longer than wide; stigmas linear oblong,

spreading, ca. '/3 the length of the style.

11. Peduncles absent or to 1 cm long; plants terrestrial, 1.5-2.5 m tall.
12. Leaves charlaceous, the lateral veins ca. 4; peduncles to 10 mm

long 23. H. rexmontis

12'. Leaves membranous; lateral veins 10-11; peduncles padlike and

scarcely measurable, occasionally to 8 mm long

21.//, psychotriifolia

11'. Peduncles to 4.5 cm long; plants probably epiphytic, ca. 0.25 m tall

4. //. hidlata

10'. Fruits less than 2 times longer than wide; stigmas if linear oblong, very

short and spreading.

13. Corolla lobes 2 or more limes longer than the tube; leaf blades often

with patches of raphides on upper surface forming a thin crust

14. //. lenticellala

13'. Corolla lobes less than 2 times the length of the tube; leaf blades
lacking patches of raphides forming a crust.
14. Calycine lobes to 12 mm in fruit; seeds visible through the wall

of the fruit 16. //. longicalyclna

14'. Calycine lobes 1-3 mm long; seeds not visible through the wall

of the fruit.

224 ANNALS OF T\\\l MiSSOLRl BOTANICAL GARDLN [Vol. 67

15. Inflorescences subtended by roots; stamens 3^.

16. Leaves oblong rotund; flowers yellow to yellow brown
or yellow red; corolla lobes ca. 3 times the length of
corolla tube; anthers 4; stigmas clavate rotund

7. //. davidsoniae

16'. Leaves oblong or obovate elliptic; flowers red; corolla
lobes less than 2 times the length of the corolla tube;
anthers usually 3; stigmas narrowly oblong .. 8. //. eUasii
15'. Inflorescences not subtended by roots; stamens 4(-6), often

3 in //. ostaurca.

17. Leaves in unequal pairs (anisophyllous); flowers 1-2

per axil 27. H . wifherin^ioidcs

17'. Leaves in pairs similar in size; flowers many per axil.

18. Anther connective not produced apically or ba-
sally.

19. Flowers ca. 2 cm long at maturity; calycine

lobes 4-5(-6); stamens 3-4; ovaries 3-4 loc-

ulate

15. //. U'W'isiuna

19'. Flowers to 1.5 cm long at maturity; calycine
lobes regularly 4; stamens (3-)4(-5): stigmas
2-3; ovaries 2-3 loculate.
20. Leaves membranous; stamens 3(-4)

19. H. osianrea

20'. Leaves thin chartaceous to subcoria-
ceous; stamens 4(-5).
21. Stigmas lanceolate, sharp tipped

13. H. landsti^iua

2\' . Stigmas capitate rotund to linear
oblong, blunt tipped.
22. Leaf blades rubescent, the
pigment extractable on boil-
ing; flowers subsessile

24. //. rubripi^menta

22'. Leaf blades not rubescent,
not yielding pigment on boil-
ing; flowers obviously pedi-
cellate.

23. Leaves chartaceous to
subcoriaceous, not mem-
branous and translucent
on drying.

24, Leaves widely ob-
long to ovate sub-
rotund, to 1 1 cm
long, subcoriaceous,
the lateral veins 7-9;
inflorescences com-
pressed, the flow-
ers few

6. H. cercidifolia

24'. Leaves elliptic or
obovate elliptic, 9-
35 cm long, char-
taceous, the lateral
veins (I I-)15-20;
inflorescences usu-
ally spreading, the
flowers numerous

20. //. pitticri

23'. Leaves membranous and
translucent on drying.
25. Leaves to 25 cm

long; pedicels 0.5

19801 DVVYFR— { LORA 01- PANAMA (Family 179. Ruhiaccae) 225

mm wide; calycine
lobes to 1.1 mm
long; corolla tube

pubescent: sta-
mens 3(^)

19. H. ostaurea

25'. Leaves 10-16 cm
long; pedicels ca.
0.25 mm wide; ca-
lycine lobes 0.4-
0.8 mm long; corol-
la tube glabrous;
stamens 4

5. //. capilhicca

18'. Anther connective produced apically or basally.
26. Leaves to 9.5 cm wide; anther connective

produced basally; flowers purple

9. H. fimhrianthcra

26'. Leaves usually 3 cm wide; anther connective

produced apically; flowers yellow

28. //. woocLsonii

6'. Leaves sessile.

27. Inflorescences with peduncles 3-6 cm long.

28. Leaves rounded toward the apex; calycine lobes 3-4 mm long

11. //. haxdcnii

_^ ^ ^ ^ ^ ^ *-h -^ -^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ -— — ^ .mm. ^ ^ -^ .- ■^- — ■- ^ ^ **- BF

28'. Leaves acute or acuminate toward the apex; calycine lobes less than I

mm long.

29. Inflorescences solitary in axils; plants prostrate .. 25. //. suhaiiricuUila

29'. Inflorescences several per axil or cauliflorous; plants upright

2. //. araneopcdarhi

27'. Inflorescences with peduncles ca. 30 cm long 17. H. inaiuissatani

1. Hoffmannia aeruginosa Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18:
1313. 1938. type: Costa Rica, Austin Smith A86 (F).

Shrubs to 4 m tall, the branchlets slender, terete, the bark pale brown, fer-
rugineous villosulose, the hairs deciduous leaving a minutely tuberculate surface.
Leaves oblong, to 20 cm long, to 8 cm wide, short acuminate at the apex, widely
to narrowly cuneate at the base, the costa subplane and slender above, promi-
nulous beneath, the lateral veins ca. 15, arcuate, prominulous, the intervenal
areas often appearing lineolate under magnification, stiffly chartaceous, scarcely
discolorous, drying green or brown, glandular shiny, glabrescent, minutely foveo-
late, the raphides moderately abundant, white, divergent, to 0.5 mm long, more
conspicuous beneath, the costa ferrugineous pubescent, sometimes glabrescent;
petioles 0.5-0.8 cm long, curved or reflexed, densely ferrugineous villosulose, to
3 mm wide; stipules not seen. Injloreseences in the uppermost axils, subsessile
glomerules of 3-10 flowers, the reduced peduncles ferrugineous pubescent or
purple villose. Flowers sessile, with the hypanthium to 1.7 mm long, villose with
elongate, soft and ferrugineous hairs; calycine teeth subulate, to 1.5 mm long;
corolla densely villose outside, the lobes acute; anthers ca. 3 mm long, acute at
the apex, the connective not produced at the apex or base; stigmas 2, rotund or
clavate, the ovary 2-loculate. Fruit not seen.

The species occurs in Costa Rica and Panama. The collection of Austin Smith
(H659) from Guadalupe de Zarcero, Costa Rica, has the notes on the label:
''Growing in deep forest shade; gregarious but very local. Base of the stem dull.

226 ANNALS OF THE MISSOURI BOTANICAL GARDF.N [Vol. 67

dark brown. Buds deep chokeberry red. Petioles and some of the veins of the
under surface of the leaves tinged with pinkish red/'

BocAS DEL TORo: Chinqui Lagoon, We del 2289 (MO).

2. HofTmannia araneopedaria Dwyer.^^ type: Panama, Gentry & Mori 14166
(MO, holotype).

Shrubs to 2 m tall, stem simple, lignose, terete, to 1.8 cm in diam.; bark with

inconspicuous longitudinal lines, glabrous; nodal scars ringlike, the nodes well

spaced. Leaves sessile, oblong, 12.5-32 cm long, 6-9 cm wide, short acuminate,

obtuse or widely deltoid toward the apex, the acumen 1-1.2 cm long, basally

attenuate and elongate cuneate below the middle, ultimately 8-10 mm wide and

truncate or scarcely auriculate, the costa scarcely prominulous to immersed

above, prominulous beneath, the costa slightly prominulous above, prominulous

on the lower side only near the apex, prominent at the base, 2.5 mm wide, the

lateral veins arcuate, the undulate submarginal veins inconspicuous, either absent

or to 0.5 cm from the margin, the intervenal areas reticulate, stiffly papyraceous,

concolorous, drying brown, glabrous above, minutely puberulent on the costa

and veins beneath. Inflorescences cauliflorous or axillary; peduncles 8-12, 0.8-

3 cm long, wiry, unequal, radially disposed from a single axil, arising from a

compressed hemispherical swollen pad, ca. 1 cm wide, 0.5 cm long, the flowers

to 6 on each axis, secundly disposed. Flowers (in bud) with pedicels red when

living, 5-7 mm long, the hypanthium turbinate, 1 mm long, glabrous, ca. 8-ribbed,

the calycine cup ca. 0.2 mm long, subturgid, the 4 lobes widely triangular, ca.

0.3 mm long, glabrous; corolla yellow, the tube carnose, glabrous, the 4 lobes

triangular oblong, 2.8 mm long; anthers subbasifixed, the filaments short; ovarian

disc cushionlike, narrower toward the apex, ca. 0.6 mm long, the style slender,

the 2 ovate stigmas to 0.8 mm long. Fruits (perhaps immature), turbinate or

oblong rotund, ca. 0.6 cm long, truncate, glabrous, not ribbed, white in living
state.

Hofffuannia araneopedaria is known only from Panama. It is most closely
related to //. suhauriculata Standley which has similarly shaped, sessile leaf
blades. The specific name araneopedaria means ''resembling a spider's legs'' a
reference to the several wiry peduncles suggesting the legs of a ''Daddy long-
legs." Some inflorescences are cauliflorous, an uncommon condition in Hoff-
niannia.

daru'n: Cerro Mali on Colombian border, S slope just below summit, lower montane wet forest-
prcmontane wet forest transition, 1400 m, Gentrx & Mori 14166 (MO).

^^ HoJJniannia arancopeJana Dwyer, spec. nov. Frutices ad 2 m alti. caule simplici glabro. folia
sessilia oblonga, 12.5-32 cm longa, 6-9 cm lata, versus apicem obtusa vel lato-deltoidea, venis late-
ralibus ca. 20. rigido-papyracea supra glabra subtus minuto-puherula. Inflorescentiae pedunculis 8-
12 ab una axilla radiatis et ab area tumida compresso-haemosphaerica, ca. 1 cm lata. ca. 0.5 cm alta,
floribus in axe utroque ad 6 secundatim dispositis. Flares pedicellis, cupula calycis ca. 0. 1 mm longa,
dentibus 4 lato-triangularibus, ca. 0.3 mm longis, glabris; corolla lutea tubo glabro lobis 4 oblongo-
triangularibus, ca. 2.8 mm longis. Fructus his fortasse juveniles turbinati vel oblongo-rotundi, ca. 0.6
cm longi, truncati glabri non costati, in vivo albi.

19801 DWYHR— FLORA OF PANAMA (Family 179. Ruhiacectc) 227

3. Hoffmannia areolata Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 52.
1940. type: Panama, Davidson 40 (F, holotype; isotype, MO).

Herbs to 2 m tall, the stems smooth, glabrous or villose. Leaves elliptic, to
25 cm long, to 8 cm wide, attenuate acuminate at the apex, the acumen to 3 cm
long, cuneate at the base, the costa immersed above, piano compressed beneath,
the lateral veins ca. 25, arcuate, glabrescent above, appressed villosulose on the
costa beneath, chartaceous, concolorous, the raphides numerous, scattered,
white, linear, to 0.3 mm long; petioles 1-4 cm long: stipules not seen. Inflores-
cences sessile, glomerules in the uppermost axils, to 2 cm long or longer, often
wider than long. Flowers yellow or greenish, the hypanthium oblong, covered
with elongate ferrugineous hairs to 1 .6 mm long, the calycine lobes 4, ca. 3.5 mm
long, obtuse; corolla ca. 4.5 mm long, the tube cylindrical, ca. 1 mm wide, the
lobes shorter than the tube; stamens 4, ovate, ca. 1.5 mm long, the connective
not produced at the apex or the base; stigmas 2, oblong, spreading, ca. 1.5 mm
long, the style ca. 2.5 mm long. Fruits scarlet (fide Davidson).

Hoffmannia areolata is known only from Panama.

CHiRiQUi: Boquete, near Bajo Chorro. Davidson 40 (F, GH, MO): Woodson & Schery 661 (F,
GH, MO).

4. Hoffmannia bullata L. O. Williams, Fieldiana: Botany 36: 52. 1973. tyf'f:
Guatemala, Steyermark 38719 (F, holotype).

Suhshruhs to 0.3 m tall, the stems simple, the lowermost nodes rhizophorous,
the hairs dense, ferrugineous, the lenticels many, the nodes moderately well
spaced. Leaves narrowly obovate oblong, 5-15 cm long, 1.5-5.0 cm wide, often
slightly falcate, widely deltoid or rounded at the apex, the acumen inconspicuous,
obtuse or cuneate toward the base, often abruptly contracted toward the base,
the costa plane above or immersed, prominulous beneath, the lateral veins 8-14,
arcuate, conspicuous and prominulous beneath, chartaceous, usually bullate at
least on drying, discolorous, drying tan or silvery, glabrous above, the costa and
veins below with golden, dense hairs, the raphides minute, white; petioles ca. 1.3
cm long, pubescent; stipules not seen. Inflorescences axillary; peduncles slender,
to 2.5 cm long, ca. 0.7 mm wide. Flowers with hypanthium oblong or turbinate,
glabrous, lightly costate, often seemingly alate on pressing, the teeth 4, to 1.5

(W

4 mm long, the

hypanthium turbinate, glabrous, the calyx lobes triangular or lance triangular,
acute, sparsely villosulose: corolla about 1 cm long, glabrous, the lobes oblong,
obtuse or acute, 2-3 times as long as the tube; stamens with short filaments, the
anthers exserted." Fruits red; pedicels to 2 mm long, oblong or turbinate, 6-7
mm long, truncate at the apex, 4-angled, smooth, glabrous, lightly costate, the
raphides abundant, the persistent calyx lobes obtuse.

Hoffmannia hullata is known from Mexico to Panama. Unfortunately the
single Panamanian collection cited below lacks flowers so the floral description
above is largely from Louis Williams' original diagnosis. Steyermark 37241 (F),
from the slopes of Volcan Tajumuico, Guatemala, 1,300-1,500 m, notes that the
flowers have: "calyx pale green; corolla lobes spreading, salmon-brick color;
anthers pale yellowish white, as in stigma."

228 ANNALS OF THF MISSOURI BOTANICAI GARDHN |Vol. Ul

Hoffnuuinia hullata is closely related to //. pusulata Dwyer, which has more

numerous lateral veins and has been collected at a much lower elevation, less
than 200 m.

Panama: Cerro Campana, 800 m, Croat 12137 (MO).

5. HofTmannia capillacea Dwyer, Ann. Missouri Bot. Card. 56: 274. 1969. type::
Panama, Elias 1783 (MO, holotype).

Suhshruhs. the stems lignose, smooth, glabrous, the petiole scars somewhat
conspicuous, the nodes ca. 1 cm apart. Leaves oblong, 10-16 cm long, 5-7 cm
wide, subacute and short acuminate at the apex, the acumen to 1 cm long, acute
at the base and scarcely symmetrical, the costa subplane on both sides, the lateral
veins ca. 10, subprominulous, the intervenal areas prominulous and spreading,
membranous, discolorous, glabrous; petioles to 5 cm long, ca. 0.2 cm wide, lightly
alate, strict; stipules not seen. Inflorescences cymes in the uppermost axils, gla-
brous; peduncles 1-15 mm long, ca. 0.3 mm wide, capillaceous; flowers 2-4 on
spreading branches, often somewhat radially disposed. Flowers (in bud) with the
pedicels 6-8 mm apart, capillaceous, to 0.4 mm wide, glabrous; hypanthium ca.
3 mm long, squarish, glabrous, carnose, the lobes 4, acute, glabrous; stamens
4, the anthers 2.5-3 mm long, the connective not produced at the apex or base,
the filaments short, ca. 0.7 mm wide; stigmas 2, oblong rotund, ca. 1.5 mm long,
wider than the style. Fruits not seen.

Hoffnuuinia capillacea is known only from Panama,

cocI-E: Alto Calvario. 7 km N of El Cope, Folsom 3314 (MO), san bi as: Headwaters of Ri'o
Mulatupo, Filias 1783 (MO).

6. Hoffmannia cercidifolia Dwyer, Ann. Missouri Bot. Card. 56; 275. 1969.
TYPHI Panama, Duke 15597 (MO).

Shrubs to Wi m, the branchlets slender, smooth, glabrous. Leaves ovate
subrotund or widely oblong, 7-1 1 cm long, 5.5-8 cm wide, short acuminate and
deltoid apically, the acumen falcate or straight, to 1 cm long, ultimately subacute,
basally obtuse or truncate, the costa subplane above with a thin median ridge,
subplane beneath, the secondary veins 7-9, arcuate, prominulous beneath, dis-
colorous, dark red brown and glabrescent above, beneath glabrous or golden
puberulent on the costa and veins, glistening, above honeycomb-like under mag-
nification; petioles slender, to 4.5 cm long, ca. 0.15 cm wide; stipules not seen.
Inflorescences to 1 cm long, the flowers few, nodding, the peduncle reduced.
Flowers (buds only) with the pedicels short, the hypanthium short, campanulate,
ribbed, glabrous, the calycine cup short and scarcely measurable, the 4 triangular
teeth ca, 1 mm long, carnose, glabrous except puberulent marginally; corolla
white with the tube short, ca. 2.5 mm long, subcarnose, pubescent outside, gla-
brous within, the lobes 4, oblong, ca. 4 mm long; stamens 4, the anthers ca. 3.5
mm long, ca. 0.8 mm diam., the connective not produced at the apex or base,
the filaments compressed; style ca. 5.5 mm long, the 2 stigmas clavate capitate.
Fruits not seen.

The species is known only from Panama.

19801 DWYF.R— FLORA OF PANAMA (Family 179. Ruhiaceae) 229

darikn: Cloud Forest, Cerro Campamento, S of Cerro Pirre, Duke 15597 (MO), vfraguas: 16
km NW of Santa Fe, road to Calovebora, 300-500 m, Mori & KuUunki 6119 (MO).

7. Hoffmannia davidsoniae Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22:
53. 1946. typf: Panama, Davidson 216 (F, holotype; isotype, MO).

Herbs to 1.5 m tall, the stems terete, smooth, glabrous, usually drying tan,
the internodes to 9 cm long. Leaves oblong rotund, 8-17 cm long, 4-12 cm wide,
rounded and acuminate at the apex, the acumen 0.5-1.8 cm long, basally rounded
or tapering widely, abruptly cuneate, the costa prominulous to subplane above,
prominent beneath, the lateral veins 12-15, widely arcuate, prominulous, 0.5-1.2
cm apart, thinly and stiffly succulent, lustrous, brown, white puberulent to gla-
brous above, minutely puberulent on veins beneath; petioles 2-7 cm long; stipules
not seen. Inflorescences cymose, mostly subtended by roots, the peduncles elon-
gate, strongly ascending, 5-10 cm long, 0.1-0.2 cm wide, rubescent, glabrous,
terminated by 3-4, usually radially disposed and spreading branches, the cymules
ascending, each branch 7-17-flowered. Flowers with the pedicels stiff, glabrous,
to 5 mm long, ca. 0.3 mm wide; buds obviously acute at the apex; hypanthium
strongly 6-8-ribbed, emerald green to red in living state, to 3 mm long, the ca-
lycine lobes flaring, deltoid, 0.3-0.5 mm long; corolla to 6 mm long, the lobes 4,
much longer than the tube, glabrous; anthers 4, oblong, to 3.8 mm long, the
connective not produced at apex or base, the filaments to 0.6 mm long; ovary
2-loculate, the style 4 mm long, the stigmas 2, clavate oblong, ca. 1.1 mm wide,
each somewhat uncinate. Fruits oblong to 3.5 mm long, truncate, the persistent
calycine teeth scarcely measurable, glabrous, delicately costate.

Hoffmannia davidsoniae is known only from Panama.

My own collection notes on this species {Dwyer 8789) are more elaborate than
usual: ''To 1 m tall; leaves smooth, lustrous, brown, stiffly succulent; inflores-
cence 17-flowered, glabrous, the peduncle thick, red green; hypanthium strongly
angular, green; ovarian disc cushionlike; corolla rose, the tube glabrous, in bud
flushed with red or pink, the anthers 4, the stigmas 2, clawlike, angular."

CHiRiQUi: Cerro Horqueta, 7000 ft, Blum & Dwxcr 2623 (MO). Rm Chiquero, irArcy 10983
(MO). Bajo Chorro, 6000 ft, Davidson 216 (F, MO). Cerro Horqueta, Dnyer & LaUathin 8789 (MO).
Bajo Mona and CJuebrada Chiquero, 1500 m, Woodson & Sclwrv 595 (MO). COCLE: Cero Pilon, 2900
ft, LaUaflun 133, 135 (both MO).

8. Hoffmannia eliasii Dwyer, Ann. Missouri Bot. Card. 56: 276. 1969. type: Pan-
ama, Kirkhride & Duke 1129 (MO).

Herbs to 10 cm high, succulent, the stems simple, horizontal at the base,
lignose, 15-20 cm long, to 1.2 cm wide; roots fibrous, lignose. Leaves oblong or
obovate elliptic, to 25 cm long, 9.5 cm wide, deltoid or subobtuse at the apex,
scarcely acuminate, cuneate to narrowly cuneate at base, the costa prominulous
above to subplane beneath, often lightly alate, the secondary veins 12-16, slightly
arcuate, prominulous, to 1.8 cm apart, the intermediate veins irregular and soon
branching, then evanescent, the ultimate venules spreading pinnatiform, the mar-
ginal vein conspicuous, ca. 7 mm from the margin, stiffly chartaceous, scarcely
discolorous, glabrescent, the hairs on the costa and the veins few; subsessile or
petiolate, the petioles to 12 cm long, often lightly alate; stipules not seen. Inflo-
rescences 1-3 cm long, to 2 cm wide, subtended by roots, the peduncles 2 per

230 ANNALS OF THE MISSOURI BOTANICAL GARDEN |Vol. 67

axil, often stiff, to 4.5 cm long, often marcescent, canaliculate, pilose, the hairs
weak and spreading. Flowers sessile or on pedicels to 4 mm long; hypanthium
campanulate, ca. 2 mm long, glabrous, black when dry, ribbed, the calycine teeth
4, somewhat unequal, erect, subulate, 1-1,8 mm long, minutely puberulent; co-
rolla glabrous, the tube cylindrical to 2.5 mm long, the lobes 4, lanceolate, longer
than the tube; stamens 3, the anthers 3-4 mm long, subsessile, the filaments ca.
0.2 mm long, the style narrow cylindrical, the connectives not produced at the
base or the apex; stigmas 2, unequal, oblong, to 0.7 mm long, scarcely wider than
the style. Fruit (immature) with pedicels strongly ascending, to 4 mm long, dif-
fusely pilose, oblong, to 5 mm long, to 2.5 mm wide, the calyx persistent, the
lobes 1-2.5 mm long, red when dry, moderately ribbed, minutely puberulent.

The species is known only from Panama and nearby Colombia.

DARir.N: Cana-Cuasf Trail, Chepigana, 4000 ft, Terry & Terry 1508 (F, GH, MO). Rio Cuasf
mainstream. 2,5 mi S of Tres Bocas, Klrkbr'ulc & Duke 1129 (MO), san blas: Headwaters Rio
Mulalupo, Ellas 1779 (MO). Mountains above Puerto Obaldia, Gentry 1504 (MO). Colombia: Cmoco:
Rfo Ncrcua, Quebrada Barrial and Quebrada Ambrosio near Camp Curiche, Duke 1 1438 (MO).

9. Hoffmannia fimbrianthera Dwyer, Ann. Missouri Bot. Gard. 56: 277. 1969.
TYPi:: Panama, Kirkhridc & Duke 556 (MO, holotype).

Siibshrtihs. the stems smooth, reed-like, glabrous, terete, the internodes to 15
cm long. Leaves oblong rotund, often subfalcate, to 19 cm long, to 9.5 cm wide,
deltoid at the apex, short acuminate, the acumen to 1.5 cm long, basally acute
or narrowly cuneate at the base, the costa plane above, subprominent beneath,
the lateral veins ca. 15, divergent-arcuate, conspicuously prominulous beneath,
the veinlets spreading pinnatiform, the submarginal vein ca, 8 mm from the mar-
gin, papyraceous, scarcely discolorous, drying grey, conspicuously honeycomb-
like under magnification, glabrescent above, granulose puberulent only on the
veins beneath; petioles slender, lax, to 7 cm long, to 1.2 mm wide, minutely
webby puberulent; stipules not seen. Inflorescences sessile, the flowers few,
(3-)5(-10). Flowers subsessile or on pedicels to 1 mm long; hypanthium rotund
oblong, ca. 3 mm long, somewhat smooth, stiffly carnose, ferrugineous puberu-
lent, the calycine lobes 4, spreading uncinate subulate, to 1 mm long, puberulent;
corolla carnose, ferrugineous puberulent outside, minutely farinose within, the
lobes 4 (?); stamens 4, the anthers oblong, to 3.2 mm long, minutely apiculate,
the connective petaloid fimbriate at the base for 0.3 mm long; style to 3.5 mm
long, 0.2 mm wide, wider toward the apex, the stigmas 2, clavate, ca. 0.35 mm
wide. Fruits not seen.

Hoffmannia finihrianthera is known only from Panama.

BOCAS Din TORo: Rio Teribe, just below Puerto Palenque, 350 m, KIrkhride &. Duke 55f) (MO).

10. Hoffmannia gentryi Dwyer. *^** typh: Panama, Gentry & Mori 14061 (MO,
holotype).

Suhshrubs to 1.5 m tall, the branchlets weak, angular when dry, villose, the
hairs persistent or deciduous, curled, swollen at the base, the nodes moderately

^'^^ Hojfniannla ^entryl Dwyer, spec. nov. Suffrifiices ad 1.5 m aiti, ramulis debilibus. Folia
oblonga vel obovato-oblonga, 12-20 cm longa, 5-10 cm lata, plerumque versus apicem delloidea.

1980] DWYHR— r-lX)RA OF PANAMA (Family 179. Ruhiaceae) 231

well spaced. Leaves oblong or ovate oblong, 12-20 cm long, 5-10 cm wide,
deltoid toward the apex, occasionally acute, acute or obtuse at the base, rarely
subtruncate, the costa prominulous on both sides, the lateral veins ca. 10, arcuate
and scarcely prominulous, the intervenal areas lightly reticulate, the undulate
submarginal vein to 5 mm from the margin, membranous, scarcely discolorous,
hairy on both sides, the hairs on upper surface few, subulate, spreading or ap-
pressed, mostly confined to the costa and the veins, below more dense, often
curled and giving a webby effect; petioles weak, slender, often curved, to 4.5 cm
long, densely hairy; stipules deciduous. Inflorescences axillary, to 2.5 cm long;
peduncle slender, to 1.5 cm long, ca. 0.3 mm wide, the cymes 4-5-flowered, the
flowers unilaterally and helicoidally disposed. Flowers subsessile, densely villose,
the hypanthium turbinate, to 2.5 mm long, the lobes 4, narrowly oblong; corolla
light green, subcampanulate, the tube about as long as the calycine teeth, perhaps
glabrous outside, raphides abundant, the lobes 4, longer than the tube; stamens
4, erect, the anthers oblong, ca. 4 mm long, subsessile, attached near the mouth;
style slender, exceeding the anthers, the stigmas connate, clavate, ca. 1.5 mm
long. Fruits not seen.

Hoffmannia gentryi is named in honor of one of the co-collectors of the type,
a distinguished student of the flora of Panama. It is known only from Panama,
although it no doubt occurs in adjacent Colombia. It is distinguished by its mem-
branous leaves adorned with adpressed and often curled hairs; the hypanthium
is replete with elongate hairs and the calycine cup has teeth as long as the corolla
tube. The flowers being unilaterally and helicoidally disposed distinguishes it from
//. aerohita to which it seems closely related.

darif.n: Cerro Tacarcuna, summit camp, along stream N of camp, 1550-1650 m. Gentry & Mori
I406I (MO).

11. Hoffmannia haydenii Dwyer, Ann. Missouri Bot. Gard. 56: 277. 1969. type:
Panama, Hayden 177 (MO, holotype).

Herbs, the stems drying quadrangular. Leaves sessile, obovate elliptic or
oblong, to 35 cm long, to 14 cm wide, rounded toward the apex, short acuminate,
the acumen to 1.5 cm long, basally narrow cuneate and ultimately auriculate, the
costa prominulous, slender and ridgelike above, beneath subplane, to 2.5 mm
wide proximally, puberulent, the lateral veins ca. 20, somewhat plane above,
prominulous beneath, coriaceous, discolorous on drying, glabrescent and brown
above, puberulent on costa and veins beneath; stipules not seen. Inflorescences
1,5_7 cm long, the peduncles 1.2-6 cm long, subtended by roots, capillaceous,
ca. 0.3 mm wide, the branches strict, puberulent, red in living state, to 2.5 mm
long, puberulent. Flowers with buds cuneate and acute at the apex; hypanthium
puberulent, scarcely ribbed, the calycine lobes 4, erect lanceolate, 3-4 mm long,
purple red when dry, puberulent; corolla lobes 4, ca. 5 mm long, puberulent
outside, longer than the tube; stamens 4, the anthers ca. 4 mm long, the connec-

venis lateralibus ca. 10, membranacea utrinque ciliata praesertim subtus; petiolis debilibus, ad 4,5 cm
longis. Inflorcscentlac ad 2.5 cm longae. pedunculo gracilissimo, cymis floribus 4-5 unilateraliter
dispositis! Florcs hypanthio dense elongato-ciliato dentibus 4 ad 3 mm longis; corolla tubo brevi lobis
ad 4 mm longis: staminibus subscssilibus, antheris 4 oblongis, ca. 4 mm longis, exsertis. Fructus non

VISL

232 ANNALS or THC MISSOURI BOTANICAL GARDI-N [Vol. 67

tive not produced at the apex or the base, the filaments short: stigmas 2. aggre-
gated. Fruits not seen.

Hofjmaniuii haydenii is known only from Panama.

CHiRigui: Finca Collins, ca. 5000 ft, Haydcn 117 (MO).

12. Hoffmannia kirkbridei Dwyer, Ann. Missouri Bot. Gard. 56: 278. 1969.
typl: Panama, Kirkhiidc & Duke 429 (MO, holotype).

Suhshruhs to 2 m tall, the stems glabrous, drying minutely rugulose. Leaves
elliptic oblong to 40 cm long, to 20 cm wide when dry, the costa prominulous
and canaliculate above, subprominent beneath, distinctly corky alate toward the
base, the lateral veins 15-25, ca. 3 cm apart, arcuate, scarcely prominulous above,
conspicuously prominulous beneath, the intermediate veins more divergent, soon
evanescent, the veinlets spreading, irregularly pinnatiform with patches of simple
appressed hairs on the costa and veins beneath, the hairs few, stellate, stiffly
chartaceous, discolorous when dry; petioles to 13 cm long, ca. 3 mm wide in the
middle, inconspicuously swollen below the base of the blade, the vescicles 0.6-
10 cm long, ca. 1 cm wide when dry. Inflorescences often 3 per axil, the peduncles
reflexed, 1-1.5 cm long, ca. 0.13 cm wide, with minute patches of crystalline
hairs and raphides, the hairs few, to 0.7 mm long, rarely stellate. Flowers (juvenile
fruit) subsessile; hypanthium short, moderately villose, the raphides thick, white;
calycine cup somewhat turgid, villose, the lobes 4, thickly carnose to 4.5 mm
long, to 3.5 cm wide, ciliate, the veins few, ascending, the wall of the hypanthium
thick. Fruits (immature) rhomboid, ca. 6 mm in diam.; seeds thickly reticulate.

Hoffmannia kirkbridei is known only from Panama.

BocAS DKi TORo: Biiena Vista, Aimirante, Cooper 225 (F). Quebrada Huron, ca. 300-400 ft,
Kirkhiidc & Duke 429 (MO).

13. Hoffmannia lancistigma Dwyer, Ann. Missouri Bot. Gard. 56: 279. 1969.
type: Panama, D^\yer d- Hayden 7735 (MO, holotype).

Shrubs, the stems canelike, terete, smooth, the hairs minute, dense, ferru-
gineous, moniliform, deciduous, the nodes 2.5-3.5 cm apart. Leaves strongly
ascending, well spaced, oblong, to 19 cm long, to 9 cm wide, obtuse at the apex,
scarcely short acuminate, ultimately obtuse, cuneate at the base, often asym-
metrical, the costa prominulous above and beneath to 0.15 cm wide, the lateral
veins ca. 15, arcuate, prominulous beneath, the intermediate veins spreading
pinnatiform, papyraceous or stiffly chartaceous, minutely granulose and honey-
comb-like under magnification, the hairs few beneath, except on the densely
pubescent costa; petioles to 7.5 cm long, ferrugineous; stipules not seen. Inflo-
rescences usually 1 per axil, often arising from a leafless part of the twig, the
peduncle ascending, to 10 cm long, slender, ca. 2 mm wide, strict or arcuate,
glabrescent or with few crystalline hairs. Flowers glabrous (?); hypanthium ob-
long, to 3.8 mm long, ribbed, the calycine teeth 4, erect, triangular, ca. 1 mm
long; corolla tube thin carnose, the lobes 4(-6), ca. 4.5 mm long, carnose; stamens
4, the anthers ca. 3.8 mm long, to 1.2 mm wide, the connective not produced at

1980] DWYF.R— FLORA OF PANAMA (Family \79. Riihiaccac) 233

apex or base, the filaments ca. 1 mm long, flat, to 1 mm wide; style ca. 4 mm
long, the stigmas 2, lanceolate, ca. 2.8 mm long, to 0.8 mm wide, thick, wider
than the style, constricted medially. Fruits not seen.

Hoffiminnia lancistigma is known only from Panama.

CHiRiQuf: Cqvvo Horqiieta. ca. 6000 ft. Dwyer & Hayden 7735 (MO. SCZ. UC).

14.

M

Bottcri 974; Bourgeon 2992; Linden /, none seen.

Shrubs to 1.5 m, the branchlets smooth, minutely puberulent to glabrescent,
the nodes well spaced, the pith porate. Leaves elliptic to oblong elliptic, 8-15 cm
long, 2.5-5 cm wide, widely deltoid or rounded at the apex but distinctly short
acuminate, the acumen to 1.5 cm long, slender, often falcate, attenuate acute
toward the base, slightly decurrent along the petiole, the lateral veins 14-20,
arcuate, plane and somewhat inconspicuous above, prominulous beneath, not
forming a distinct marginal vein, the veinlets inconspicuous, rigidly papyraceous,
scarcely discolorous, presumably glabrous above, glabrescent to minutely pu-
berulent on the midvein and the lateral veins, often the upper surface with small
patches of white round crystals; petioles 0.5-2.5 cm long; stipules not seen.
Inflorescences axillary, sessile or on reduced peduncles, the cymes few flowered.
Fhnvers not seen. Fruits somewhat congested, puberulent, to 4 cm long, the
peduncle absent or to 0.5 cm long; pedicels to 2 mm long, the pericarp oblong,
to 4 mm long, truncate, the lobes 4, erect triangular or oblong, to 1 mm long,
acute to obtuse, puberulent, concave within, rigid, the persistent ovarian disc
compressed annular, the fruit body with 8 narrow but prominent ribs.

Hoffmannia lenticellata extends from Mexico to Panama.

Many collections labelled H

Middle

crystals on the leaf blades as a distinguishing character needs assessment. Stand-
ley in his key to species of Hoffmannia in North American Flora used the char-
acter that the corolla lobes are 2 times as long as the tube; no flowers were
available on the single collection from Panama seen.

CHiRiQL'f: Road Between Cerro Piinta and Las Nubes. 1800-2100 m. Croat 26399 (MO).

15. Hoffmannia lewisiana Dwyer, Ann. Missouri Bot. Card. 56: 279. 1969. type:
Panama, Ridguay & Solis 2413 (MO).

Shrubs, the branchlets smooth, glabrous, canaliculate. Leaves oblong, 17-19
cm long, 7-8 cm wide, somewhat attenuate acute and acuminate at the apex, the
acumen to 1.5 cm long, acute and almost symmetrical at the base, the costa
subplane above, canaliculate, prominulous beneath, to 0.15 cm wide, the lateral
veins ca. 12, plane above, prominulous beneath, arcuate, ultimately spreading
reticulate, papyraceous, concolorous, glabrous above, pubescent beneath only
on the costa and veins, petioles to 4.5 cm long, glabrous; stipules not seen.
Inflorescences to 6.5 cm long, spreading, the peduncles short, the branches nu-
merous, 2.4 cm long, 0.8-1.8 mm wide, the cymes few, alternate or often sub-

234 ANNALS OF IHE MISSOURI BOTANICAL GARDEN (Vol. 67

radiate. Flowers with the buds clavate; pedicels 1-7 mm long, ca. 0.4 mm wide,
densely white puberulent; hypanthium oblong, to 7 mm long, carnose, puberulent
and conspicuously ribbed, the teeth 4-6, erect, deltoid, unequal, often one con-
spicuously reduced, to 2.5 mm long, puberulent; corolla with the tube to 11 mm
long, carnose, the lobes 5, to 5.5 mm long, acute, puberulent outside; stamens
4-5, the anthers ca. 3.5 mm long, ca. 1.3 mm wide, the connective not produced
at the apex or base, the filaments compressed, ca. 1 mm wide; style 12-14 mm
long, the stigmas (2-)3-4, capitate. Fruits not seen.

Hoffmannia lenlsiana is known only from Panama.

CHiRiQuf: Cerro Punta, 6800 ft, cloud forest and distrubed edge. Rid^way & Soli.s 2413 (MO).

16. Hoffmannia longicalycina Dwyer, Ann. Missouri Bot. Gard. 56: 280. 1969.
type: Panama, Kirkbride & Duke 1278 (MO, holotype).

Suhshrubs, the branchlets smooth, glabrous, the nodes ca. 1 cm apart. Leaves
oblong, tapering cuneately toward the apex, acuminate, the acumen short, to 1
cm long, ultimately acute, narrowly cuneate at the base, the costa prominulous
and ridged beneath, to 2 mm wide, the lateral veins ca. 16, spreading arcuate,
prominulous, rigidly chartaceous, slightly discolorous, green when dry, glabres-
cent above, appressed ciliate on the costa and the veins beneath, the hairs often
rigid and appressed falcate; petioles to 10 cm long, angular to compressed, ca.
0.2 cm wide, the hairs white, thickly subulate, forming a thin crust; stipules not
seen. Flowers not seen. F/7//7.V oblong, terete in cross section, to 12 mm long,
ca. 7 mm wide, delicately costate, pubescent with weak ferrugineous hairs to 2
mm long: persistent lobes of calyx 5, erect, linear lanceolate, to 12 mm long, ca.
2 mm wide, petaloid and pubescent; seeds numerous, thickly reticulate, ca. 0.5
mm in diam,, visible through the thin wall.

Hofjnuuuua longiealycina is known only from Panama.

darirn: Cana-Cuasf Trail between Cerro Campamento and La Escalera to '^paramo," E of Tres
Bocas, Kirkbride & Duke 1278 (MO).

17. Hoffmannia manussatani Dwyer, '^' type: Panama, Allen 4978 (MO, holo-
type). — Fig. 50.

Herb to 3 ft tall, the stem perhaps unbranched, 4-angled, smooth, glabrous,
the nodes well spaced. Leaves sessile, obovate oblong or spatulate, ca. 20 cm
long, 6.0-6.5 cm wide, narrowly acuminate at the apex, the acumen to 1.5 cm
long, acute, the blade obviously contracted below the middle, at base vaguely
auriculate, to 2.5 cm wide basally, the costa prominulous above with a central

**^ Hoffmannia manussatani Dwyer, spec. nov. Herhae caulibus 4-angiilarihiJS glahris. F<flia
sessilia obovato-oblonga vel spathulata, ca. 20 cm longa, 6.0-6.5 cm lata, acuminata, basi infra me-
dium altenualo-spathulata proximalitcr modice auriculata venis lateralibus ca. 18 chartacea glabra.
Infl()re.\( enfiae axillares pedunculo elongato ad 30 cm longo, floribus tcrminalibus in cacspitibus pauci-
floris congestis ad 2.5 cm latis, jugis tribus bractearum bene distanlibus in pedunculo, bracteis
magnis plerumque ovato-oblongis, ad 9 mm longis. Flares hypanthio ferrugineo-villosuloso, lobis
calycis 4, ca. 3 mm longis: corolla lobis 4. ad 4 mm longis; staminibus sessilibus antheris 4, ad 4 mm
longis. Fructus non visi.

1 980J

DWYER— FLORA OF PANAMA (Family 179. Ruhiaccac)

235

Figure 50. Hojfmannia manussataiu Dwyer.— A. Habit (x>/2).—B. Flower bud { x5).— C. Sta
mens (x5).— D. Style (x5).— E. Ovary cross section (x5). [After Allen 4978.\

depression, prominulous beneath, ca. 0.2 mm wide proximally, the lateral veins
ca. 18, arcuate, the submarginal vein inconspicuous and proximate to the margin,
the veinlets pinnatiform, chartaceous, concolorous, drying reddish, glabrous;
stipules widely triangular, to 0.6 cm long, wider than long, thickly chartaceous,
glabrous. Inflorescences axillary, elongate pedunculate, the peduncles fiagelli-

236 ANNALS OF- THl; MISSOLIRI BOIANICAI. GARDEN [Vol. 67

form, to 30 cm long, twisted, angular, puberulent, 1.6-2.3 mm wide basally, with
4 pairs of bracts, the basal bracts much reduced, rotund, to 3.5 mm long, the
other 2 pair well spaced, one pair at the apex of the lower Vs of the peduncle, the
other at the base of the upper 'Z^, rotund or ovate rotund, 1.6-2.5 cm long, obtuse
or rounded at the apex, stiffly chartaceous, ascending, the costa and veins evi-
dent, disposed as in the leaves, the flowers few, loosely aggregated in a terminal
rotund or compressed rotund head, to 2.5 cm in diameter, presumably of few
cymules; bracts oblong or ovate oblong, 7-9 mm long, stiff, ciliolate marginally
and on the keel, usually narrowing toward the apex. Fl<»vcrs sessile (in bud), the
hypanthium turbinate, ca. 2 mm long, densely ferrugineous villosulose, the ca-
lycine cup short, the calycine lobes 4, narrowly triangular, ca. 3 mm long, ca. 1
mm wide, obtuse, thickly carnose especially dorsally, ferrugineous ciliolate at
the base outside and marginally, glabrous within; corolla lobes 4, oblong, ?4 mm
long, carnose; stamens sessile, the anthers 4, rectangular oblong, to 4 mm long,
to 1.2 mm wide; style slender, to 5.5 mm long, ca. 0.4 mm wide, obtuse at the
apex, the stigmas not distinct, the ovarian disc compressed, to 1.8 mm wide, the
ovules numerous, subrotund, ca. 0.05 mm wide, compressed, reticulate.

Hoffmannia manussatani is known only from Panama. A combination of ses-
sile leaves and a peduncle of extraordinary length bearing 3 pairs of bracts along
its length readily distinguish it from all other species oi Hoffmannia.

BOCAS DLL TORo: Robalo Trail, slopes of Cerro Horqiiela, 6000 7000 ft, AUcn 4978 (MO).

18. Hoffmannia morii Dwyer.*'- type: Panama, Mori & KoUunki 5915 (MO, ho-
lotype).

Siihshruhs to 1 m tall, the stem simple, obtusely angular, rimose, glabrous or
glabrescent, covered with mosses, the nodes to 4 cm apart. Leaves widely oblong,
13-25 cm long, 7-12 cm wide, deltoid at the apex, obtuse or widely cuneate
toward the base, the costa subplane above, prominent beneath, the lateral veins
ca. 10, arcuate, not joining to form a conspicuous submarginal vein, thin coria-
ceous when dry, slightly discolorous, glabrous above, puberulent beneath on the
veins and costa, often moss covered; petioles to 3 cm long, to 1 mm wide, not
vesciculiferous or with poorly developed vescicles. Inflorescences axillary and
terminal, the flowers numerous in globose heads, 2 cm in diam. Flowers with
pedicels ca. 2.5 cm long; hypanthium subrotund or turbinate, ca. 2 mm long, the
hairs elongate, to 3 mm long, the calycine cup ca. 2 mm long, the lobes 4, narrowly
oblong, ca. 4.5 mm long, thickly petaloid or subcoriaceous with purple hairs to
2 mm long outside; corolla tube narrowly cylindrical, glabrous, 5 mm long, 2.5
mm wide, thickly petaloid, the lobes 4, oblong, triangular, ca. 3 mm long, cu-
cullate; stamens 4, the anthers oblong ca. 3 mm long, the filaments ca. I mm

''- floffhiaiinia Diorii Dwyer, spec. nov. Suffrmices ad 1 m alti. Folia lato-oblonga, 13-25 cm
longa, 7-12 cm lata, apice deltoidea, venis lateralibus circa 10 arcuatis, tenui-coriacca supra glabra
subtus in venis et costa puberula saepe. Inflorescentidc axillares et terminales fjoribus dense confertis
in capitiila globosa circa 2 cm diam. Flares pedicellis 2.5 mm longis; cupula calycis 2 mm longa, lobis
4 angusto-oblongis 4.5 mm longis, extus ciliis purpureis elongatis ad 2 mm longis ornatis, intus glabris;
corolla tubo 5 mm longo, lobis 4 oblongo-triangularibus, 3 mm longis; staminibus 4 antheris oblongis
3 mm longis filamcntis 1 mm longis; stylo 6.5 mm longo stigmatibus ovatis.

1980]

OWYKR— FLORA OF PANAMA (Family 179. Riihiaccac) 237

long, attached above the middle of the tube; style slender, ca. 6.5 mm long,
slightly expanded toward the apex, the stigmas 2, ovate, ca. 2-8 mm long, ca.
1.5 mm wide, conspicuously thickened. Fruits not seen.

Hoffmannla inorii is known only from Panama. It is named in honor of Dr.
Scott Mori whose collections of Panamanian Rubiaceae have been of great as-
sistance.

The new species is closely related to H. vesciculifcra Standley. The latter has

conspicuous inflations or vescicles on the petioles which are not present or at
least not well developed in the new species, although some smaller leaves of the
new species have conspicuously dilated petioles. The leaf blades average 10 lat-

H. vesciculifi

nn

much shorter than in Standley's species.

This species is known only from the type collection.

CHiRiQuf: N of San Felix at Chiriqiii-Bocas del Toro border, road to Cerro Colorado mine. 5000-
5500 ft, Mori & Kalliinki 5915 (MO).

19. Hoffmannia ostaurea Dwyer, Ann. Missouri Bot. Gard. 56: 280. 1969. type:
Panama, Kirkbridc & Duke 501 (MO, holotype).

Succulent suhshruhs, the branchlets canaliculate when dry, glabrescent, with
crustose patches of hairs (under magnification), the juvenile branchlets with many
raphides, the internodes 1.5-3 cm long. Leaves elliptic, to 25 cm long, to 7.5 cm
wide, deltoid and short acuminate apically, the acumen to 1 cm long, the costa
plane on both sides, the lateral veins 12-15, spreading and arcuate, slender, the
intermediate veins irregular pinnatiform with intervenal areas conspicuously
spreading, thinly membranaceous, concolorous, glabrous but with vermiform ra-
phides to 0.3 mm long; petioles arcuate ascending, to 8 cm long, 0.2-0.15 cm
wide, glabrous, the raphides to 0.3 mm long; stipules not seen. Inflorescences to
1.5 cm long, the peduncle capillaceous, the flowers few. Flowers (buds) with the
hypanthium oblong rotund, ca. 2 mm long, smooth, the calycine lobes 4, unequal,
subulate, to 1.2 mm long, the raphides thick, white; corolla subcarnose, the lobes
presumably 4; filaments short; stigmas 2, capitate, ca. 1 mm long. Fruits on
pedicels to 5 mm long, rotund, to 5 mm in diam., red and red purple when dry.

Hoffmannia ostaurea is known only from Panama.

BOCAS DEL loRo: Ri'o Tcribc between Qiiebrada Huron and Quebrada Schlunjik, Kirkbridc &
Duke 501 (MO).

20. Hoffmannia pittieri Standley, Contr. U.S. Natl. Herb. 18: 140. 1916. type:
Panama, Pittier 3247 (US).

//. pamimcnsis Standley, Contr. U.S. Natl. Herb. 20: 204. 1919. type: Panama, I'ittier 3056 (US).

Shrubs to 8 m tall, the stouter branches smooth, drying tan, terete, glabrous,
the pith porate. Leaves elliptic or obovate elliptic, 9-35 cm long, 8-12 cm wide,
2.5-3 times longer than wide, acute to cuneate, rarely subrotund at the apex,
short acuminate, the acumen to 2 cm long, basally acute to attenuate acute, the

238 ANNAl.S OF- IHH MISSOURI BOTANICAL GARDEN [Vol.67

costa prominulous above, prominulous to prominent beneath, often ligneous, to
2.5 mm wide, the lateral veins (li-)15-20, widely arcuate, I(-2.2) cm apart, with
1-2 conspicuous irregular veins between each pair of lateral veins, chartaceous,
usually slightly discolorous, usually drying tan or brown, glabrous; petioles 5.5-

7 cm long, 1.5

Infl,

axils, spreading, 3-9 cm long, the primary peduncle compressed as a nodal pad,
or elongate, the fasciculate branches 1-4, the flowers usually spreading. Flowers
with the pedicels to 15 mm long, stiff, glabrous; hypanthium carnose, ribbed,
puberulent to glabrous, the calycine teeth 4, triangular, 0.6-2 mm long; corolla
yellow to greenish white, to 13.5 mm long, subcarnose, glabrous to sparsely
puberulent, the lobes 4(-5), 2.3-4.3 mm long, 1.7-3 mm wide; stamens 4, the
anthers 2.3-4.5 mm long, the connective not produced at apex or base, the fila-
ments to 1.8 mm long; ovary 2-4-loculate, the style to II mm long, the stigmas
usually ball shaped, 1.2-1.8 mm long, wider than the style, each stigma ovate
oblong to cordate. Fruits subrotund; calycine lobes persistent, to 6.5 mm long,
glabrous, drying red brown, scarcely ribbed; seeds brown, foveolate.

Hoffmannla pitticri is known only from Panama. Folsom 221 1 and 2212 have
exceptionally large fruits, to 1.4 cm long, and drying tan. Apparently the fleshy
portion surrounding the seeds did not shrink as usual on drying. The fruits of the
2 collections are described as "white and pink."

CHiRiQuf: Robiilo Trail. N slope of Cerro Horqueta, 6()(K)-7()00 ft, Allen 4962 (MO). Cciro Hor-
queta, 7000 ft. Blum ct Dwyer 2599 (MO). Cerro Horqueta, Blum £?■ Dnvcr 2643 (MO). Cerro Hor-
queta, 7000 ft, Blum ct Dwyer 2647 (MO). Las Nubes, 5.5 km N W of Rio Chiriqui Viejo, W of Cerro
Punta, 2200 m, Bii.sey 679 A (MO). Cerro Horqueta, Cochrane el al. 6244 (MO). Monte Rey above
Boquete, Croat & Porter 15655 (MO). Between Boquete & Monte Rey, Croat 15823 (MO). Alto
Respinga and above. 2800 m, D'Arey 9967 (MO). Above Boquete on La Popa, 5400 ft, D'Arex &
D'Arey 6371 (MO). Above Boquete, road to La Popa, D'Arey & D'Arey 6372 (MO). Bajo Chorro,
Boquete, 60(K) ft, Davidson 47 (MO). Bajo Chorro, Boquete, Davidson 51 (MO). Cerro Horqueta,
Dnke et al. 13651 (MO): Dwyer 8765. 8790 (both MO). Boquete, Finca Collins, ca. 5000 ft, Dwxer
& Hayden 7654 (GH, K, MO, UC, US). NW of Boquete, Cerro Horqueta, 5000-5800 ft, Dwyer et
al. 525 (MO). Boquete. Finca Collins, Dwyer & Hayden 7665 (GH, F, K, MO, NY, UC, US).
Boquete, Cerro Horqueta, Dwyer & Hayden 7745 (GH, MO). Cerro Horqueta, Dwxer & Lallatliin
8783 (MO). Boquete, Folsom 2211. 2212 (both MO). Boquete, Finca Colhns, Haxden 163 (MO).
Boquete, Finca Collins, Kirkhride 117 (MO). Cerro Horqueta, 5000-7000 ft, Kirkbride 147 (MO).
Between Pinola & Quebrado Hondo, toward summit on Chiriqui Trail, Kirkhride & Dnke 894 (MO).
Cerro Punta. 6800 ft. Rid^wax A Solis 2393 (MO). Upper Ri'o Chiriqui Viejo, near Monte Lirio, 1300-
1900 m, Seihert 165 (MO). Palo Alto, E of Boquete, Stern et al. 1025 (MO). Boquete, Finca Collins,
5500 ft. Stern et al. 1127 (MO). 6 mi above Cerro Punta, Boquete Trail, ca. 750 ft. TYson 7168 (MO).
Above Jamarillo Arriba, N slopes of Cerro Palo Alto, 4900 ft, Webster 11674 (MO). Valley of upper
Ri'o Chiriqui Viejo. White & White 2. 26 (both MO). Rio Chiriqui Viejo Valley, White 74 (MO). Bajo
Mona & Quebrada Chiquero, 1500 m, Woodson & Sehery 538 (MO). Bajo Mona & Quebrada Chi-
quero, 1500 m, Woodson & Scherx561 (MO). Casita Alta. Volciin de Chiriqui, Wo,>dson et al. 839 (MO).
coci e: Cerro Pilon, 2900 ft, l.allathin 434 (MO).

21. Hoffmannia psychotriifolia (Benth.) Griseb., Fl. Brit. W. Ind. 321. 1861.
Fig. 51.

Hiiiiiinsia psyehotriaefolia Benth. in Oerst. Vidensk. Meddel. Dansk Naturhist. Foren. KJ0benhavn
1852: 50. 1852. type;: Costa Rica, Oersted 11507. not seen.

Shrubs to 2.5 m tall, the stems ultimately slender, terete, glabrous, rubescent.
Leaves oblong, to 20 cm long, to 8 cm wide, usually 3-4 times longer than wide,
acute and finally attenuate acuminate at the apex, the acumen to 2.5 cm long,

1980J

DWYHR— FLORA OF PANAMA (Family 179. Riihiacciw)

239

Figure 51. Uoffmannia psychotriifolia (Benth.) Griseb.— A. Habit (x.45).— B. Flower, opened
to show interior (x4.95).— C. Ovary cross section (x4.95). [After Cnuit 15976.]

acute and oblique at the base, the lateral veins 10-12, widely arcuate, stiffly
chartaceous, concolorous or discolorous, glabrescent, usually with minute, scat-
tered white raphides on both the surfaces; petioles to 9 cm long, and 2 mm wide,
stipules not seen. Inflorescences in the axils of the uppermost leaves, the pedun-

240 ANNALS or THE MISSOURI HOTANICAI. GARDEN (Vol. 67

cles reduced to a pad or to 8 mm long, the flowers radially disposed, ca. 7 per
inflorescence. Flowers on slender pedicels to 4 mm long, white, to 12 mm long;
hypanthium 2-4 mm long, scarcely ribbed, often with abundant red raphides, the
calycine teeth 4, deltoid, 0.8-2 mm long, acute; corolla 8-11 mm long, the lobes
4, thickly petaloid, often longer than the tube; stamens 4, the anthers 2-2.8(-4)
mm long, the connective not produced at the apex or the base, the filaments 0.5-
1 mm long; ovary 2-3-loculate, the style to 5.5 mm long, the stigmas 2, linear,
spreading at maturity, 2-3.3 mm long. Fruits on radiate, slender pedicels, shorter
than the fruit; fruit oblong to ovate oblong, obtuse or truncate at apex, to 8 mm
long, at least 2 times longer than wide, at first delicately ribbed, smooth, glabrous,
often drying yellow, finally fleshy, in vivo china red, pink or purple.

The species ranges from Guatemala to Panama, and it also occurs in the West
Indies.

CHiRiQuf: Central valley of Rio Chiriqui Viejo, Allen 1329 (MO). Cerro Horqueta, 7000 ft, Bh/m
cC Duycr 2626 (MO). Monte Rey above Boquete, Croat ct Porter 15702 (MO). NW side of Cerro
Pando, Croat 15976 (MO). Between Concepcion and EI Hato de Volcan, D'Arcy4254A (MO). Bajo
Chorro, Davidson 198 (MO). NW of Boquete, Cerro Horqueta, 5000-5800 ft, Dwyer et al. 5IS (GH,
K, MO, NY, UC, US). Boquete, Finca Collins, Dnyer A Harden 7653, 7666, 7681 (GH, MO, US).
Boquete, Finca Collins, Dwyer A Hayden 7664 (GH, K, MO, UC, US). Boquete, Fhini^er67l (MO).
Finca Collins, Boquete, Hayden 162 (MO). Finca Collins, Boquete, 6000 ft, Kirkbride 120 (MO). 27
km NW of El Halo del Volcan, 500-5300 ft, Mori c£ Bolten 7194 (MO). Palo Alto, E of Boquete.
5000 ft. Stern et al. 1015 (MO). Quebrada Velo, 1800 m, Woodson ct Schery 258 (MO). Bajo Mona
& Quebrada Chiquero, Woodson A Schery 564 (MO). Rio Chiriqui Viejo Valley, White 53 (MO).
Casita Aha, Volcan de Chiriqui, 1500-2000 m, Woodson et al. 984 (MO), cocle: Between Cerro
Pilon and El Valle, Duke A Duver 13972 (MO).

22. HofTmannia pustulata Dwyer.'^"* type: Panama, Dressier May-June 1972 (MO-
217890, holotype).

Suhshriihs to 0.3 m tall, the stems subprostrate and rooting at the nodes, the
bark somewhat rough, the lenticels minute. Leaves deciduous, but crowded to-
ward the apex of the twiglets, narrowly obovate oblong, 4.5-8 cm long, 1.6-4.5
cm wide, deltoid to obtuse toward the apex, acute at the base, the costa prominent
above and beneath, the lateral veins 22-25, plane above, prominulous beneath,
the intervenal areas conspicuously buUate above, stiffly chartaceous, concolor-
ous, ferrugineous villosulose beneath; petioles 0.4-1 cm long, the hairs many,

ferrugineous; stipules oblong, ca. 3 mm long, leaf-like. Infloreseenees axillary,

the peduncles 0.5-5 cm long, slender, ca. 1.2 mm wide, the hairs few, the cymules
on reduced branches, the flowers few. Flowers scarcely pedicellate; calycine cup
much reduced, the lobes 4, thickly subulate, to 1 .5 mm long, the hairs few, to 0.8

*'^ Hoffmannia pnstnlata Dwyer, spec. nov. Snffrutiees ad 0.3 m aiti, caulibus in parte inferiori
subprostratis et in nodis radicibus. Folia anguste obovato-oblonga, 4.5-8 cm longa, 1.6-4.5 cm lata,
versus apicem deltoidea, venis lateralibus 22-25 interveniis supra conspicue bullatis rigido-charlacea
subtus ferrugineo-villosulosa; petiolis 0.4-1 cm longis; stipulis oblongis, hie ca. 3 mm longis, foliosis.
Inflorescentiae pedunculis 0.5-5 cm longis, gracilibus, cymulis in brevissimis ramis dispositis floribus
paucis. Flores breviter pedicellati. cupula calycis brevissima dentibus 4 crasso-subulalis, 1.5 mm
longis, ciliis paucis. ad 0.8 mm longis, glandulis 2-3 minutis inter dentes dispositis: corolla lubo
brevissimo, 1 mm longo, glabro, lobis 4 oblongis, ad 2 mm longis: staminibus 4, antheris oblongis,
ca. 2 mm longis, filamenlis brevibus, ca. I mm longis; stigmatibus, ca. 1.2 mm longis, stylo ad 2 mm
longo. Fruetus immaturi oblongi-rotundi, ca. 5 mm in diam., glabri, raphidibus multis.

19S()| DWYI-R— FLORA OF PANAMA (Family 179. Ruhimcac) 241

mm long, weak, with 2-3 minute glands between adjacent lobes, these 0.1 mm
long, red; corolla pinkish green, the tube much reduced, to 1 mm long, glabrous,
the lobes 4, petaloid, oblong, to 2 mm long, obtuse, slightly cucullate and carinate,
the raphides abundant; stamens 4, the anthers oblong, ca. 2 mm long, obtuse, the
filaments short, ca. 1 mm long; stigmas 2, thickly subulate, ca. 1.2 mm long, the
style to 2 mm long, thick, the ovules numerous, the ovarian disc pulvinate, to 0.4
mm long, 1 mm wide. Fruits (immature), oblong rotund, ca. 5 mm in diam,,
glabrous, the raphides many.

The new species is known only from Panama. Dressier has supplied important
notes concerning the plant: ''Shady bank in forest; leaves pink beneath, dark red
green above; sepals red, petals pinkish green, anther and stigma cream." The
dominance of red and pink in several plant parts is striking. Dressier pressed the
plant after cultivating it.

Hoffmannia pustulata is closely related to the recently described H. huUata
L. O. Williams which also has buUate leaves. The wide-ranging //. bullata (Mex-
ico to Costa Rica) has the leaves petiolate and the lateral veins 8-15 pairs, while
//. pustulata has leaves with petioles to 1 cm long and with 22-25 pairs of lateral
veins. The corolla of//, hullata is ''dull brick red measuring about 10 mm long,"
while //. pustulata has a pinkish green corolla about 3 mm long.

colon: Rfo Guanche, Dressier May-Jun. 1972 (MO).

23. Hoffmannia rexmontis Dwyer.'^^ type: Panama, Croat 15934 (MO, holotype).

Suhshruhs 1-1.5 m tall, the stems succulent, olive colored when dry, con-
spicuously rimose, minutely puberulent, the nodes well spaced; stipular scars
conspicuously annular. Leaves widely oblong lanceolate, 13-18 cm long, 6-8 cm
wide, deltoid and short acuminate at the apex, the acumen to 1,5 cm long, 0.15-

0.3 cm wide, the costa plane or ridged above, prominulous beneath, deltoid to
obtuse at the base, the lateral veins ca. 14, widely arcuate, the submarginal vein
undulate, 0.5 mm from the margin, the intervenal areas spreading reticulate,
membranous, concolorous, minutely white farinose on both sides; petioles 6-7
cm long, slender, to 0.15 cm wide; stipules not seen. Inflorescences axillary, the
peduncle to 1 cm long, 0.1 cm wide, the flowers few, the bracts subulate, to 1.5

mm long. Flowers not seen. Fruits pedicellate, the pedicels 0.3-10 mm long, the
scars of the pedicels rotund, to 2 mm in diam.; fruits elliptic oblong or obovate

oblong, to 0.9 cm long, to 0.38 cm wide, often truncate at the apex, attenuate
toward the base, more than twice as long as wide, yellow green when dry, gla-
brous, the ribs 8, slender; persistent calycine cup to 1 mm long, the lobes trian-
gular or subulate, to 1.5 mm long, glabrous.

^'"^ Hoffnuinnia rexmontis Dwyer, spec. nov. Suffrutices 1-1.5 m alii, caulibus succulcnlis. Folia
lato-oblongo-lanceolata, 13-18 cm longa, 6-8 cm lata apice dcltoidea, venis lateralibus ca. 14 lato-
arcuatis, membranacea utrinque minute-albido-farinosa: petiolis ad 6 cm longa. Inflorescenfiae pe-
dunculo ad I cm longo, floribus paucis; bracteis subulalis ad 1.5 mm longis. Fructus pedicellis 0.3-
2 mm longis elliptico-oblongi obovato-oblongi, ad 0.9 cm longi, glabri costis gracilibus 8; calycis
dentibus, ad 1.5 mm longis.

242 ANNALS OF THH MISSOURI BOTANICAL GARDLN [Vol. 67

Hoffmannia rex-montis is known only from Panama. It is distinguished by the
fruit drying a dull yellow green and measuring about 2(4 times longer than wide.
The fruits in the living state are red, pink or white (fide Croat). The only other
species of Hoffmannia in Panama with similar fruit measurements is H. psycho-
triifolia (Benth.) Griseb. The pedicels leave a buttonlike scar which is a distin-
guishing character. The leaves tend to be concolorous on drying, but a few of the
dried leaves suggest that they are darker above in the living state.

CHiRiQUi: NW side of Cerro Pando, cloud forest, Croat 15920, 15934 (both MO).

24. Hoffmannia rubripigmenta Dwyer, Ann. Missouri Bot, Gard. 56: 283. I960.
type: Panama, Duke 10870 (MO, holotype).

Herbs, woody, low growing, the branchlets glabrous, the raphides minute,
oblong. Leaves obovate elliptic, to 23 cm long, to 9.5 cm wide, widely deltoid at
the apex, ultimately obtuse, narrowly cuneate at the base, scarcely decurrent on
petiole, the costa prominulous above and beneath, the lateral veins ca. 8, arcuate,
the intermediate veins more divergent, ramose and evanescent; petioles to 1 cm
long, delicately alate; stipules not seen. Inflorescences a little longer than the
petioles, peduncle slender erect, glabrous, to 2 cm long, ca. 0.5 mm wide, red
when dry, the flowers ca. 8, Flowers (in bud) subsessile, narrowly oblong, ca. 1
mm wide; hypanthium oblong, to 2.4 mm long, carnose, the wall thick, to 3.5 mm
wide, glabrous, smooth, the teeth erect, triangular, to 1.5 mm long; corolla with
the tube presumably short, the lobes 4, glabrous; stamens 4, the anthers ca. 4
mm long, the connective not produced at the apex or base; style cylindrical, the
stigmas erect, oblong, ca. 1.3 mm long, scarcely thicker than the style. Fruits
not seen.

Hoffmannia rubripigmenta is known only from Panama.

darien: Between upper Ri'o Membrillo and Camp 7 on construction road to San Bias, 100-800
m, Duke 10870 (MO).

25. Hoffmannia subauriculata Standley, J. Wash. Acad. Sci. 18: 179. 1928. type:
Costa Rica, Standley & Rojas 50596 (US, holotype).

Herbs, short, woody, the stems strongly quadrangular, smooth, glabrous, the
roots fibrous. Leaves sessile, elliptic to obovate elliptic or obovate spatulate, 9-
30 cm long, 5-10 cm wide, acute and short acuminate at the apex, basally cuneate,
often widely so, to 2.5 cm wide at the base, ultimately obtuse or vaguely auricu-
late, the costa porcate above, prominent beneath, the lateral veins 10-25, widely
arcuate, prominent beneath, usually 1.5-2.5 cm apart, with usually a conspicuous
intermediate vein between a pair of lateral veins, chartaceous, glabrous, the
raphides numerous, linear; stipules not seen. Inflorescences with peduncles wiry
but not stiffly erect, arising at or near the ground level, to 4 cm long, the cymules
few flowered. Flowers (in bud) with the hypanthium glabrous, the calycine teeth
4, subulate, less than 1 mm long, puberulent; stamens and petals not seen; ovary
2-locular. Fruits oblong, to 6 mm long, the ribs 4; seeds ca. 1 mm long, coarsely

foveolate.

1980J DWYFR— FLORA OF PANAMA (Family 179. RuhuHcae) 243

Hoffmannia suhauricidata is known from Costa Rica and Panama.

BOCAS DEL TORo: Without other locality, Wcdcl 361 (MO), darifn: Cerro Pirre, Bristau 627

(MO).

26. Hoffmannia vesciculifera Standley, Publ. Field Columbian Mus., Bot. Ser. 4:
285. 1929. type: Panama, Cooper 231 (F).— Fig. 52.

Shrubs to 1,5 m tall, probably unbranched, the hairs densely ferrugineous and
arachnoid flocculose. Leaves oblong rotund to obovate oblong, to 25 cm long, to
15 cm wide, briefly acuminate at the apex, cuneate to rounded at the base, the
costa immersed above, prominulous to piano compressed beneath, the lateral
veins 12-25, widely arcuate to substrict, chartaceous, discolorous, drying brown,
usually foveolate above, the hairs diffuse, to 5 mm long, often calcified at the
base, confined to veins and venules; petioles to 4.5 cm long, conspicuously in-
flated for most of their length, the inflations to 1 cm wide, located nearer the
blade than the node, densely ferrugineous flocculose; stipules not seen. Inflores-
cences to 5 cm long, the peduncles to 2 cm long or absent, the flowers numerous
in densely bracteate axillary heads, these rounded or compressed rotund, the
bracts linear lanceolate, to 1.5 cm long, 0.3-1.5 mm wide, with 1-2 irregular
ascending submarginal veins, the margins glandular pectinate ciliate, the hairs to
4 mm long. Flowers yellow; pedicels to 4 mm long, glabrous; hypanthium oblong,
truncate, glabrous, delicately 4-ribbed, the calycine lobes 4-5, unequal in length
and width, lanceolate, 7.5-10 mm long, tapering above the middle, 1.5-2 mm
wide, the hairs ferrugineous, marginal, to 2 mm long, usually densely aggregated
above the middle, with a minute gland at junction of adjacent calycine lobes;
corolla with the tube cylindrical, to 20 mm long, glabrous, the lobes 4, shorter
than the tube, hirsute outside; stamens 4-5, the anthers oblong, ca. 4.5 mm long,
the filament ca. 0.3 mm long, the connective not produced at the apex or base,
the style slender, the stigmas clavate, spreading at maturity, wider than and about
V4 the length of the style. Fruits oblong rotund, to 5 mm long, ca. 4.5 mm wide,
delicately ribbed, the ribs ca. 10, the pericarp thin walled, the raphides abundant,
the hairs deciduous; the persistent calycine lobes 4, linear lanceolate, to 10 mm
long, the hairs deciduous in patches; seeds rectangular to cuneate, coarsely re-
ticulate.

Hoffmannia vesciculifera Standley is known from Costa Rica and Panama.

BOCAS DF I TORo: Ccpro Bonyic, above Quebrada Huron, Kirkhridc & Duke 61 1 (MO). Chiriqui'-
Calderas Trail, between Quebrada Higueron and Gutierrez, Kirkhridc & Duke 761 (MO). Water
Valley, Chiriqui Lagoon, Wcdcl 1514 (MO). Fish Creek Mts., Chiriqui Lagoon, Wcdcl 2284 (MO).
CHiRiQUi: Cerro Colorado, 50 km N of San Felix, 1200-1500 m, Mori & Dressier 7784 (MO). Buena
Vista Camp, Chiriqui Trail. 750 m, G. F. Cooper 231 (F, holotype). Cerro Colorado, upper mining
road, 20-28 mi from San Felix, Croat 33386 (MO). Cerro Colorado, 34.1 km beyond bridge over Rio
San Felix, Croat 37307 (MO), cocle: El Valle, Dressier 4071 , 4386 (both MO). Cloud forest of Cerro
Caracoral, Duke tC- Dnver 15150 (NY). La Mesa, Dwyer & Nee 11952 (MO). Cerro Colorado, Bocas
Road, 1500 m, Folsom Si Collins 1821 (MO). Cerro Caracoral Gaital, Kirkhridc 1117 (MO), colon:
Rio Iguanita, 300 m. DArcx 11256 (MO), vi.raguas: NW of Santa Fe, 11 km from Escuela
Agricola Alto de Piedra, Valley Rio Dos Bocas. 450-550 m, Mori ct al. 3810 (MO). 16 km from Santa
Fe, Atlantic slope near Rio Caloveborita. Mori et al. 6693 (MO). NW of Santa Fe, II km from
Escuela Agricola, Alto de Piedra, Mori et al. 3847 (MO). Guabal, 16 km NW of Santa Fe, 500 m.
Dressier 4796 (MO).

244 ANNALS OF- Till- MISSOURI BOIANICAL GARDI-N |Vol. 67

27. Hoffmannia witheringioides Dwyer."^ type: Panama, Croat 25551 (MO, ho-
lotype).

Suhshruhs to 1.5 m tall, the branchlets angular, black brown when dry, pu-
bescent, the hairs dense, curled, the nodes well spaced. Leaves ovate, oblong,
rotund or oblong rotund, at first seemingly anisophyllous but actually pseudo
anisophyllous, the smaller leaves on short axillary branchlets, the blades then to
4 cm long, to 2.3 cm wide, acuminate, the larger leaves to 13 cm long, to 8 cm
wide, rounded or obtuse toward the apex, attenuate cuneate toward the base, the
costa slender and prominulous above, subprominent beneath, the lateral veins
ca. 10, the venules irregular pinnatiform, arcuate, chartaceous, discolorous, dark
brown above, liver colored beneath, the hairs on both sides short, vermiform
subulate, turgid toward the base, rough to the touch; petioles 3.5-4 cm long, to
1.5 mm wide, with dense, curled hairs; stipules not seen. Flowers 1-2 per axil,
the peduncle absent, the pedicels to 0.7 cm long, slender with curled hairs; hy-
panthium turbinate, to 4 mm long, truncate, the hairs curled, lightly costate, the
calycine cup 0.5 mm long, the lobes 4, deltoid, to 1 mm long; corolla dark
yellow green, the tube cylindrical, to 4 mm long, scarcely expanded near the
mouth, petaloid, villose outside, glabrous within, the lobes 4, ovate oblong, to 4
mm long, obtuse; stamens 4, the anthers subsessile, oblong, to 2.2 mm long, the
connective to 0.5 mm long at the apex, rounded; style to 5 mm long, the stigmatic
lobes oblong, to 1.2 mm long, to 0.5 mm wide, obtuse. Fruits not seen.

Hoffmannia witheringioides is known only from the solitary collection in Pan-
ama. It bears a striking resemblance to the solanaceous genus Witheringia. The
pseudo-anisophylly of the species is striking. The reduction of the flowers to one

of Middle America.

>fj

VhRA(iUAs: Rio Segundo Braso, 8 km beyond Escuela Agricola Alto Piedra beyond Santa Fe
750 m, Croat 2555! (MO).

28. Hoffmannia woodsonii Standley, Ann. Missouri Bot. Card. 28: 471. 1941.
type: Panama, Woodson & Schery 503 (MO, holotype).

Shrubs 2-4 m tall, the twigs glabrous, smooth, the raphides dot-like white.

Leaves oblong, 5-12 cm long, l-5(-6) cm wide, attenuate acuminate at the apex,

the acumen to 1 cm long, basally attenuate cuneate or acute, the costa piano
compressed beneath, the lateral veins 4-8, prominulous above, subprominent
beneath, membranous to thinly chartaceous, glabrous or puberulent, the raphids
scattered, white, subgranular above; petioles 2-6 cm long, to 0.8 mm wide; stip-

*^-' Ilojfniannla withcrini^ioUles Dwyer, spec. nov. Snffruticcs ad 1.5 m aiti ciliis densis crispisque.
Folia ovato-oblongo-rotimda vel oblongo-rotunda primo visa ut anisophylla sed vere pseudo-aniso-
phylla, foliis minoribus in ramulis axillaribus brevibus dispositis, laminis tunc ad 4 cm longis. ad 2,3
cm latis, foliis maionbus ad 13 cm longis, ad 8 cm latis, venis lateralibus circa 10, chartacea discoloria
ciliis utrinque brevibus vermiformi-subulatis; pctiolis 3.5-4 cm longis. Flares 1-2 per axillam, pe-
dunculo nullo, pcdicellis ad 0.7 cm longis ciliis crispis ornatis, cupula calycis circa 0.5 mm longa,
lobis 4 deltoideis. ad I mm longis; corolla lubo ad 4 mm longo, extus villoso intus glabro, lobis 4, ad
4 mm longis; antheris 4, subsessilibus oblongis, ad 2.2 mm longis, apice connectivo ad 0.5 mm longo;
stylo ad 5 mm longo, lobis stigmatis oblongis 1.2 mm longis.

1980]

DWYHR— FLORA OF PANAMA (Family 179. Ruhiaceac)

245

FiGURL 52. Hoffnuinnia vc'scUulifcra Slandley. — A. Habit (x'/i). — B. Flower fx I). — C. Fruit
(x2.5). [After Dressier 4386.]

Infi

2 per axil, cymes to 1.5 cm long, the peduncles

reduced to a pad or to 6 mm long, ca. 0.5 mm wide, glabrous to ferrugineous
puberulent, the flowers usually 2-6 per cymule. Flowers with the pedicels 5-7
mm long, often radially disposed or deflexed candelabraform capillaceous, gla-
brous to ferrugineous puberulent; hypanthium square in cross section; subcar-
nose, glabrous (puberulent?), the calycine teeth 4(-5), cuneate subulate, 0.5-1.3

246 ANNALS Ol- Tflh MISSOURI BOTANICAL GARDLN [Vol. 67

mm long; corolla white, cream, yellow, to occasionally whitish pink or orange,
7-8 mm long, the lobes 4, elongate, delicately carnose, glabrous, often delicately
ribbed; stamens 4, the anthers partly exserted, 3-4.7 mm long, the connective
produced at the apex, the acumen ovate deltoid, ca. 1 mm long; ovary 2-locellate,
the style 4-5.5 mm long, the stigmas 2, oblong capitate, 1-1.2 mm long, wider
than the style. Fruits (not seen in fleshy mature stage) oblong subrotund, truncate,
to 4.5 mm long, red; persistent calycine lobes glabrous, delicately ribbed, the ribs
well spaced, varying in prominence.

Hoffnuinnia woodsouii is known only from Panama. Most collections are from
above 600 m but a few collections have been made near sea level.

BOCAS DEL TORo: Bctwccn Guttierrez Quebrada and E slope of La Zorra, Ri'o Mali, Chiriqui
Trail, Kirkhride & Duke 715 (MO), canal zone: Farfan Beach, Dwyer & Lallathin 8791 (MO). Barro
Colorado Island. Croat 10125, 10213, 14567 (all MO). CHiRiQuf: Methodist camp, near Nueva Swissa,
Croat 1351 1 (MO). Between Boquete & Monte Rey, Croat 15634 (MO). 10-11 mi W of Puerto
Armuelles, Croat 22003 (MO). 6 mi beyond Escuela Agricola Alto Piedra, 730 m, Croat & Folsom
34024, 34025 (both MO). Road between Alto Piedra School and Ri'o Calovebora, 1.8 mi beyond
school, 500 m, Croat & Folsom 34112 (MO). Cerro Horqueta, Duke et al. 13634 (MO). 12 mi W of
Puerto Armuelles, Liesner 202 (MO). Dos Lagunas, 4 km W of El Hato del Volcan, 1300 m, Mori
& Bolten 7396 (MO). Quebrada Velo, Woodson & Scherv 260 (MO). Callejon Seco, Volcan de
Chiriqui. Woodson & Schery 503 (MO), cocle: Cerro Pilon. Duyer .?■ Lallathin 8659, 8681 (all MO).
Cerro Gaital Caracoral, Dwyer S: Correa 8910 (MO). Club Campestre to La Mesa, Dn\er 10526
(MO). El Valle, Lewis etal. 1732 {GH. MO, US). Cerro Pilon, 5 km NE of El Valle, Mori 6560 {UO).

Rio Guanche, ca. 3-5 km above Portobelo road bridge, Croat 36998 (MO); COLON: Mori & Kallunki
5189 (MO). Ri'o Guanche, 2.5 km upriver from bridge, road to Portobelo, Mori et al. 6447 (MO), los
SANTOS: Loma Prieta, Duke 11849 (MO). Panama: Cerro Campana, Croat 14723, 14783 (both MO).
El Llano-Carti Road, 9.6 km from Panamerican Highway, Mori <& Kallunki 1855 (MO). Cerro Cam-
pana, 870-1000 m, Mori & Kallunki 3564 (MO). 5-10 km NE of Altos de Pacora, 750 m, Mori ct
Kallunki 4987 (MO), 16 km N of Panamerican Highway, El Llano-Carti Road, 400-450 m, Nee A
Dressier 9341 (MO), veraguas: Between Santa Fe and Rfo Calovebora, 1.8 mi beyond Escuela
Agricola, 735 m, Croat <Sc Folsom 34268 (MO).

49. ISERTIA

Isertia Schreb., Gen. PI. 1: 342. n. 602. 1789. type: L coccinca (Aubl.) Vahl.

Creatantha Standley. Field Mus. Nat. Hist., Bot. Ser. 8: 344. 1931. type;: C, peruviana Standley.

Trees or shrubs. Leaves petiolate, the stipules free, interpetiolar, erect, per-
sistent. Inflorescences large, terminal, cymose corymbose or cymose paniculate.
Flowers large, pedicellate, the hypanthium usually subovoid, the calycine cup
short, the teeth usually 4-6; corolla elongate, tubular funnelform or salverform,
rigidly petaloid, hairy within, the lobes 4-6, valvate; stamens exserted, the an-
thers linear; ovary 2-6-celled. Fruits baccate, globose, the seeds small, numer-
ous, foveolate.

Isertia occurs throughout Middle America and tropical South America. There
are about 25 species in the genus.

a. Leaf-blades usually dull grey green beneath; lateral veins 15-22; corolla tube to 20 mm

long: fruits to 0.7 cm in diam 1. /. haenkeana

aa. Leaf-blades bright gray green, grey or tan beneath when dry, usually described as whitish
beneath; lateral veins ca. 22; corolla tube to 50 mm long; fruits 1.5 cm in diam.

2. /. Iiypoleuca

I980J DWYER— FLORA OF PANAMA (Family 179. ^/(/'/V/rfflf) 247

1. Isertia haenkeana DC, Prodr. 4: 437. 1830. type: Mexico, Haenke, not seen.

/. spkaeformis Hemsl., Biol. Centr. Amer. Bot. 2: 31. 1881, not DC. 1830.

Trees to 6 m tall or shrubs, the branchlets angular, usually densely villosulose
or villose, the nodes well spaced, often constricted. Leaves oblong, widely lan-
ceolate, occasionally obovate oblong, to 46 cm long, to 28 cm wide, cuspidate,
the cusp to 2.5 cm long, deltoid to acute at the apex, subobtuse to cuneate at the
base, often attenuate, occasionally slightly inequilateral, the costa slender, sub-
immersed or slightly prominulous above, prominent beneath, especially proxi-
mally, the secondary veins 15-22, rigid, ascending, the intervenal areas with
perpendicular veinlets, membranous, papyraceous, rarely subcoriaceous, usually
dull grey green beneath when dry, glabrous above, appressed villosulose beneath;
petioles 0.5-6 cm long, the lamina decurrent adaxially, pubescent; stipules tra-
peziform or ovate elliptic, to 1.6 cm long, acute, carinate, villose. Inflorescences
terminal, solitary, puberulent, thrysiform, pyramidal, oblong or subrotund, rarely
wider than long, shorter than the terminal leaves, to 21 cm long, to 10 cm wide,
the peduncles absent or to 5 cm long, often stout, to 0.6 cm wide, angular, the
basal branches usually opposite, the lower branches of epedunculate inflores-
cences more remote from the upper branches, these appearing whorled, densely
flowered. Flowers with the hypanthium and calycine cup subrotund, ca. 4 mm
diam., to 2 mm long, sparsely pubescent outside, but with patches of glands at
or near the apex of the cup within, the teeth 4-5, scarcely evident; corolla yellow,
the tube narrowly cylindrical, ca. 20 mm long, glabrous outside, pubescent above
the middle within, the lobes ca. 6 mm long, slightly cucullate, about V3 the length
of the tube or less, golden villose within; stamens 5, the anthers oblong, ca. 3.5
mm long, acute at the apex, the filaments crassate, about as long as the anther,
attached near the corolla mouth; ovarian disc prominent, padlike, the style linear,
ca. 25 mm long, the stigmas 5, linear oblong, ca. 5 mm long. Fruits depressed
rotund, to 7 mm in diam., glabrous to puberulent; seeds brown, to 1 mm long,
favose.

Isertia haenkeana occurs in Central America, extending into Colombia and
Venezuela. It also occurs in Cuba. "Canelito," "Fruta de Mono," "Huevo de
Mono," "Murcielago," Pansus Tumatti" (San Bias).

BOCAS DtL TORo: Almirantc. Bhiin 1388 (MO); McDamel 5049 (MO). Chiriqui Lagoon, Scihcrt
1546 (US). Little Bocas, WeJel 2488 (MO, US), 2539 (MO). Santa Catalina, Blachvell el al. 2703
(MO. UC). c ANAL zone: Madden Dam Road, Bartlett & Lasscr 16442 (MO). Summit Gardens, Croat
14980 (MO). Pipeline Road within 5 mi of Gamboa Gate, DAny 9275 (MO). Drowned Rio Azote
Caballo, Dodi^c ct al. 16842 (MO). Fort Kobbe, Duke 3929 (MO). Between Puentos de Las Americas
& Fort Kobbe. Duke & Mussell 6552 (MO). George Greene Memorial, Dwyer 6756 (MO). Miraflores
Locks, Dwver & Ri>b\ns 2 (MO). Albrook, U.S. Army Tropic Test Site, Dwyer & Rohyns 52 (MO).
Chagres, Fcmller 43 (MO). Road to Corozal, Gervais 129 (F). Ancon, Gervais 149 (US). Mount
Hope, Johan.sen 57 (MO). K-10 & K-15 Road, Luteyn 1584 (MO). Headwater Rio Providencia, 7.5
km SW of Gatun Locks, Nee & GeiUrx 8678 (MO). Miraflores & Corozal, Pittier2l99 (US). Gamboa,
Pitlier 3786 (US). Agua Clara, Pittier 5931 (US). Ancon Hill, Seiherf 389 (F, MO). Road K-15, Smith
et al. 3285 (F). Pipeline Road, Smith & Smith 3303 (F). Corozal Road, StaiuUey 26832 (US). Summit,
St amllex 26940 (MO). Balboa, Staiullev 270II (US). Rio Tapia, StaiuUey 281 11 (US). Fort Randolph,
Staiullex 28605 (US). Balboa, StaiuUey 29570 (US). Summit, StaiuUey 29640 (US). France Field,
Staiullev 30340 (US). France Field, StaiuUey 30426 (US). Cerro Luisa, above Pedro Miguel Locks,
Stern & Chambers 2 (MO). Fort Sherman, Stimson 5100 (MO, US). Albrook, Stimson 5062 (MO).

248 ANNAl.S {)}• THH MISSOURI BOIANICAL GARDHN (Vol. 67

Miraflores Locks area, Tyson 1144 (MO). Gamboa Pipeline Road, Twson 1488 (MO). Beacon Hill,
Albrook Air Force Base, 7va(;// 2040 (MO, US). Farfan Beach, White 141 (F, MO). Summit Garden,
Witlicrspoon & Withcrspoon 8692 (MO). Ancon Hill, Woodson ct al. 1333 (F, MO), barro Colorado
island: Avilcs 29 (F, MO); Bani^ham 370 (US); Brown 73 (F); Croat 4055, 4063, 5076, 6107, 6331,
7050, 1/6/1 (all MO): Dodi^c 3466 (US); Dwxer 8475 (MO); Ilaxdcn 23 (MO): Kcnoxer 548 (US);
Liitcxn 78/ (F); Sliattuck 625 (F, MO), 765 {¥): Standlcx 4/056 (US): Starry /33 (F, MO); Woodson
at Sc/icry 964 (F, MO); Woodwort/i ct Vestal 352 (MO). cocLL: Cerro Pilon, Du/.e /2/77 (MO). El
Valk% WUhitr ///40 (F). colon: Camp Pina, Allen 3589 (F). Nuevo Limon, Allen 5141 (F). Guasimo
on Rio Miguel de la Borda, Croat 9997 (MO). Sabanita, Croat / /072 (MO). Santa Rita Ridge, Croat
/3/75 (F). Ocean Trail between Ri'o Indio and Miguel de la Borda, Croat 36923 (MO). Rio Guanche
near Coastal Highway, D'Arcx 10207 (MO). Road to Portobelo between Rio Piedras & Portobelo.
Elias cC Kir/./?ride 1638 (GH, MO, UC. US). Pina, Kirkhride & Hayden 3/6 (MO). 6 mi SW Portobelo.
Lutexn /420 (MO). Santa Rita Ridge, Lewis et al. 5254 (MO, MOCZ, US). 7 km NE of Puerto Pilon,
road to Portobelo, Nee 6680 (MO). 3-4 km NW of Salamanca, 13 km NE of Buenos Aires, Nee 9097
(MO). Santa Rita Hills, Smitli & Smith 3441 (F). darien: Pinogana to Yaviza. Dnke 5/66 (MO). Rio
Tuira between Pucro & Rio Ponusa, Duke 5388 (MO). S of El Real on Rio Pirre, Duke 5418 (MO).
Cana, Stern et al. 653 (MO). Tucuti, Terry & Terry /404 (F, MO), Cana. Williams 754 (US). Panama:
Pacora, Allen / /2/ (F, US). Without exact locality, Andersson (F). Burunga, Austin /4 (MO). Cam-
pana, Castillo 27 (MO). San Jose Island, Correa 89 (MO). El Llano-Carti' Road, ca. 10 km N of
Panamerican Highway, D'Arey 10660 (MO). 5-6 mi E of Chepo, Duke 4050 (MO). Between El Llano
and Rio Mamoni, Duke 5606 (MO). San Jose Island. Duke /2544 (MO); Erianson 264, 306 (both US).
Arraijan, Guerra 8 (MO). Capira, Jaramillo 36 (MO). San Jose Island, Johnston 77 (US). Chiman,
Lewis et aL 3241 (MO). Capira, Maurice 803 (MO). 6 km SW Arraijan, road to Bique, Nee 6903
(MO). Capira, Paul /35 (US). Las Sabanas, Piftier 6705 (US). Juan Franco Race Track, Standlex
2774/ (US). Tumba Muerta Road, Standley 29753 (US). San Jose Island, Stimson (MO). Arraijan.
Woodson et al. /366 (F, MO), san blas: Perme, Cooper 653 (F, US). Ustupo, D'Arcx 9500 (MO).
Mulatupu (Rio IbedO, Duke 8485 (MO). Isla de Pinas, Kirk/iride 2/4 (MO). Achituppu, Lewis et al
//8 (MO). Ailigandi, Lewis et al. 138 (GH, MO, UC, US). Niatuppu, Stier 20/ (MO), vlraguas:
Rio Concepcion, Lewis et al. 2788 (COL, DUKE, MO, UC, VEN).

2. Isertia hypoleuca Benth., Hook. L. Jour. Bot. 3: 220. 1841. type: British
Guiana, Schotuhnrghk 281 (K, not seen).

/. coccinea var. hypoleuca (Benth.) Schum. in Mart.. Fl. Bras. 6(6): 286. 1889.

Cassupa panamensis Standley, Contr. U.S. Natl. Herb. 18: 135. 1914, type: Panama, Pittier 3889
(US).

Isertia panamensis (Standley) Standley, Field Mus. Nat. Hist., Bot. Ser. 8: 346. 1931.

Trees to 20 m tall, the branches stout, grooved, densely puberulent. Leaves
oblong, to 40 cm long, to 27 cm wide, deltoid at the apex, shortly cuspidate, the
cusp slender, to 2.5 cm long, often falcate, obtuse, deltoid to cuneate at the base,
the costa subplane to prominulous above, slender, prominent, especially proxi-
mally beneath, the secondary veins ca. 22, strict, subcoriaceous, discolorous,
usually chocolate brown above, grey green, drying grey or tan brown beneath,
glabrate above except puberulent on the costa and the veins, densely puberulent
beneath; petioles to 0.9 cm long, to 0.4 cm wide, stout, often curved, puberulent,
delicately winged on the adaxial side; stipules oblong lanceolate, 1-2 cm long,
acute, coriaceous, pubescent marginally. Inflorescences terminal, solitary, pu-
berulent, shorter than or as long as the terminal leaves, to 35 cm long, to 15 cm
wide, oblong, ovate or subrotund in outline, thyrsiform; peduncles stout, to 7(-8?)
cm long, to 0.7 cm wide, lignose, angular, the branches alternate or subverticillate
except the lowermost which are often disposed as 2 opposite pairs, stiff, well
spaced; bracts and bracteoles minute, widely subulate. Flowers sessile or con-
spicuously pedicellate; hypanthium oblong, truncate, 2.0-3.5 mm long, coria-
ceous, glabrous, marcescent, the teeth scarcely evident or absent, with small
patches of obtuse glands within the cup, corolla white, turning brown to red, the

1980]

DWYl.R— FLORA OI- PANAMA (Family 179. Ruhiaccac)

249

FiCftRE 53. Isertia liypolcuca Bcnth. — A. Hahil (x'/i). — B. Flower opened to show interior
(x%).— C. Ovary cross section {xV/i). [After Slimson & Gallciios 525^. |—D. Fruit (x'/i). [After
Croat 1 67 56. \

250 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

tube narrow cylindrical, to 50 mm long, 2.0-3.5 mm wide at the middle, puber-
ulent outside, densely villose within above the middle, coriaceous, the lobes 5,
oblong, 9-15 mm long, cucullate, subcoriaceous, villose within below the middle;
stamens 6, the anthers oblong, ca. 7.5 mm long, acute, minutely apiculate, dor-
sifixed, the surface honeycomb-like, the filaments short, attached a few mm below
the mouth; style linear, 25-27 mm long, ca. 0.7 mm wide, ciliolate at the apex;
the stigmas 2, erect, oblong, ca. 5.5 mm long, 1.8-2.5 mm wide, the ovary oblong

rotund in cross section, the wall thick, replete with reddish glands, the septum
thick, 2-loculate, the placentas T-shaped, crassate, the ovules minute. Fruits

rotund or oblong, to 1.5 cm in diam., smooth, glabrescent, drying black.

The species is known from Panama, Colombia, Surinam, French Guiana, Brit-
ish Guiana, Venezuela, and Brazil.

BOCAS DEL roRo: Chiriquicito, Lewis et al. 2054 (GH, MO, US). Nances Cay Island, Wedel 5H0
(MO). Old Bank Island, Wedel 1937 (MO, US), canal zone: Pipeline Road, Folsom 2848 (MO). Rio
Pina-Rfo Media Divide, Johnston 1603 (MO), cocll: La Mesa, Gentry 6845 (MO). Above El Valle,

Road to La Mesa, 2500 ft, Tyson 6930 (MO), colon: Rfo Piedras, road to Puerto Pilon, Correa 143
(MO). Guasimo, Croat 9929 '(MO). 1 mi N of Maria Chiquita, Croat 11368 (MO). Santa Rita Ridge,
Croat 13175 (MO). Maria Chiquita, Dwxer ct Kirkbridc 780! A (MO), 7812 (GH, MO, US). Santa Rita
Ridge, Gentry 1856 (MO). Gatiin, Lao 127 (MO). Between Piiia and Gati'in, Luteyn LW9 (MO). Santa
Rita Ridge. Stimson c< Gallei^os 5254 (MO). Achiote, Tyson et al. 4551 (MO), dariln: Quebrada
Venado and Peje Swamp, Bristan 1056 (MO). Cerro Pirre, Brisian 1228 (MO, US). Rio Paya, Duke
L^ KirkhriJe 14069 (MO). Puerto St. Dorothea, Dnver 2201 (MO). Panama: Cerro Jefe, Duke 9393
(MO). Cerro Jefe and La Eneida, Dwxer 8213 (MO). Capira, Bro. Maurice 803 (US). Cerro Azul,
Tyson 2196, 4051 (both MO). Cerro Jefe, Tyson 3330 (MO), san bias: Puerto Obaldia, Croat 16756
(MO). VI RAGUAs: Cerro Tute, Gentrv 6218 (MO). Colombia: choco: Rio Mulata, Rio El Valle, base
of Alto De Buey, 100-150 m. Gentry & Fallen 17303 (MO).

1980

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

251

Indfx oi Latin Name:s

Numbers in bold face type refer to descriptions; numbers in roman type refer to synonyms
numbers with daggers (t) refer to names incidentally mentioned.

Aeginetia 55, 55t

capitata 108t
Alibertia4t, 19

edulis 19t, 19, 2lt
garapalica 21
longistipulata 19
panamensis 19

Allenanthus 4"', 22
erythrocarpus 22"*",

Alseis 3t, 24, 271-

blackiana 24
darienensis 26
floribunda 241
Amaioua 4t, 27
cormbosa 27. 30t
fagifolia 27
guianensis 27t, 27

magnicarpa 29

pedicellata 30
Amphidaysa 39

ambigua SOt, 31

neblinae 33t

spathulata 33
Anisomeris 95

brachyloba 104

englesingii 102

purpusii 104

record ii 102

spinosa 951"
Antirhea

panamensis 100
Antirhoea 34
Antirrhoea 34

22

borbonica 34
trichantha 34

J.
I

Appunia 3t, 41", 51', 36
seibertii 36

tenuiflora 36t
Arcytophyllum 5t, 38

blaerioides 38t
lavarum 38
Aspidanthera 177
rudgeoides I77t

Barleria
hirsuta I90t

Bathysa 5t, 40

stipulata 40t

veraguensis 40
Bellardia 108
Bellynkxia 36

angulata 36t
Berthiera 42t
Bcrtiera 3t, 5i, 42

aequaliramosa 43

angustifolia 43

guianensis 421", 43
— var. leiophylla 44

palustris 43

parviflora 43t

tenuis 44
Billardieria

paniculata 132
Borojoa 4t, 51", 44

atlantica 46

panamensis 461", 47

patinoi 441 , 461", 47

universitatis 46
Borrera 49
Borreria 21", 48

articularis 49t

capitata
— var. suaveolcns 54

densiflora 49

laevis 50

latifolia 52

ocimoides 53

parviflora 53
perrottetii 52

pringlei 53
pumilio 54
radicosa 49
spinosa 50t
suaveolcns 481", 54
tampicana 53
tenella 54
— var. suaveolcns 54
Bouvardia 2t, 55
glabra 55

var. gracilis 55
var. ohtusa 55

triphylla 55t
Buena 190

panamensis 1901", 193

skinneri 1 18
Callicocca

ipecacuanha 74

Calycophyllum 3t, 57

candidissimum 57t, 57
Campylobotrys 222

discolor 222t
Carapichea 59

guianensis 59t
Carinta 186
repens 188

— var. americana 188
Caruelina
filipes 104

spinosa 104
Cascarilla 106

Cassupa

panamensis 248
Cephaelis 2t, 4t, 59, 67t, 781
acuminata 75t
axillaris 61, 61t

252

ANNALS OFTHH MISSOURI BOIANICAI. GARDHN

[Vol. 67

bristanii 62
camponutans 63
chiriquiensis 70
correae 64
costaricensis 70

« ^

Croat 1 1 64
dichroa 65

dimorphandrioides 66
discolor 67
dressier! 68, 79t
elata 70
gaugeri 71

glomeriilata 71
guapilensis 67
insueta 73
ipecacuanha 2t, 74
ketinedyi 75
lastistipulata 76
mucosa 59i , 76t
nana 77

nicaraguensis 67
panamensis 77

panchocoensis 78
punicea 70
reniformis 188
tomentosa 79
tonduzii 67
violacfolia 187
vultusmimi 81
Chimarrhis 4"', 82
cymosa 82t
decurrens 22
latifolia 83

parviflora 84

turbinata 82t
Chiococca 4t, 84. 84
alba 84t, 86
durifolia 87
jefensis 88

macrocarpa 86
pachyphylla 87t
pctensis 86

phacnostcmon 90
raccmosa 84"i*, 86
staminea 90
vestita 86

var. glaberrima 86
Chione 3+, 4t, 90
allenii 93
buxilblia 92, 93t
campanensis 92
chambersii 100

costaricensis 93
daricnensis 94
glabra 90t
panamensis 94
Chomelia 95, 98t
atlantica 96

brachyloba 104
coclensis 97

englesingii 102

filipes 104

grandicarpa 97
leucophylla 98

longicaudata 104
panamensis 100

peninsularis 101
psilocarpa 102

purpusii 104

recordii 102

— var. paucipubescens 103

— var. recordii 102

spinosa 104

tenuiflora 96t
Chorisies 143

obtusitblia I43"i
Chrysobalanaceae I I7"i
Cinchona 2t, 5i\ 106, 106i

cordifolia 106
officinalis 106'
pubescens 2"f', 106
— a cordata 106

tucuyensis 106
Coccocipsilum 108t

Coccocypselum 108, 108"^
canescens 1 1 1
cordifohum 108

glabrum 1 10
herbaceum 109
hirsutum 110

— var. glabrum 1 10

— var. hirsutum 110, 110
lanceolatum 1 11

macropodum 187
pleuropodum 109
repens I08t, 109
rothschuhii 109
Coffea 2•^. 112
arabica 1 12'r, 113

liberica 112i\ 113
Condalia 108, 108-E-

lanceolata 1 1 1

repens I08t, 1091'
Condaminea 4t. 5t, 113

corymbosa 1 131 , 115

peliolala 117

Cordiera 19
edulis 19

triflora 19:
Cosmibuena 4i, 5t, 117. 117-^

J.

obtusitblia 1 17
ovalis 118
paludicola 1 18
skinncri 1 18

Coussarca 4t, 5t, 120, 125t, |29t, 1311

Carolina I35t
cerroazulensis 121

curvigemmia 122
darinensis 132
durifolia 124
enneantha 125, 132t

froelichia 132
garciae 127
impetiolaris 124t

jefensis 128
latifolia 128
loftonii 129

1980

DWYER— FLORA OF PANAMA (Family 179. Ruhiaccac)

253

morii 130

nebulosa 131. 134t

ncei 132

paniculata I30t, 13lt, |32. 1351

roseo-cremea 133
veraguensis 134

villosula 135
violacea 120t

Coutarea 3t, 4t, 135

flavescens 136

hexandra 136

pubescens 136

speciosa 135t
Crcatantha 246

peruviana 246t
Crusea 2t, AK 90. 137

brachyphylla 139
— fo. microcephala 139

chiapensis 139

coccinea 137

— var. chiriquiensis 137

glabra 90t

longiflora 139

rubra 137t
Declieuxia 4t, 140

chiococcoides 140t

fruticosa 141
Deppea 4t, 143

costaricensis 143
erythrorhiza 143

floribunda 143
grandifiora 143

longipes 143

panamensis 145
Didymaea 4"^". 145

alsinoides 146

var. alsinoides I46t
var. australis 146, I46t
var. mollis 146

J.

mexicana 145

J.

Didymochlamys 147. 1491
connellii 147

whitei 147+, 149*^
Dimorphandra 67t
Diodia 2t, 5t, ISO

denudata 150

prostrata 152

sarmentosa 152

teres 152

virginiana 150t
Dressleriopsis 4*!", 153

panamensis 153"^, 154
Duggena 190

bracteosa 191

hayesii 193

leptantha 193

ovatifolia 192

richardii 190''
Duggenia

rudis 196
Duhamelia 204
Duroia 3t, 4t

panamensis 221

Elaeagnia 156
auriculata 156

nitidifolia 157

ruiz-tcranii I58t

utilis 156t
Enkylista 57

spruceana 571"
Ereicoctis 38
Evea 59

axillaris 61

dichroa 65

elata 70

guapilensis 67

guianensis 59t

ipecacuanha 74

nana 77

tomentosa 79
Evosmia 222

caripcnsis 222t
Exostema 3+, 4t, 159

mexicanum 159

parviflorum 159t

Faraniea 3t, 4t, 5t, I29t, 161, 164i. 171t

allipetens 162

bocataurensis 163

bullata 164

caput-anguis 164

corymbosa 161 1

eurycarpa 165

jefensis 166, I76■^
liesneri 167
loftonii 129
luteovircns 167, 1711
martini 132
occidentalis 4*^", 168
ovalis 170
papulata 171

pauciflora I63t, 172
platycarpa 165
scalaris 172
suerrensis 174
talamancarum 741", 175

terreycae 175

trinervia
— var. suerrensis 174

woodsonii 167'!', 175

zetcki 168
Ferdinanda 177^
Ferdinandea 177
Ferdinandusa 177

panamensis 3"^, 178

speciosa 177t

Fereira 215

vellozoana 215t
Froelichia

paniculata 132
Galium 2t, 4t, 178

aschenbornii 179
galeottianum 179
gemmiflorum 179
mexicanum 179
— var. glabratum 179
var. platyphyllum 179

254

ANNALS or THE MISSOURI BOTANICAL GARDEN

[Vol. 67

moilugo 178t

nelsonii 181

obstipum 181

uncinulatum 181

uropctalum 179
Garapatica

edulis 19, 21
Gardenia

edulis 19

gcnipa 182

hexandra 27
Gcnipa 181

americana 181 1, 182
— var. caruto 182

fo. grandifolia 182

barbata 182

caruto 182

curviflora 184

edulis 19

cxcclsa 182

grandifolia 182

humilis 182

oblongifolia 182

pubcscens 182

williamsii 185t, 185
Geocardia 186

herbacea 188

pleuropoda 108

violaefolia 187
Gcophila 4t, 186

cordata 188

croalii 187

herbacea I86t, 188

— violaefolia 187

macropoda 187
pleuropoda 108

J.

4.
1

renitormis 186

repens 188

violaefolia 187
Gomphosia 177
Gon/alagunia 190,

acutifolia 196

bracteosa 191
cornifolia 195
dependens 190t

kailunkii 192
leptantha 193
ovatifolia 192
panamensis 193
rosea 192t, 195
rudis 196
veraguensis 197

Gon/alea 190
bracteosa 191
cornifolia 193
hayesii 193
leptantha 193
ovatifolia 192
panamensis 193

Guettarda 198

armata 104
chiriquiensis 199
costaricensis 104

190t

foliacea 200

macrosperma 201

odorata 202

famuliflora 203

sanblasensis 204

speciosa I98t
Halesia 198
Hamelia 2•^, 3t, 4"^, 204, 204t

axillaris 205

boyacana 206

calycosa 206

chiapensis 206

grandiflora 207

haughtii 212

klugii 206

macranlha 207

magnifolia 208

magniloba 212

patens 208

var. coronata 21 1
var. glabra 209

var. patens 209

pauciflora 212, 212"*

purpurascens 21 1

rovirosae 211

rowlcei 212

sanguinca 212

xerocarpa 212
Hamellia 204
Hedoytis

rupestris 38t
Hemidioidia 213

ocimifolia213i, 213
Higginsia 222, 222

obovata 222t
Hillia 5t, 215, 2l8t, 219t

chiapensis 216

subsp. chiapensis 216t
subsp. grandifolia 216
— var. chiapensis 216

t

chiriquiensis 216
hathewayi 219
ligulifolia 218
maxonii 220

palmana 219
panamensis 219

J.

parasitica 215

tetrandra 219t. 220
Himantandraceae 22t
Hippotis 221

albiflora 221
trifolia 2211
Hoffmannia 2t. 3t, 222, 226t, 233t, 236t,
242t, 244t
aeruginosa 225
araneopedaria 226
arcolata 227, 23 It
bullata 227, 24 It
capillacea 228
cercidifolia 228

davidsoniae 229
eliasii 229

fimbrianthera 230

\

\

1980]

DWYHR— FLORA OF PANAMA (Family 179. Rnhiaceae)

255

gentry! 230
haydenii 231

kirkbridei 232
lancistigma 232
lenticellata 233
lewisiana 233
longicalycina 234
manussatani 234
morii 236
ostaurea 237
panamensis 237
pedunculata 222t

pitticri 237

psychotriifolia 238, 2421-

pustulata 228t, 240

rexmontis 241

rubripigmenta 242

subauriculata 226t, 242

vesciculifera 237+, 243

witheringioides 244

woodsonii 244
Houstonia

fruticosa 141
Ipecacuanha 59

officinalis 59t, 74
Isertia 3t, 246

coccinea 246t

var. hypoleuca 248

haenkeana 247

hypoleuca 248

panamensis 248

spicaeformis 247
Ixora 21", 4t

occidentalis 168

spinosa 104
Joosia 31", 5
Kinkina 106
Ladenbergia 3i', 5t
Laugeria 198

odorala 198t, 202

Leguminosae 98t

Lindenia 4"^, 5'^

nivalis 2t
Lipostoma 108

capilata 108i
Lonicera 204, 204t

alba 84+, 86
Machaonia 5+, 95+
Macrocnemum

candidissimum 57

corymbosum 1 15
Mallostoma 38

lavarum 38
Mapouria

cordata 188

herbacea 188
Matthiola 198

scabra I98t
Milracarpus 5t
Morinda 3+, 4t, 38t
Mussaenda

Candida 57

candissima 57

J.

Nerter 4t

alsinoides 146
O'Higginsia 222

obovata 222t

Orchidaceae 70t
Ouragoga

ipecacuanha 74
Pentagonia 3+
Pittierothamnus 31

Pittoniotis

trichantha 34
Portlandia

hexandra 136
Posoqueria 3t, 4+, 5t
Pseudochimarrhis 82

turbinata 82

-1.

Pseudorhachicallis 38

Psychotria 2+, 3+, 4t, 60+, 62+, 78+

acuminata 4+

aubletiana 61, 62+

axillaris 61 + , 62+

boquetensis 3t

glomerulata 71

herbacea 186+, 188

ipecacuanha 75+

macropoda 187

poeppigiana 79

repens 188

Pterocarpus 981
Quinquina 106
Rachicallis 38
rupestris 38

4,
I

Randia 2+, 4+, 95t
Rhamnaceae 108+
Richardia 2+, 5+
Ronabea
erecta 62t

Rondeletia 2+, 3t, 5t
repens 188

Rubia

acuminata 179

Rubiaceae 1, 2+, 3+

4+,

5+

22+, 27+, 92+,

95t, 135+, 154+, 190+, 223+, 237+

subfam. Rubioideae 5+

tribe Cinchoneae 4+, 5t

tribe Condamineae 5t

tribe Coussareae 5t

tribe Gardenieae 5+

tribe Hamelieae 5+

tribe Mussaendeae 5

tribe Naucleeae 5+

tribe Oldenlandieae 5+

tribe Rondeletieae 5

J.

t

4.
)

Rustia 5t, 43t
Sabicea

ambigua 30+, 31

edulis 19
Sacconia 90
Saldanha 215

illustris 215t
Sicelium 108
Simira 3+, 4+, 5+
Solenandria 159+

256

ANNALS OF THL MISSOURI BOTANICAL GARDRN

[Vol. 67

Solendra L*^9
ixoroidcs 159'

Spcrmacocc 2"^
echioidcs 50
guianensis 50
laevis 50

lalifolia 52

longiflora 139

ocimifolia 2L^"i*, 2L^

ocimoides 53

spinosa 49

suffruticosiim 49t

tenuiflora 139

Stachyarrhena 4"^

Stenostomum 34
lucidum 34t

Stcudelago L59

Slurmia 34

lucida 34"

Styracaccac 198'

Tangaraca 204

Tapogomea 59
axillaris 61

data 70
lomentosa 79
violacca 59'i"

Tardavel 49

laevis 50

latifolia 52

spinosa 49
Tepesia 204

dubia 204t
Tetramerium

odoratissimum 168
Tocoyena 3^, 4t, 5-"
Tontanea 108t

canescens 1 1 1

glabra 110

herbacea 109

hirsuta 1 10

plcuropoda 108
Totanea 108

guianensis IOS't
Uncaria 5t. 95"
U ragoga

punicea 70
Viburnum 12()'''

costaricanum 120'i'
Withcringia 244'r

THE BOTANY AND NATURAL HISTORY OF PANAMA

A symposium to signal completion of the Flora of Panama will be held in
Panama 14-17 April 1980. Sessions will include the physical setting, archaeology,
vegetation, flora, fauna and demography of Panama and nearby countries as well
as other natural history related topics. Conservation of endangered landscapes
will be a theme. Both invited and contributed papers will be given, and either
Spanish or English may be used. Limited funds may be available to assist costs
of attending this meeting. The symposium is sponsored jointly by the Missouri
Botanical Garden, the University of Panama, and the Smithsonian Tropical
Research Institute.

Contact: Prof. Mireya D. Correa A.

Escuela de Ciencias Naturales y Farmacia
Estafeta Universitaria
University of Panama
Republic of Panama

The previous issue of the ANNALS of the Missouri Botanical Garden, Vol.
66, No. 4, pp. 591-904 + i-v was published on 22 February 1980.

OF THE

m

VOLUME 67

1980

NUMBER 2

FLORA OF PANAMA

BY Robert E. Woodson, Jr., and Robert W. Schery

and Collaborators

Part IX

Family 179. RUBIACEAE— Part Ilf

John D. DwYERt

49. IXORA
Ixora L., Sp. PI. 110. 1753; Gen. PI., ed. 5. 48. 1754. type: /. coccinea L.

Sideroxyloides Jacq., Sel. Stirp. Amer. 19. 1763. type: S.ferreum Jacq.
Siderodendron Schreb., Gen. PI. 1: 71. 1789. type: S. floribunduin A. Rich.

Trees or shrubs. Leaves occasionally ternate, usually thick; petioles occa-
sionally wanting; stipules free, usually acuminate. Inflorescences terminal or ax-
illary, corymbose, the pedicels of the flowers bracteolate. Flowers variously col-
ored, the hypanthium ovoid, the calycine cup 4(-5)-lobed; corolla salverform,
slender, the lobes 4-5, contorted; stamens 4(-5), the anthers linear, dorsifixed,
exserted; ovarian disc swollen, the style slender, the stigmas 2, exserted, the
ovary 2-celled, the ovules solitary, peltately attached to the middle of the septum.
Fruits baccate, leathery or fleshy; seeds concave convex, the testa membranous.

Ixora, a genus of about 400 species, is widely distributed throughout the trop-
ics of the world.

a.

Inflorescences capitate; corolla tube 30-45 mm long.

b. Leaves sessile or subsessile; corolla red or orange I. /. coccinea

bb. Leaves obviously petiolate; corolla white 2. I. findlavsoniana

t Part I appeared in Volume 67(1) of the Annals of the Missouri Botanical Garden.
t Missouri Botanical Garden, Box 299, St. Louis, Missouri 63166, and St, Louis University,
221 North Grand, St. Louis, Missouri 63103.

Ann. Missouri Bot. Gard. 67: 257-522. 1980.

0026-6493/80/0257-0522/$26.65/0

258 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

aa. Inflorescence cymose paniculate, spreading; corolla tube 3-5 mm long.

c. Leaves 16-24 cm long; peduncles ca. 2.2 cm long; flowers sessile or subsessile; fruits

ca. 1 cm in diam 3. I.florihunda

cc. Leaves 7-16 cm long; peduncles 5-9 cm long: flowers with pedicels to 5 mm long;

. 4. /. nlcaraguensis

fruits ca. 0.4 cm in di

1. Ixora coccinea L., Sp. PI. 110. 1753. type: ?Ceylon, herb. Linn. (LINN 131.1,
not seen, microfiche MO).

Shrubs 0.5-2.0 m tall, much branching, the twiglets glabrous, the nodes well
spaced. Leaves sessile or subsessile, oblong, ovate oblong, elliptic or occasion-
ally obovate oblong, 2.0-8.5(-16.0) cm long, 1.2-6.2 cm wide, rounded, deltoid
to subacute at the apex, often with a terminal bristle to 1 mm long, cordate,
obtuse, basally truncate or subobtuse, the lateral veins 10-13, joining to form an
irregular, undulate submarginal vein, the intervenal areas spreading reticulate,
coriaceous, glabrous; stipules oblong, to 6.5 cm long, usually shorter, the median
awn as long as the body. Inflorescences terminal, the flowers in dense corymbs,
the aggregate to 7 cm long, occasionally wider than long, the peduncle usually
suppressed or to 1.2 cm long, occasionally with 1-2 pairs of opposite branches,
the lowermost to 2 cm long; bracts and bracteoles ovate, to 0.8 cm long, acute.
Flowers sessile; hypanthium subrotund to oblong rotund or turbinate, glabres-
cent, the calycine cup reduced, the lobes 5, ascending, triangular, ca. 1 mm long,
acute; corolla red or orange, the tube linear infundibuliform, 3.5-4.5 cm long, 0.1
cm wide, glabrous or minutely puberulent, the lobes ovate oblong or oblong, 1.2-
1.5 cm long, 0.8 mm wide, acute to widely deltoid, glabrous; stamens 4, the
anthers soon deciduous, narrowly ovate oblong, 3.5-4.2 mm long, caudate acu-
minate for ca. 1 mm, sagittate basally, the filaments shorter than the anthers,
attached at the mouth; style scarcely exserted or exserted for 5 mm, the stigmas
1 mm long. Fruits not seen.

Ixora coccinea is native to Asia. It is commonly cultivated in Central and
South America, in the West Indies and in Madagascar. The inflorescence is usu-
ally spreading and the peduncle is obvious. Croat 15624 notes: ''fruits red to dark
violet, mesocarp sweet, pulpy; seeds 2/' One collection, Correa 1142, has large
leaves reaching to 16 cm in length and distinctly petiolate. "Buquet de Novia'"";
^Tacho de Toro^^; ''Jazmin de Coral."

BOCAS DEL TORo: Bocas, airport runway, Lazor et al. 2430 (MO), canal zone: Diablo Heights,
Correa 1140, 1142 (both MO). Frijoles, Ebinger 311 (MO). Coco Solo, Gentry 6063 (MO). Gaillard
Highway between Summit Garden and Gamboa, Sullivan 21 (MO). Miraflores Locks, Tyson 1138
(MO). Fort Sherman, Tyson & Blum 3788 (MO), barro COLORADO island: Croat 4061 A, 7051,
15264 (all MO); Hay den 15264 (MO); Lourteig et al 3027 (MO), cocle: Penonome, Quiros 5 (MO).
Turnoff from main highway to Cerro Azul, Reece I (MO), dariln: El Real, Lazor & Correa 3368
(MO). F'anama: Juan Diaz, Mendez s.n. (MO).

2, Ixora findlaysoniana Wall, ex G. Don, Gen. Syst. 3: 572, 1834. type: not seen.

Shrubs or trees, to 4.5 m tall, the twigs terete, smooth, glabrous, ultimately
angular, the pith porate, the nodes 1.5-5.0 cm apart. Leaves narrowly obovate
to oblong, occasionally subfalcate, 7-18 cm long, 2.5-6.5 cm wide, obtuse to
cuneate at the apex, occasionally rounded, basally cuneate, often attenuate acute,

19801

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

259

FiGURt 54. Ixoraflorihiinda (A. Rich.) Griseb. Habit (xi/2). [After Dwyer & Robyns 35. \

the costa prominulous or immersed above, subprominent beneath, especially
proximally, the lateral veins ca. 9, arcuate, thinly coriaceous, concolorous, gla-
brous; petioles to 2 cm long, 0.10-0.22 cm wide; stipules free or occasionally
connate, oblong or hemispherical, 0.4-0.7 mm long, with 1-2 subulate awns long-
er than the body, arising from 1-3 scarcely elevated longitudinal ridges on the
body. Inflorescences terminal, capitate, to 4.5 cm long, ca. 9 cm wide, rounded.
Flowers glabrous; hypanthium short, the calycine cup ca. 0.5 mm long, eglan-

260 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

dular, the calycine lobes foliaceous, widely elliptic, to 3.5 mm long, to 2 mm
wide, acute, thickly petaloid; corolla white, the tube narrowly cylindrical, to 32
mm long, 1 mm wide, petaloid, the lobes 4, spreading or reflexed, oblong, 8 mm
long, to 3 mm wide; stamens 5, exserted, the anthers linear ovate, 1.8 mm long,
acuminate at the apex, sagittate at the base, the filaments short, attached at the
mouth; ovarian disc doughnut shaped, compressed, the style linear, as long as
the corolla tube, the stigmas narrowly oblong, to 2 mm long. Fruits not seen.

Ixora findlaysoniana is widely cultivated in the tropics; it is probably native
to Southeast Asia. ''Lila.''

CANAL zone: Summit Gardens, Nee 10859 (MO). Panama: Arraijan, Carrasquilla & Rincon 295
(MO), Paitilla, Correa 40 (MO), san blas: Trail to inland village of Armila, 3-8 km SW of Puerto
Obaldia, Mori et al. 6805 (MO). Niatuppu, Stier 222 (MO), veraguas: San Francisco, Dwyer 1299
(MO).

3. Ixora floribunda (A. Rich.) Griseb., Cat. PL Cub. 134. 1866.— Fig. 54.

Siderodendron floribundum A. Rich, in Sagra, Hist. Cuba 11: 24. 1850. type: Cuba, Richard (?P, not

seen).

Small trees, to 5 m tall, the twigs terete, smooth, glabrous, the nodes well
spaced. Leaves oblong or obovate oblong, 16-24 cm long, 7-10 cm wide, widely
cuneate to subobtuse at the apex, acuminate, the acumen to 1.5 cm long, ulti-
mately acute or obtuse, attenuate acute to somewhat rounded at the base, the
costa slender, immersed, plane or somewhat prominulous above, proximally
prominent beneath, the lateral veins 9-13, arcuate, thinly coriaceous, concolor-
ous, glabrous; petioles stout, often curved, to 2.5 cm long, to 0.25 cm wide in the
middle with a delicate adaxial wing, glabrous; stipules deltoid, to 5 mm long, ca.
1 mm wide at the base, acute, stiffly scarious, rugulose, densely villose on the
margin and within, otherwise glabrous. Inflorescences terminal, pyramidal pan-
iculate, shorter than the terminal leaves, epedunculate or with peduncles to 2.2
cm long, angular, pubescent, the branches disposed as 2-3 pairs, to 2 cm long,
the flowers aggregated into subrotund cymules, ca. 1 cm in diam. Flowers sessile
or shortly pedicellate; hypanthium subrotund, ca. 1 mm long, villosulose, with
numerous oblong glands within, ca. 0.1 mm long, the lobes 5, oblong, obtuse, ca.
0.5 mm long; corolla white, the tube narrowly cylindrical, ca. 4 mm long, ca. 0.7
mm wide, glabrous outside, densely white villose within, the lobes 4, oblong, ca.
4 mm long, concave, ca. 1.5 mm wide, petaloid, glabrous; stamens 5, the anthers
narrowly oblong, ca. 3.3 mm long, dorsifixed, the filaments plane, 0.6-1.5 mm
long; stigmas 2, unequal, oblong, connate, ca. 1.5 mm long, the ovary oblong in
cross section, the wall thick, the septum thick, the ovules several. Fruits sessile,
rotund, ca. 1 cm in diam., the persistent calycine tube ca. 0.5 mm long, drying
red, smooth, sparsely villose.

Ixora floribunda occurs in Salvador, Nicaragua, Costa Rica, Panama, and
Colombia, and the Antilles. The stipules of the type of Siderodendron floribun-
dum A. Rich, are described as ''obtussimis," which does not agree with my
description of the stipules as ''acute.'' ^'Cacho de Toro'' {Holdridge 6219).

1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 261

CANAL zone: Albrook Site, U.S. Army Tropic Test Center Site, Dwyer & Rohyns 35 (MO).
darien: Between Pinogana and Yaviza, Allen 298 (MO). Los Santos: Coabal, Progreso, W slope
above Rio Cobachen, Holdridge 6219 (MO). Panama: Pacora, Paul 785 (MO).

4. Ixora nicaraguensis Standley, Wernham, J. Bot. 50: 243. 1912. type: Nicara-
gua, Sccmann 1 17 (?BM, not seen).

/. rauwolfioidcs Standley, Trop. Woods 11: 25. 1927. type: Panama, Cooper & Slater 93 (US, ho-
lotype).

Shrubs or trees, to 7.5 m tall, the twiglets drying grey, smooth, glabrous,
somewhat twisted. Leaves elliptic, oblong or oblanceolate, 7-16 cm long, 2-7 cm
wide, short to long acuminate at the apex, acute at the base, the lateral veins 10-
15, slender, arcuate, prominulous, the intervenal areas spreading reticulate, char-
taceous, discolorous, glabrous; petioles 0.4-1.0 cm long; stipules free, triangular,

0.5

Infl^

cymose paniculate, spreading, sessile or pedunculate, the peduncles 5-9 cm long,
the branches paired, few, well spaced, many flowered; pedicels to 5 mm long.
Flowers with the hypanthium turbinate, ca. 1 mm long, puberulent, the calycine
cup 0.5 mm long, stiffly carnose, rubescent, glabrous, eglandular within, the teeth
absent or scarcely developed; corolla white, the tube cylindrical, 3-5 mm long,
glabrous outside, glabrous within except villose at the mouth, the lobes 4, oblong,
3-4 mm long, rounded, rubescent, subcarnose; anthers 4, narrowly ovate oblong,
3 mm long, acuminate, the acumen petaloid, the filaments 1 mm long; ovarian
disc ca. 0.3 mm long, the style linear, 3 mm long, the stigmas oblong, ca. 1.5 mm
long. Fruits pink, rotund to oblong rotund, to 4 mm in diam., smooth, glabrous.

Ixora nicaraguensis extends from Belize to Panama.

BOCAS DEL TORO: Changuinola Valley, Cooper & Slater 93 (US). CHiRiQuf: Puerto Armuelles,
Woodson & Schery 851 (MO). San Bartolome, Burica Peninsula, Woodson & Schery 851 (MO). San
Bartolome, Burica Peninsula, Woodson & Schery 880 (MO), darien: Rio Urucea, Bristan 1451 (MO).

50. JOOSIA

Joosia Karst., Fl. Colomb. 1: 9. Tab. 5. 1858. type: Colombia, 7. umbeUifera
Karst.

Trees or shrubs. Leaves usually elliptic or oblong, acute to long acuminate at
the apex, tapering basally; petioles short; stipules deciduous except usually per-
sistent at the tips of the twigs, elongate, free. Inflorescences terminal, cymose
umbellate, the flowers secundly disposed. Flowers with the hypanthium short,
the calycine cup and teeth short; corolla white, the lobes valvate, ovate, bifid at
the apex, the margins conspicuously scalloped; stamens with anthers often ses-
sile, attached below the middle of tube; ovary 2-celled, the ovules numerous, the
placenta adnate to the middle of the septum. Fruits capsular, linear or narrowly
oblong, splitting into 4 spirally coiled valves; seeds imbricate, membranous,
winged, the wing usually bifid at the base, entire or dentate at the apex.

Joosia, a genus of 4 species, is known from Panama, Colombia, and Peru.

262 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

1. Joosia panamensis Dwyer.*'*' type: Panama, Croat 25932 (MO). — Fig. 55.

Shrubs or small trees, the branchlets glabrescent, often covered with mosses.
Leaves elliptic or rarely obovate trapeziform, 16-22 cm long, 6-9.5 cm wide,
acute or widely deltoid toward the apex, cuneate or acute at the base, slightly
inequilateral, the costa prominulous above, prominent beneath, the lateral veins
10-15, arcuate, the veinlets dense, arising at right angles from the costa, stiffly
chartaceous, glabrescent above, beneath with white, appressed hairs; petioles to
1.5 cm long, villose; stipules free, ovate oblong, ca. 2 cm long, ca. 0.5 cm wide
at the base, obtuse, red when dry, the hairs few, white. Inflorescences terminal,
perhaps solitary, cymose umbellate, the peduncle, rigid, slender, to 8 cm long,
ca. 0. 15 cm wide, often twisted, densely pubescent, the flowers secundly disposed
and crowded on short branches arising radiately from the apex of the peduncle;
pedicels ca. 2 mm long. Flowers with the calyx cup ca. 2.5 mm long, ca. 3 mm
wide, cup shaped, expanded, densely ciliolate outside, glabrous within, the lobes
4, oblong, ca. 2 mm long, to 1.5 mm wide, rounded at the apex, glabrous within;
corolla white, the tube narrowly cylindrical, ca. 9 mm long, ca. 1.8 mm wide,
carnose, the hairs outside appressed, minute, glabrous within, the lobes 5, ovate
oblong, to 3 mm long, with 2 ovate elliptic appendages within, to 2.8 mm long,
petaloid, glabrous, scalloped; anthers 5, sessile or subsessile or filaments to 2.5
mm long, narrow oblong, ca. 3.5 mm long, attached to the tube at or below the
middle, often near the base; style (2-)4-10 mm long, the stigmas 2, unequal,
scarcely wider than the style, to 2 mm long. Fruits linear oblong, to 2 cm long,
to 0.35 cm wide, terete, scarcely angular, the ribs few, prominulous, minutely
glandular punctate; seeds spindle shaped ca. 6 mm long, the wing bifid or ero-
sulose at the base, acute, entire or fimbriate at the apex.

Joosia panamensis is known only from Panama. It is related to J. umhellifera
Karst., a well-known Colombian species. It differs in having much shorter fruits.

llifi

seeds, and presumably shorter stipules.

BOCAS DHL TORo: Chiriqiii Trail between Buena Vista Coffee Finca & Cerro Pilon, Kirkhride &

Duke 680 (MO), cocle: Alto Calvario around Rivera sawmill, 7 km N of El Cope, 700-900 m, Folsom
3240 (MO), colon: 2-3 mi up Ri'o Guanche, Kennedy & Foster 2136 (MO). Panama: 16-20 km above

Panamerican Highway road from El Llano to Carti-Tupile. 400 m. Kennedy 2694 (MO). El Llano-
Carti Road, 18 km from Panamerican Highway, Mori et al. 4592 (MO), veraguas: Road between

Escuela Agricola Alto de Piedra above Santa Fe and Ri'o Dos Bocas, 730-770 m, Croat 25932 (MO).

Valley of Rio Dos Bocas between Alto Piedra and Calovebora, 350-400 m, Croat 27471 (MO).

Caribbean slope above Rio Primero Brazo, 5 mi NW of Santa Fe, Liesner 1014 (MO). 8.8 km from

Escuela Agricola Alto de Piedra, Mori & Kallunki 3220, 3239 (both MO). 16 km from Santa Fe, near
Rio Caloveborita, Mori 6688 (MO).

*'*^ Joosia panamensis Dwyer, spec. nov. Fntiives vel arbores parvae. Folia elliptica vel interdum
obovato-trapeziformia, 16-22 cm longa, 6-9.5 cm lata, venis lateralibus 10-15, arcuatis; petiolis ad
1.5 cm longis; stipulis ovato-oblongis, ad 2 cm longis. Infloresccntiae terminales cymoso-umbellatae,
pedunculo ad 8 cm longo, floribus in ramis brevibus secundatim dispositis, ramis radiate dispositis.
Flores pedicellati, pedicellis ad 2 mm longis, cupula calycis ca. 2.5 mm longa, extus dense ciliolata,
lobis 4, ca. 2 mm longis; corolla alba, tubo ca. 9 mm longo, extus pubescente, lobis 5, ovato-oblongis,
ad 3 mm longis, ad marginem fimbriatis; stamina 5, antheris sessilibus vel subsessilibus, angusto-
oblongis, ca. 3.5 mm longis, infra medium tubi affixis. Fructus lineari-oblongi, ad 2 cm longi, teretes,
costis paucis prominulis; seminibus ellipticis, ad 6 mm longis.

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

263

Figure 55. Joosia panamensis Dwyer. — A. Habit (x'/i). — B. Flower opened to show interior
(x2). — C. Ovary longitudinal section (x5). [After Kennedy & Foster 2136.]

51. LADENBERGIA

Ladenbergia Klotzsch in Hayne, Arzn. Gew. 14: pL 15. 1846. lectotype; L.

dichotoma Klotzsch.

Cascarilla Wedd., Ann. Sci. Nat. Bot. Ser. 3. 10: 10. 1848, not CascareUa Adanson, 1763, not
Cascarilla Raf. 1838. lectotype: not selected.

264 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

Trees or shrubs. Leaves opposite, occasionally verticillate, coriaceous, peti-
olate; stipules large, often deciduous, glandular within at the base. Inflorescences
terminal, paniculate. Flowers with the hypanthium ovoid or turbinate, the caly-
cine cup cupuliform, corolla with the tube somewhat funnelform or salverform,
replete with fibers, glabrous within, the lobes 5, valvate, papillose within and on
the margins; stamens 5, the anthers linear, included, dorsifixed; ovarian disc
pulvinate, the ovary 2-celled, the style slender, the stigmatic lobes 2, included,
slender, the placentas adnate to the septum, the ovules numerous. Fruits cap-
sular, narrowly clavate or cylindrical, septicidally bivalvate from the apex to the
base; seeds numerous, imbricate, peltate, the body oblong, surrounded by a mem-
branous, dentate or laciniate wing.

Ladenbergia is known from Costa Rica, Panama, and tropical South America.

It is a genus of about 30 species.

Mori & Dressier 7850 (MO) is probably a new species but has only immature

leaves. It is the only material seen from the Province of Chiriqui although La-

denbergia has been collected several times in Costa Rica near the Panamanian

border. It was collected in the Province of Chiriqui on Cerro Colorado, 50 km N

of San Felix on the continental divide in cloud forest, alt. 1,200-1,500 m.

a. Leaf blades bullate 1. L. hullata

aa. Leaf blades not bullate.

b. Leaves 8-15 cm long, 3.0-4.3 cm wide; lateral veins ca. 10 2. L. kmrifoUa

bb. Leaves (IO-)20-45 cm long, more than 6 cm wide; lateral veins 10-12

3. L. macrocarpa

1. Ladenbergia bullata (Wedd.) Standley, Trop. Woods 34: 41. 1933.

CascariUa hullata Wedd., Ann. Sci. Nat. Bot. Ser. 3. 10: 11. 1848. type: not seen.

Trees or shrubs to 5 m tall, the twigs pubescent or glabrescent, ultimately
angular, the nodes well spaced below, somewhat crowded terminally. Leaves
oblong rotund, 11.5-38.0 cm long, 8-26 cm wide, rounded or obtuse at the apex,
not acuminate, basally rounded, obtuse or truncate, the costa plane or immersed
above, prominent beneath, to 0.35 cm wide, the lateral veins 8-11, widely arcuate,
strict, prominent beneath, the intervenal areas spreading reticulate, coriaceous,
drying dark brown above, bullate, glabrescent to golden villosulose above, golden
appressed pilose beneath; petioles 2.0-4.5 cm long, rigid, lignose, golden villo-
sulose; stipules not seen. Inflorescences terminal, solitary, golden villosulose,
cymose paniculate, to 14 cm long, to 10 cm wide, the peduncles 0.5-4.5 cm long,
the lowermost branches opposite, 1-2 cm long, angular, usually with 2 pairs of
opposite, strongly ascending, superior branches. Flowers sessile or subsessile;
hypanthium subturbinate, ca. 1 cm long, densely golden villosulose, the calycine
cup cupuliform, ca. 3 mm long, coriaceous, pubescent outside, glabrous within,
the lobes 5, 1-2 mm long, oblong to rotund, obtuse or rounded, coriaceous,
golden villosulose on margins and outside, glabrous within, drying much darker
than the hypanthium; corolla white, the tube narrowly cylindrical, 3.5-5.0 cm
long, 0.35-0.50 cm wide, densely villosulose outside, the hairs occasionally tuft-
ed, glabrous but rugulose within, coriaceous, the lobes narrowly oblong, 2.0-2.5
cm long, crassate, carnose and rugulose within; stamens 5, the anthers with tips

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 265

barely exserted from tube, narrowly oblong, ca. 7 mm long, the filaments ca. 2.5
mm long, glabrous, somewhat geniculate at point of attachment, attached ca. 1
cm below the mouth; style ca. 25 mm long, piano compressed, villose, the stig-
matic lobes 2, ca. 1.8 mm long. Fruits with pedicels to 1.5 cm long, narrowly
oblong, to 9 cm long, the valves at dehiscence to 2 cm wide, obtuse or rounded
at the apex, cuneate at the base, smooth, rugulose, glabrescent at maturity, with
a few slender longitudinal prominulous lines outside, the calycine cup persistent
in unopened fruit, finally deciduous in open fruit.

PANAMA: 8 mi S of Goofy Lake toward Cerro Jefe, Dwyer 7080 (MO). Cerro Campana, Gentry
1840 (MO). Cerro Jefe, Gentry et at. 3407 (MO); Gentry & Dnyer 5542 (MO). Cerro AzuK just E of
Goofy Lake, Gentry & Mori 13403 (MO). El Llano-Carti Road, Liesner 1260 (MO); Mori et al. 4681
(MO).

2. Ladenbergia laurifolia Dwyer, ^^ spec. nov. type: Panama, Croat 25586 (MO).

Trees to 5 m tall, the twigs terete, the bark grey when dry, lenticellate, the
nodes well spaced, the scars of the pedicels oblong rotund, ca. 4 mm in diam.
Leaves narrowly oblong or narrowly obovate oblong, 8-15 cm long, 3.0-4.3 cm
wide, deltoid toward the apex, acuminate, the acumen triangular, to 0.6 cm long,
obtuse, widely cuneate or obtuse at base, the costa prominulous above, sub-
prominent beneath, dark red, the lateral veins ca. 10, subplane on both sides,
arcuate, the submarginal vein undulate and inconspicuous, the margin lightly
revolute; petioles lignose, rigid, glabrous, to 2 cm long; stipules deciduous. In-
florescences terminal, to 11 cm long, golden puberulent, dark red when dry, the
peduncle to 0.2 cm wide, the branches 3, terminating a small branch, the lateral
branches shorter and with 3 secondary branches at the apex, the cymes few
flowered, the middle branch similar to the lateral branches but with the cymes
spreading and often on branches to 1.5 cm long; bracts oblong ovate, to 2 mm
long, puberulent, the bracteoles deciduous, to 1 mm long. Flowers sessile; hy-
panthium turbinate, ca. 4 mm long, densely sericeous, the calycine cup somewhat
swollen, ca. 0.5 mm long, the lobes 5, widely triangular or ovate, 0.3-1.2 mm
long, obtuse, carnose, glabrescent outside except at the base and the margins,
red when dry, minutely punctate; corolla white, the tube narrowly hypocrateri-
form, to 1 cm long, to 0.5 cm wide, densely sericeous outside, stiffly carnose,
the lobes 5, oblong, to 1.2 cm long, to 0.3 mm wide, glabrous within; stamens 5,
the anthers linear oblong, ca. 5 mm long, ca. 0.6 mm wide, obtuse, the filaments
ca. 1 mm long, attached near the middle of the tube; ovarian disc cushion like,
ca. 0.4 mm long, the style slender, ca. 1.5 cm long, the stigmas 2, linear oblong,
ca. 2.2 mm long, puberulent(?), the ovules membranous, winged, piano com-
pressed, oblong, ca. 0.3 mm long. Fruits not seen.

Ladenbergia laurifolia is known only from Panama. The general aspect of the

®^ Ladenbergia laurifolia Dwyer, spec. nov. Arbores ad 5 m altae. Folia angusto-oblonga vel
angusto-obovato-oblonga, 8-15 cm longa, 3.0-4.3 cm lata, versus apicem deltoidea venis lateralibus

ca. 10: petiolis ad 2 cm longis. Inflorescentiae terminales, ad 1 1 cm longae, aureo-puberulae, cymis

paucifloris; bracteis oblongo-ovatis, ad 2 mm longis. Flores sessiles; cupula calycis ca. 0.5 mm longa,

lobis 5, 0.3-1.2 mm longis; corolla alba tubo ad 1 cm longo, lobis 5, ad 1.2 cm longis; antheris ca. 5

mm longis, subsessilibus. Fructus non visi.

266

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 67

Figure 56. Ladenhergia macrocarpa (Vahl) Klotzch. Habit (xVio). [After Mori & Kallunki 5501 .]

leaf resembles that of a Lauraceae, the color suggesting the dried blades of a
Myristicaceae. L. kmrifolia has the smallest leaves among the Central American
species of the genus. The flowers are markedly small.

VERAGUAS: Beyond Rio Tres Brazos, 9 km from Santa Fe, Croat 25586 (MO).

3. Ladenbergia macrocarpa (Vahl) Klotzch in Hayne, Arzn. Gew. 14: 15. 1846.
Fig. 56.

Cinchona macrocarpa Vahl, Skiv. Naturh. Selsk. I: 120. 1790. typr: not seen.

Cascarilla macrocarpa (Vahl) Wedd. Ann. Sci. Nat. Bot. Ser. 3. 10: 13. 1848.

Trees to 40 ft tall, the trunks often buttressed, the branches smooth, glabrous,
usually angular. Leaves oblong or obovate oblong, 10-45 (or more) cm long, 6-
24 cm wide, obtuse at the apex, acuminate, the acumen short or evanescent, the
costa plane or prominulous, to 0.5 cm wide, expanded toward the petiole, prom-
inent beneath, the lateral veins 10-12, prominent beneath, coriaceous, glabrous
to densely golden pubescent above, glabrous to densely golden villosulose or

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 267

appressed pilose beneath, often the axils barbate, shiny, dark brown when dry;
petioles to 4.5 cm long; stipules crowded toward the apex of the twigs, persistent
or deciduous, oblong rotund, 1.5-3.5 cm long, to 2.5 cm wide, rounded at the
apex, coriaceous, glabrescent, venose, shiny. Inflorescences terminal, solitary,
minutely pubescent or glabrescent, to 20 cm long, to 22 cm wide, presumably
epedunculate, the several branches sharply ascending and terminating the twig-
lets, 4-9 cm long, angular. Flowers scarcely pedicellate; hypanthium narrowly
turbinate, to 0.8 cm long, tapering gradually into the pedicel, drying blackish,
smooth, glabrous or puberulent, the calycine cup bowl shaped, to 0.7 cm long,
the teeth triangular to rotund, 1.5-2.8 mm long, often obtuse; corolla with the
tube narrowly cylindrical, 3.5-5.0 cm long, apically 3-4 mm wide, the lobes
narrowly oblong, ca. 3.5 cm long, ca. 0.35 cm wide, densely golden pubescent;
anthers 5, subsessile, ca. 6 mm long, ca. 1 mm wide, attached near the mouth.
Fruits scarcely pedicellate, narrowly oblong, often slightly falcate, 7.0-13.5 cm
long, 0.8-1.2(-1.8) cm wide, obtuse at the apex, attenuate toward the base, lig-
nose, glabrescent to densely golden pilose, drying black brown or golden tan,
delicately longitudinally striate ribbed, the persistent calycine cup 6-8 mm long,
the calycine lobes ca. 1 mm long, the valves at dehiscence to 1.8 cm wide; seeds
narrowly oblong or subfusiform, 1.0-2.5 cm long, the body 1-5 mm long, the
wings often split or erosulose at the distal pole, acute at the proximal pole.

A collection of this species, Mutis 6219 (F), from Colombia has a manuscript
name L. colombiana which has not been validly published.

Ladenbergia macrocarpa is known from Panama, Colombia, and south to
Boliva. ^^Fruta Mono," ^'Michurajo."

darien: Cloud Forest on Cerro Pirre, 3700 ft, Duke 6571 (MO). W slope of Cerro Pirre, Duke
6590 (MO). Cana-Cuasi Trail between Cerro Campamento and La Escalera, E of Tres Bocas, Kirk-
hride & Duke 1313 (MO). N slope of Cerro Pirre, lower montane rain forest, 700-950 m, Mori &
Kallunki 5501 (MO).

52. LIMNOSIPANEA
Limnosipanea Hook. f. in Hook., Icon. PI. Ser. 3. 1(11): 38,/?/. 1050, 1868.

type: Brazil, Limnosipanea spruceana Hook. f.

Sipania Seemann, Bot. Voy. Herald 136. 1853. typh: S. palustris Seem,, not Sipanea Aubl. 1775,
(Rubiaceae).

Herbs, slender, erect, branching above, the branches dichotomous. Leaves
3 to numerous at a node, verticillate, the uppermost occasionally opposite; stip-
ules obsolete, often reduced to a subulate projection. Inflorescences cymose with
flowers solitary or in few-flowered terminal clusters in the angles or along the
length of usually dichotomous branches. Flowers with the calycine lobes some-
what foliaceous, persistent; corolla variously colored, hypocrateriform, the tube
narrow, glabrous within, the mouth glabrous or variously pubescent, the lobes
5, oblong, contorted; stamens 5, inserted at the mouth, the anthers oblong, dor-
sifixed, the filaments slender: ovary 2-celled, the ovules numerous, disposed on
a median septum. Capsule ovoid, scarious, loculicidally dehiscent, the seeds
numerous.

268 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

Limnosipanca is a genus of about 7 species, one of which is in Panama, the
others in tropical South America.

Literature:

Standley, P. C. 1928. Flora of Panama Canal Zone, Contr. U.S. Natl. Herb. 27:

355.
Steyermark, J. 1974. Limnosipanca Hook. f. Flora de Venezuela 9(1): 384.

Wernham, H. F. 1917. The Genus Sipanca. J. Bot. 50: 171-175.

1. Limnosipanea palustris (Seem.) Hook. f. in Hook., Icon. 1050 (/?. 38). 1868.

Sipania palustris Seem., Bot. Voy. Herald 136. 1853. type: Panama, Scemann 347 (BM, excl. spec-
imen M/c7-(;//(7V/, Melastomalaceae, not seen; MO, photo of holotype).

Perennial? herbs, the stem slender, erect, branching only in the upper '/i,
pubescent, the nodes well spaced. Leaves ternate, the uppermost opposite, ses-
sile, erect, except the basal leaves often deflexed, oblong, ca. 8 mm long, ca. 2
mm wide; stipules not seen. Flowers small, solitary or in few-flowered cymules,
terminating a pair of dichotomously disposed, angular, ascending branches, pre-
sumably a solitary flower in the angle of the dichotomy.

Limnosipanea palustris is known only from the Seemann collection (Panama)
and another from Colombia (Goudot?), neither seen by the author except for a
photograph of the first. As the species is known to me only by a photograph,
floral details have been omitted in the diagnosis.

The type collection {Seemann 347, BM) has two different elements, one a
Melastomataceae which superficially looks like Limnosipanea palustris but is
probably a species of Microlicia, The specimen, on close examination, seems to

have been first mounted on a separate piece of paper, this then pasted to the
overall sheet containing the type material of Limnosipanea palustris,

Standley (1928) said about Limnosipanea palustris: 'The only North Amer-
ican species, L. palustris (Seem.) Hook. f. was collected once in a swamp near
Panama City, but has not been found again. It is a perennial herb with hirsute
stems, ternate, sessile ovate-lanceolate leaves, and terminal cymes of small pink
flowers.'' He may have drawn freely on Seemann's original description of Sipania
palustris: ''caule hirsuto basi repente, foliis ternis sessilibus ovato-lanceolatis

acutis glabris, floribus in dichotomia solitariis sessilibus, laciniis calycinis lineari-

lanceolates acutis bi-aristatis, corolla glabra vel intus subpuberula, capsula subro-

tunda, setoso-pilosa. In swamps, near Panama, growing together with Noterophila

inundata Mart.'' (Melastomataceae). Noterophila has been reduced to synonymy

under MicroHcia,

On the other hand Standley may have seen the type sheet under discussion

in the British Museum. Tn his diagnosis (repeated in North American Flora) he

described the leaves as * 'ovate-lanceolate'' when in actuality they are oblong.

The leaves of Microlicia are strikingly ovate-lanceolate, as is easily seen on the

photograph of the sheet under discussion.

Thus, it is possible that both Seemann and Standley used the Microlicia for

parts of their diagnosis of Limnosipanea palustris, and they may have failed to

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

269

Figure 57. Lindenia nvalis Benlh. Habit (x%). [After Stern et al. 7.]

270 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

note that the sheet has been annotated with the name Noterophila (Melastoma-
taceae). The enlargement on the photograph taken by Nowicke (MO) is of the
Melastomataceae rather than the Rubiaceae element.

53. LINDENIA
Lindenia Benth., PI. Hartw. 84, 351. 1841. type: L. rlvalis Benth

Siplionia Benth., PL Hartw. 84. 1841, not Schreber, 1791, nomen nudum.

Shrubs, glabrous or pubescent, the branchlets terete. Leaves short petiolate;
stipules connate, short. Inflorescences terminal, cymose, the flowers few; bracts
subulate. Flowers pedicellate, large, showy; hypanthium turbinate, 5-ribbed, the
calyx with 5 elongate lobes, equal or unequal, persistent in fruit; corolla salver-
form, elongate, the lobes 5, oblong, contorted in bud; stamens 5, inserted at the
top of the corolla tube, the anthers sessile, linear, exserted; style slender, the
stigmatic lobes 2, pilose, erect. Fruits clavate or pyriform, 2-celled, septicidally
bivalvate, the valves splitting apically, often elastically coiled; seeds numerous,
angulate.

Lindenia is a genus of 2 species, one in the Old World and one in the New
World.

Literature:

Darwin, S. P. 1976. The genus Lindenia (Rubiaceae). J, Arnold Arb. 57: 426
449.

1. Lindenia rivalis Benth., PI. Hartw. 84. 1841, type: Mexico, Hartweg 581 (K,
not seen; photo, MO). — Fig. 57.

L. acuiiflora Hook., Icon. PL 475. 1842. type: Mexico, Linden 358 (not seen).

Subshrubs to 1.5 m tall, branching freely above, the leaves terminally aggre-
gated, the branchlets stout, terete, smooth, drying black, often with an cricoid
appearance, glabrous or minutely puberulent. Leaves salicoid or narrow elliptic,
occasionally subfalcate, 4-12 cm long, 0.8-2.5 cm wide, acute at the apex and
the base, occasionally vaguely acuminate, the costa subplane to prominulous
above, prominulous beneath and rubescent, the lateral veins 7-8, strongly ar-
cuate, reaching the margin or anastomosing to form a submarginal undulate vein,
smaller intermediate veins soon evanescent, papyraceous to thinly coriaceous,
glabrous above, glabrous or puberulent beneath especially on the costa and the
veins, the margin often revolute; petioles absent or 0.2-1 cm long, the lamina
slightly decurrent, dilated proximally. Flowers solitary, terminal, large, showy;
hypanthium oblong, ca. 1 cm long, puberulent, the calycine cup scarcely mea-
surable, the calycine lobes subulate, subequal, ca. 20 mm long, thinly carnose,
puberulent with several rows of minute rubescent glands located basally within;
corolla white, the tube slender, to 20 cm long, ca. 0.35 cm wide in middle, scarcely
expanded at the apex and the base, pubescent outside, glabrous within, the lobes
5, elliptic oblong or rhombic oblong, 1.5-2.5 cm long, to 1.2 cm wide, subacute
or obtuse at apex; stamens 5, the anthers linear ovate to oblong, ca. 1 cm long,

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhlaceae)

271

c

Figure 58. Machaonia acuminata H. & B. — A. Habit (xVi). — A. Flower opened to show
interior (x5). Fruit (x5). [After Tyson et al. 4273.]

glabrescent with a few short hairs, the filaments attached near the mouth, the
style slender, about as long as the tube, villose below middle, the stigmatic area
ca. 1 cm long, barely exserted, divided above middle into 2 lobes, these oblong,
ca. 0.2 cm wide, obtuse. Fruits capsular, pyriform, to 4 cm long (including per-
sistent calycine teeth), minutely pubescent, lignose, costate, drying dark, at first
rupturing terminally at the ovarian disc position.

272 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

Lindenia rivalis is common throughout Central America and Mexico; it has
been erroneously reported from the island of Tobago. The only other species in
the genus, L. vitiensis Seem, occurs in Fiji and New Caledonia. '*Guabita de
Quebrada."

CANAL zone: Road K-6 E of Arraijan, Croat 15036 (MO). K-9 Road, Dwyer 3008 (MO). Pipeline
Road, 16.6 km from beginning of road, Mori & Kallunki 2043 (MO). Rio Cocoli. Road K-9, Stern et
aL 7 (MO), darien: Rio Morti Drill Site 7, ca. 250 m, Duke 14208 (MO), herrera: Las Minas, Rio
Las Trancas, Stern et al. 1788 (MO). Panama: Rio La Maestra, Allen 24 (MO). Rio Maje above first
waterfall on steep wooded slopes, Croat 34411 (MO). Rio Maje 4-5 mi above waterfall near new
Bayano Lake, 100 m, Croat 34710 (MO). Panamerican Highway, near Jenene, Rio Canita, Duke 3842
(MO). Rio Bayano, crossing Panamerican Highway, Duke 3984 (MO). Rio Pasiga, to above waterfall
on 2nd main fork, Gentry 2298 (MO).

54. MACHAONIA

Machaonia H. & B., PI. Aeq. 1: 101. 1806. type: A/, acuminata H. & B.

Shrubs or trees, often armed. Inflorescences terminal, cymose paniculate, the
flowers bracteate. Flowers with the calyx obovoid or turbinate, the calycine cup
cupuliform, 4-5-lobed; corolla with the tube funnelform, hairy within, the lobes

4-5; anthers dorsifixed, included or exserted, the filaments short or long; ovarian
disc fleshy, the style slender, the stigmas 2, the ovary usually 2-celled, the ovules
solitary, pendulous. Fruits of 2 trigonous, indehiscent 1-seeded cocci separating
from below upwards from the persistent axis; seeds elongated, cylindrical, the

testa thin.

Machaonia is found in Mexico, Central America, tropical South America, and
the West Indies. It is a genus of about 30 species.

Literature:

Standley, P. 1934. 71. Machaonia H. & B. in North Am. Flora 32(3): 215-222.

a. Leaves with 8-10 lateral veins; buds with the corolla 1. 0-1.5 mm wide at the apex; anthers

0.3-0.5(-0.6) mm long _ __ 1. M. acuminata

aa. Leaves with 6-7 lateral veins; buds with the corolla ca. 2.5 mm wide at the apex; anthers

ca. 0.6 mm long 2. M. martinicensis

1. Machaonia acuminata H. & B., PI. Aeq. 1: 101. 1806. type: Ecuador, Hum-
boldt & Bonpland, not seen. — Fig. 58.

M. tysonii Dwyer. Ann. Missouri Bot. Gard., 53: 106. 1%6. type: Panama, Dwyer 5061 (MO, ho-
lotype).

Shrubs or small trees, occasionally armed with spines at nodes, to 1.2 cm
long, lignose, occasionally with very small spinules, the branchlets terete,
smooth, occasionally lenticellate, glabrous or puberulent, the nodes well spaced,
frequently distichous. Leaves oblong, ovate oblong, ovate rotund, 4.5-10.0 cm
long, 1.5-6.0 cm wide, deltoid to obtuse at the apex, often falcate cuspidate, the
cusp usually short, the base acute, obtuse or rounded, occasionally slightly in-
equilateral, the costa immersed or prominulous above, prominent beneath, the
lateral veins 8-10, arcuate, immersed or prominulous above, prominulous be-
neath, chartaceous, often stiff, usually concolorous, glabrate to villosulose above

1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 273

and beneath, the intervenal areas reticulate, the veinlets immersed beneath; pet-
ioles 0.5-2.0 cm long, slender, to 0.1 cm wide, puberulent, the hairs often dense
only on the adaxial side; stipules free, small, compressed rotund, ovate oblong
or triangular, 2.0-3.5 mm long, acute or constricted terminally into a subulate
extension about as long as the body, pubescent. Inflorescences terminal, ex-
ceeding the uppermost leaves, thrysoid paniculate, 4-15 cm long, 3-9(-15) cm
wide, the peduncles 2-4 cm long, to 0.2 cm wide, pubescent, the lowermost 2-
4 pairs of branches usually opposite, usually strict and divergent, to 7 cm long,
usually with 1 or 2 pairs of opposite secondary branches or with branches ter-
nately disposed, the cymules terminal, densely flowered. Flower buds apically
1.0-1.5 mm wide, white, 4.2-5.0 mm long, the hypanthium turbinate or oblong,
to 2 mm long, carnose, densely villosulose, the calycine cup scarcely measurable,
glabrous and eglandular within, the lobes 5-6, erect, unequal, oblong, 0.6-1.0
mm long, villosulose outside and marginally; corolla tube compressed cylindrical,
to 1.5 mm long, sparsely puberulent or occasionally glabrous outside, densely
white villose within; stamens 5, the anthers oblong or subrotund, 0.3-0.6 mm
long, dorsifixed, the filaments about as long as the anthers, attached near the
apex of the tube; stigmas 2, erect, linear, 0.3-0.6 mm long, the style 1.0-3.5 mm
long. Fruits turbinate, to 7 mm long, puberulent.

This species extends from Mexico southward to Brazil.

CANAL zone: Farfan Beach, Dwyer & Hayden 7533 (GH, K, MO, UC, US); Dwyer & Lallathin
8788 (MO). Coco Solo, Gentry 6055 (MO), cocle: Rio Hato Airstrip, Blum & Dwyer 2468 (MO). Ca.
14 mi W of Rio Hato, D'Arcy & D'Arcy 6277 (MO). Rio Hato Airstrip, Dwyer 5061 (MO). La Pintada,
Escobar U (MO). Aguadulce, Stimson 5301 (MO); Woodson et al. 1205 (MO), colon: Miguel de la
Borda, Croat 9843 (MO). 2 m S of Portobelo, Croat 11413 (MO), darien: Puerto St. Dorothea,

HERRERA

PANAMA: Bejuco, Woodson et al. 1680 (MO), san blas: Mulatuppu (Rio IbedO, Duke 8468 (MO).

veraguas

MO, UC), 7553 (MO, UC). San Francisco, Rio Los Chorros, Dwyer 1354 A (MO). Santiago, Stern
et al. 993 (MO). Sona, Tyson et al. 4265 (MO). 5 mi E of Santiago, Tyson et al. 4273 (MO).

Machaonia martinicensis (DC.) Standley, Publ. Field Mus
Ser. 22: 193. 1940.

Tertrea martinicensis DC, Prodr. 4. 481. 1830. type: not seen.
Schiedia martinicensis (DC.) A. Rich., Mem. Soc. Hist. Nat. Paris 5: 187. 1834.
Machaonia rotundata Griseb., Fl. Brit. W. \. 348. 1861. type: Jamaica, not seen.
M. rotundata war. dodgei Standley, Field Mus. Nat. Hist., Bot. Ser. 18: 1326. 1938. type: Costa Rica,
Dodge 7752 (F, holotype).

Shrubs, vines, or trees, to 10 m tall, the branchlets occasionally armed at the
nodes with spines to 1.3 cm long, lignose, the twiglets terete, smooth, glabrate,
the nodes 2-5 cm apart. Leaves opposite or rarely alternate, widely oblong,
occasionally elliptic, 4-11 cm long, 1.8-4.5 cm wide, acute or rounded at the
apex, usually contracted into a short acumen 0.3-1.0 cm long, basally widely
cuneate to subobtuse, the costa slender, plane or prominulous above, prominu-
lous to prominent beneath, the lateral veins 6-7, arcuate, the intervenal areas
minutely mosaiclike, chartaceous, usually concolorous, glabrous above and be-
neath except often minutely barbate in the axils; petioles to 1.2 cm long, 0.5-1.2
mm wide, glabrous or puberulent on the adaxial side; stipules minute, 2.5-3.0

274

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 67

Figure 59. Macrocnemum ^labrescens (Benth.) Wedd. — A. Habit (xVi). — B. Flower opened

to show interior (x2'/6). [After Busey 404.]

mm long, the body compressed rotund to triangular, the subulate process longer
than or shorter than the body. Inflorescences terminal, thrysoid paniculate, to 11
cm long, the branches plane or angular, pubescent, the lowermost 2-4 pairs of
branches usually opposite, strict and divergent, mostly with 1 or 2 pairs of op-

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 275

posite secondary branches; bracts usually linear, 5 mm long, sometimes larger
and leaflike, the cymules terminal, densely flowered. Flowers 5.5-6.5 mm long,
the buds usually 2.5 mm wide at the apex; hypanthium turbinate, the hypanthium
and calyx ca. 2.5 mm long, glabrate, the hairs small and somewhat webby or
larger and subulate, to 0.2 mm long, white, drying red, the calycine lobes 5, erect,
unequal, to 1.2 mm long, obtuse, villosulose and often sparsely pubescent on the
margins; corolla white or pale green, the tube turgid, compressed cylindrical, to
2.5 mm long, sparsely villose outside, densely villose at the mouth within; sta-
mens 5, the anthers oblong, ca. 0.6 mm long, the stigmas 2, linear, to 1.2 mm
long, the style ca. 3.5 mm long, frequently exserted. Fruits turbinate, to 6 mm
long, glabrate.

Machaonia martinicensis is found in Jamaica, Panama, Costa Rica, and Co-
lombia. Duke 9719 notes that "the flowers are very attractive to ants and bees."

BOCAS DEL TORo: Macaw Hills, Wedel 548 (MO). Nances Cay Island, Wedel 585, 2862 (both
MO). Water Valley, Wedel 834. 1751, 1788 (all MO). Fish Creek Hills, Wedel 2446 (MO). Panama:
Coco Solo Weather Station, Duke 4287 (MO). Colombia: antioquia: Punta Las Barcas Turbo, sea
level, Duke 9718 (MO).

55. MACROCNEMUM

Macrocnemum P. Br., Hist. Jam. 165. 1756. type: M. jainaicense L.

Trees or shrubs, the branchlets terete. Leaves opposite, petiolate; stipules
oblong or obovate. Inflorescences terminal or axillary, paniculate. Flowers with
the calycine cup cup-shaped, the teeth 5, persistent; corolla funnelform or sal-
verform, the lobes 5, in bud drawn in so as to give a maltese cross pattern in top
view, spreading at anthesis, wider than long, pubescent within, valvate or redu-
plicate; anthers oblong, included, the filaments partly villose; ovary 2-celled, the
ovules numerous. Fruits capsular, bisulcate, dehiscing loculicidally into 2 valves;
seeds winged.

Ma

West

configuration of the corolla lobes in the bud stage.

a. Corolla tube glabrous outside; anthers ca. 1 mm long 1. M. gUihrescens

aa. Corolla tube golden puberulent outside; anthers ca. 0.7 mm long 2. A/, pastoense

1. Macrocnemum glabrescens (Benth.) Wedd., Ann. Sci. Nat. (Paris) 4. 1: 76.
1854.— Fig. 59.

Lasionema ghihrescens Benth., Bot. Voy. Sulphur 105. 1844. type: not indicated.

Trees to 8 m tall, the branchlets terete, ultimately angular, glabrous, the nodes
usually well spaced. Leaves obovate oblong, spathulate, oblong, often vaguely
falcate, widely deltoid to obtuse at the apex, acuminate or cuspidate at the apex,
the cusps to 1 cm long, basally obtuse, acute or rounded, often inequilateral, the
costa plane or immersed above, prominulous beneath, the lateral veins 7-9, ar-
cuate, chartaceous to papyraceous, usually discolorous, glabrescent above and
beneath, occasionally glabrous, often barbate on the costa and the veins beneath;

276 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

petioles 0.5-1.5 cm long, usually stiff; stipules deciduous, foliaceous, oblong or
ovate oblong, 1.0-1,5 cm long, obtuse. Inflorescences terminal or axillary, thry-
soid paniculate, to 30 cm long, either sessile and with usually 3 branches arising
from the stem tip, or pedunculate and the peduncles to 1 1 cm long, glabrescent,
the branches few, well spaced, usually opposite or ternately disposed, the low-
ermost to 14 cm long, arcuate or strict, the cymules terminal, few flowered,
spreading; bracteoles subulate, divergent. Flowers sessile or subsessile, dark-
ening at maturity, the buds clavate capitate; hypanthium oblong, ca. 3.2 mm long,
ca. 2.2 mm wide, villosulose, the calycine tube somewhat inflated, 0.5 mm long,
petaloid, glabrous inside and out, venose, the calycine teeth 5, triangular, to 1
mm long, minutely puberulent on the margin; corolla pink, the tube cylindrical,
5 mm long or longer, ca. 2 mm wide, stiffly petaloid, glabrous outside and in
except usually villose at filament attachment point, the lobes 5, oblong rotund,
to 6.5 mm long, glabrous, venose; stamens 5, the anthers oblong rotund, ca. 1
mm long, dorsifixed with a glandular patch at point of filament attachment, the
filaments unequal, slender, subulate, 2.5-4.0 mm long, densely villose below the
middle, attached at about middle of the tube and continuing downwards in riblike
fashion; stigmas 2, oblong, ca. 0.5 mm long, convex in cross section, the style
thickly subulate, 6-8 mm long, ca. 0.25 mm wide in middle. Fruits elliptic or
fusiform, to 2.5 cm long, to 0.35 cm wide before opening, on opening the valves
united at the apex and the base, the interior of valves smooth, yellow, the outside
delicately ribbed, pubescent, glabrous, capped by a saucerlike calycine cup and
ovarian disc, both to I mm long; seeds membranaceous, fusiform, to 3 mm long,
often falcate, alate, the surface with crowded irregular lines.

Macrocnemum glahrescens is known from Costa Rica, Panama, and Colom-
bia. Duke {8041) indicates the fruits are eaten by loro. Busey and Mahler {351)
note that the tree was covered with ants. 'Talo cuadrado''; 'Talo bianco'';
^'Madrono''; ^^Guanabana Torete''; ^^Cedro Macho^'; ^^Cedro Chumaquino." In
the type photo collection at the Missouri Botanical Garden there is a collection
of Wagner from Panama deposited in the Herbarium Regium Monacense (Vien-
na). It is definitely M. glahrescens, but it is not the type.

BOCAS DEL TORO: Daytonia Farm, Cooper 448 (MO). Between Buena Vista and Cerro Pilon,
Kirkbride & Duke 680 (MO), canal zone: Chiva-Chiva Trail, Allen 2467 (F). Rio Puente, between
tunnel and natural bridge, Dod^e & Allen 17317 (MO). Chiva-Chiva Trail, Fosher^ 27392 (F). Summit,
Harvey 5227 (MO). Mindi Hills, Johnston 1734 (MO). Near Gamboa, Ri'o Mandinga, Kennedy et al.
2382 (MO). Without other locality, Lindsay 455 (MO). Gorgona, Maxon 4741 (F, MO). Rio Masambi
Grande, 2 km N of Summit Garden, 80-100 m, Nee 9^/7 (MO). 1 mi N of Summit, Tyson et al. 2756
(MO). BARRO COLORADO ISLAND! AvUes 50 (MO), 273 (F); Bailey & Bailey 233 (F, MO), 234 (F),
244 (MO); Ban^ham 172. 372 (both MO); Bro\vn 26, 76, 142, 193 (all F); Chapman s.n. (F); Croat
4578, 4806, 4944, 4955, 5806, 5933, 7190, 7848, 8699, II256A, 13169 (all MO); D'Arcy 3929 (MO);
Du'xeret al, 8459 {M0)\ Ehin^er293 (MO); Foster 694, 698 (both F); Frost 26. 135 (both F); Shattuck
423] 659 (both F, MO), 811 (MO); Wetmore & Ahhe 218 (F); Wilson I (F); Woodnorth &. Vestal 333
(F, MO). CHiRiQuf: Corotii, 6 km SW of airport, Puerto Armuelles, Busey 404 (MO). 9 km S x SW
of Puerto Armuelles, Busey 728 (MO). 12 mi W of Puerto Armuelles, Croat 22155 (MO). 2 mi SW
of Puerto Armuelles, Croat 22454 (MO), cocle: El Valle, Allen 114 (F, MO), 2910, 4223 (both MO);
Hunter & Allen 302 (MO), colon: Gatuncillo, Correa et al. 734 (MO), darien: Rlo Pirre, Bristan
56 (MO). Rio Aruza, Bristan 1346 (MO). Pidiaque, Duke 8041, 8042 (both MO). Rfo Pirre, Duke 8249
(MO). Rio Sabana, ca. 4 m above Santa Fe, Duke 10203 (MO). Between upper Rio Membrillo and
Camp M on construction road to San Bias, Duke 10873 (MO). Manene near Cuasi, Kirkbride &
Bristan 1482 (MO). La Palma, Pittier 6961 (F). Panama: Upper Rio Pequeni between Salamanca
Hydrographic Station and Rio Boqueron, Allen 17276 (MO). Las Cumhres, Duke 11825 (MO). Isla

1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 277

Pedro Gonzales, Dwyer 1716 (MO). NW slope of Red Hill, San Jose Island, Johnston 1309, 1366
(both MO). Alhajuela, Pittier 2397 (F). veraguas: Santa Fe, Allen 4378 (MO). Santiago, Dwyer 1351
(MO). Isia de Coiba, Dwyer 2374 (MO).

2. Macrocnemum pastoense Karst., Linn. 30: 150. 1859. type: Colombia, Karst,,
not seen.

Trees to 8 m tall or shrubs, the branchlets terete, ultimately angular, smooth,
minutely puberulent. Leaves oblong, oblong rotund or obovate oblong, 6-17 cm
long, 4-10 cm wide, widely deltoid to rounded at apex, acuminate, the acumen
short, widely deltoid, to 0.5 cm long, ultimately obtuse or acute, the base rounded
or obtuse, the blade occasionally contracted at proximal Vs, the costa plane or
prominulous above, prominent beneath, the intervenal areas spreading reticulate,
papyraceous, tending to dry dark at least on the upper side, glabrous above,
puberulent beneath, barbate in the axils; petioles to 3 cm long, stiff, puberulent;
stipules persistent only below inflorescence, oblong, oblong rotund to ovate ob-
long, 3 cm long, obtuse or acute, rimose, puberulent. Inflorescences solitary,
terminal, occasionally terminal and axillary, cymose paniculate, to 22 cm long,
to 10 cm wide, the peduncle to 12 cm long, terminated by a pair of reduced
leaves, the lowermost branches opposite, to 7 cm long, divergent or arcuate
ascending, the branches divided into 2 branchlets. Flowers sessile, the hypan-
thium densely golden puberulent, the calycine cup short, the 5 triangular teeth
ca. 1 mm long, longer than the cup, golden puberulent; corolla purple or lilac,
the tube narrow cylindrical 6-8 mm long, puberulent outside, glabrous within,
thinly carnose, the lobes 5, in bud disposed as a hemispherical capitate mass to
0.3 cm long, 0,5 cm wide, when expanded the lobes hemispherical, wider than
long, puberulent, the margin undulate; stamens 5, the anthers oblong, ca. 0.7 mm
long, curved when mature, subversatile, the filaments slender, to 6 mm long,
villose below the middle, attached below the middle of the tube; style slender,
ca. 7 mm long, glabrous, the stigmas 2, when separated scarcely wider than the
style, 0.3 mm long. Fruits not seen.

Macrocnemum pastoense is known only from Panama and Colombia.

darien: Pifias, Duke 10610 (M(). Rfo Tuira, near mouth of Rio Paya, Gentry 4426 (MO). Yaviza
along Rfo Chucunaque at El Punteadero, Stern et aL 173 (MO). Colombia: choco: Rio Mutata
tributary Rio El Valle, between base of Alto de Buey and mouth of river, 100-150 m. Gentry &
Fallen 17459 (MO).

56. MANETTIA
Manettia L., Mant. PI. 553. 1771. Nomen conserv. type: M

Lygistum P. Br., Civ. Nat. Hist. Jam. 142. 1756. Nomen rejic. type: Petesia hgistum L.
Nacihea Aubl. Hist. PI. Guiane 1: 95. 1775. type: no type indicated.

Conotrichia A. Rich., Mem. Soc. Hist. Nat. (Paris) 5: 277. 1829; Richard, Mem. Fam, Rub. 197.
1830. type: C. alha (Aubl.) A. Rich.

Herbs or vines, sometimes woody. Leaves petiolate, the stipules small, per-
sistent. Inflorescences axillary, cymose, occasionally terminal, the flowers oc-
casionally solitary. Flowers variable in color, the calycine lobes usually 4, short
or long, usually with teeth or small glands between the lobes; corolla tubular or

278 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

funnel shaped, 4-lobed; anthers exserted or included; ovary 2-celled, the ovules
numerous. Fruits capsular, obovoid or turbinate, septicidally bivalvate, splitting
from the apex; seeds compressed, winged.

Manettia occurs sporadically in Central America, abundantly in tropical South
America, and the West Indies. It is a genus of about 130 species.

Literature:

Wernham, H. F. 1918. The genus Manettia, J. Bot. 57 (Suppl.): 1-44.

a. Herbs, upright or scarcely upright.

b. Plants terrestrial; leaves sessile, the blades linear ovate or linear elliptic; flowers on

pedicels to 6 mm long I. M, bocataurcnsis

bb. Plants aquatic; leaves with petioles ca. I cm long, the blades ovate or rhombic ovate;

flowers sessile 3. M. hydrophila

aa. Vines.

c. Calycine lobes (6-)8; flowers red or magenta rarely white 4. M. reclinata

cc. Calycine lobes 4; flowers white 2. M.flexilis

1. Manettia bocataurensis Dwyer.*^*^ type: Panama, Croat & Porter 16334 (MO,
holotype).

Herbs, the stems weak, scarcely erect, smooth, glabrous, the nodes well
spaced. Leaves sessile, narrowly ovate or narrowly elliptic, 2.5-7.0 cm long, 0.3-
1.0 cm wide, acute at the apex and at the base, the costa plane above, subplane
beneath, the lateral veins ca. 5, strongly arcuate, the veins of the intervenal areas
not visible, thin membranous, glabrous except the minutely pubescent, slightly
revolute margins, concolorous, shiny, with minute white cystoliths beneath; stip-
ules compressed hemispherical, 2.0-3.7 mm long, the awns 3, subulate, equal or
unequal, to 2.5 mm long. Flowers solitary, axillary; pedicels slender, to 6 mm
long, puberulent; hypanthium subrotund, 1.5 mm long, glabrescent, the calycine
lobes subulate, to 1.8 mm long, puberulent; corolla white, the tube cylindrical,
to 1.2 mm long, glabrous, the lobes short. Fruits rotund, ca. 3.5 mm in diam.,
black when dry, glabrate, the calyx persistent.

Manettia bocataurensis is known only from Panama. It is related to M. steno-
phylla Donnell Smith, from Costa Rica, the type of which is Tonduz 12969 (F).
The latter species has some of the flowers short-pedicellate, in cymes, and to 8
mm in length.

BOCAS DKL TORo; Railroad near station at Milla 10, Croat & Porter 16334 (MO).

^'^ Manettia bocataurensis Dwyer, spec. nov. Planta vix erecta caulibus glabris. Folia sessilia
lineari-ovata vel lineari-elliptica, 2.5-7.0 cm longa, 0.3-1.0 cm lata, venis latcralibus ca. 5 forte
arcuatis, tenui-membranacea glabra praeter margines minuto-pubescentes; stipulis compresso-
hemisphaericis, 2.0-3.7 mm longis, setis 3, ad 2.5 mm longis. Flores in axillis solitariae pedicellis
gracilibus, ad 6 mm longis; hypanthio subrotundo, ca. 1.5 mm longo, lobis calycis subulatis, ad 1.8
mm longis; corolla alba tubo cylindrico, ad 1.2 mm longo, glabro, lobis brevibus. Fructus rotundi,
ca. 3.5 mm diam., glabrescentes. ^

1980 J

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

279

Figure 60. Manettia flexilis Brandegee. — Habit (xVi). — B. Flower opened to show interior
(x5).— C. Ovary cross section {xlVi). [After D'Arcy & D'Arcy 6616.]

280 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

2. Manettia flexilis Brandegee, Univ. Calif. PubK Bot. 6: 196. 1915. type: Mexico,
Purpus 7118 (UC, holotype),— FiG. 60.

A/, estreUae Standley, J. Wash. Acad. Sci. \5: 6. 1925. type: Costa Rica, Standley 39224 (US,
holotype).

Vines, the stems flexuous, the hairs short, curled, white. Leaves oblong, ovate
oblong or trapeziform, to 5(-7?) cm long, 1.5 cm wide or wider, acute or long
attenuate (Standley) at the apex, acute to obtuse at the base, the costa subim-
mersed above, prominulous to plane beneath, the lateral veins 7-8, arcuate, mem-
branous, glabrous to villosulose above, short pilose or villosulose beneath; pet-
ioles slender, to 1 cm long, to 0.5 mm wide, puberulent; stipules not seen.
Inflorescences axillary, umbelliform, the peduncle reduced, the flowers conglom-
erate. Flowers on short pedicels to 0.5 cm long; hypanthium to 0.5 mm long,
densely villose, the calycine cup to 0.4 mm long, the calycine lobes 4, lanceolate,
ca. 3 mm long, ca. 0.8 mm wide, venose, the hairs dense and curled outside, at
least 1 erect oblong gland to 0.35 mm long at a sinus; corolla white, the tube (in
bud) short, villose outside, glabrous inside, the lobes 4, to 3 mm long, ca. 0.7
mm wide, densely villose abaxially; stamens 4, the anthers ca. 0.7 mm long, the
filaments short. Fruits turbinate, 5-7 mm long, sparsely villose.

Manettia flexilis is known from Guatemala, Costa Rica, Panama and Colom-

bia.

CHiRiQUi: 5.4 km from Hato de Volcan on road to Las Lagunas, Correa & Lazor 1481 (MO). W
of Cerro Pando (Rio Chiriqiif Viejo), D'Arcy & DArcx 6616 (MO). Bajo Mono, Boquete, Davidson
468 (MO).

3. Manettia hydrophila Dwyer.*'^ type: Panama, Lewis et al, 2822 (MO).

Herbs, often prostrate and procumbent, the stems spreading or erect and
weak, the hairs dense, weak, wrinkled, the nodes well spaced. Leaves ovate or
rhomboid ovate, 2-5 cm long, 1.2-2.8 cm wide, acute toward the apex, basally
attenuate acute or subobtuse, often inequilateral, the costa subplane above, plane
beneath or scarcely immersed, the lateral veins ca. 6, arcuate, prominulous
above, sulcate beneath, membranous, shiny, with minute white cystoliths on both

sides, the costa and veins often with appressed uncinate hairs, the margins pu-
bescent; petioles ca. 1 cm long, puberulent; stipules short, widely triangular, ca.
1 mm long, black when dry. Inflorescences terminal or axillary and opposite, the
peduncle to 1.5 cm long, the cymes contracted, the flowers alternate and well
spaced on a flexuous rachis or on 2 branches terminating the peduncle, the
branches to 4 cm long. Flowers sessile; hypanthium compressed roiund, ca. 1

^^ Manettia hydrophila Dwyer, spec. nov. Herba aquatica nodis rhizophoris caulibus ciliis multis
crispis. Folia ovata vel rhomboideo-ovata, 2-5 cm longa, 1.2-2.8 cm lata, venis lateralibus ca. 6
arcuatis, membranacea cyslolithis multis albis utrinque costa venisque ciliis saepe adpressis uncinatis
praeditis; petiolis ca. 1 cm longis; stipulis brevibus lato-triangularibus, ca. 1 mmlongis. Inflorescentiae
terminates vel axillares pedunculo ad 1.5 cm longo, cymis contractis vel cymis floribus solitariis bene
distantibus, ad 4 cm longa. Flares sessiles lobis calycis 5 ovato-lineari-lanceolatis, ad 2.8 mm longis,
ca. 0.4 mm latis; corolla rosea, tubo ad 5 mm longo lobis 4-5, ca. 5 mm longis; antheris ad 1.3 mm
longis.

1980] DWYER— FLORA OF PANAMA (Family 179. Rublaceae) 281

mm long, the hairs appressed, white, the calycine lobes 5, ascending, narrowly
linear lanceolate, to 2.8 mm long, 0.4 mm wide, puberulent on both sides with a
distinct, red, rotund gland, 0.05 mm long, in the sinuses; corolla rose colored,
the tube cylindrical, to 5 mm long, 3 mm wide, glabrous, the lobes 4-5, ovate
oblong, 5 mm long, 3 mm wide, obtuse; stamens 4, the anthers oblong, 1.3-1.5
mm long, obtuse at the apex and base, the filaments ca. 1 mm long, attached ca.
1.5 mm from the base, the ovarian disc ringlike, 0.1 mm long, the style much
reduced, the stigmas lanceolate, acute. Fruits rotund, 3.5 mm in diam., the hairs
ascending, uncinate, to 0.5 mm long, drying at first brown, ultimately the surface
of the wall deciduous, the remainder scarious, white, smooth; seeds subrotund
or oblong, ca. 0.5 mm long, flat, red, shiny.

Manettia hydrophila is known only from Panama. It is unique in being a
prostrate or erect aquatic.

VERAGUAs: Mouth of Rio Concepcion, in swamp, Lewis ct al. 2822 (MO).

4, Manettia reclinata L. Mant. PI. 2: 553. 1771. type; not seen.

Vines, the stems slender, often with longitudinal ridges, glabrate or puberu-
lent. Leaves ovate oblong, 3.0-5.5 cm long, 1.0-2.5 cm wide, acute at the apex,
often acuminate, the acumen to 1 cm long, basally cuneate to obtuse, slightly
inequilateral, the costa plane to prominulous above, prominulous to subplane
beneath, the lateral veins 6-7, arcuate, the margin often villosulose, membranous;
petioles to 2 cm long, slender, ca. 0.7-1.0 mm wide, puberulent; stipules small,
widely triangular, to 1 mm long, wider than long, minutely puberulent and ru-
gulose. Inflorescences axillary or terminal, glabrous, 1-several flowers per axil,
the peduncle, if present, short, terminated by a pair of bracteoles subtending the
pedicels; pedicels capillaceous, to 3.5 cm long. Flowers with the hypanthium
oblong, ca. 3 mm long, the calycine cup ca. 1 mm long, with minute oblong glands
within to 0.15 mm long, the lobes (6-)8, subequal, subulate to narrowly oblong,
6-7 mm long, marginally ciliate; corolla red, scarlet, magenta, occasionally white,
the tube ca. 7 mm long, basally turgid, glabrate, minutely puberulent to densely
villose outside, densely villose within near the mouth and with a few hairs near
the base, the lobes 4, oblong, 7-8 mm long, 3-4 mm wide, obtuse, petaloid,
venose, marginally ciliate, glabrous adaxially; stamens 4, the anthers oblong, ca.
3 mm long, the filaments ca. 1 mm long, attached near the mouth; stigmas 2,
linear oblong, ca. 2.5 mm long, the style ca. 11 mm long. Fruits capsular, com-
pressed rotund, 0.6-0.8 cm in diam., smooth, glabrous or puberulent, each valve
with 3 delicate ribs, the calycine lobes persistent, curled; seeds ca. 3 mm in diam.,
the wing round, the margin undulate.

Manettia reclinata ranges from Mexico to northern South America. It also
occurs in Cuba and Jamaica. Duke 14934 notes that the Indians chew the fruits
which stain their teeth black. "Churco-guidave" (Choco). Sullivan 509 and Fol-
som 3891 have an average of 6 calyx lobes and the flowers are described as
"white.'' They may represent a new species.

BOCAS DEL TORO: Changuinola Valley, Dunlap 254 (MO). Chiriquf Lagoon, Wedcl 1217, 1271,
1669, 1714 (all MO), canal zone: Between Gatiin and Fort Sherman, Burch et al. 1017 (F, GH, K,

282 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

MO, NY, UC, US). Farfan Beach, Burch et al. 1407 (F, GH, K, MO, NY, UC). Rio Indio Hydro-
graphic Station and Natural Bridge, Dodge & Allen 17485 (MO). Madden Dam, Dwyer 8389 (MO).
Fort Sherman, Dnyer 8595 (MO). Madden Forest, Gentry 2819 (MO). Cerro Galera, Gentry 6664
(MO). Curundu, Harvey 5196 (MO). Cerro Ancon, Heriberto 51 (F). Road C-29, 6 km E of Gamboa,
Nee 8992 (MO). 1 km NW of Gamboa on E side of Ammunition Depot, Nee 9542 (MO). Between
Rodman Marine Base and Chorrera, Nowkke et al, 3591 (MO, UC, VEN). barro Colorado island:
Croat 4375, 6705, 7467, 12740 (all MO); Shattuck 159 (F, MO), 486 (MO). 13 mi N of Rio San Felix
bridge, 800-1200 m, Croat 33406 (MO), chiriqui: Frances Arriba School, ca. 14 km N of David,
Lewis et al. 684 (MO). 12 mi W of Puerto Armuelles, Liesner 205 (MO), cocle: La Mesa, Allen
2345, 2853 (both F); Croat 37483 (MO); Folsom 3891 (MO); Sullivan 509 (MO), darien: Ridge NW
of Yaviza, Duke 6538 (MO). Rio Morti Hydro Camp, Duke 15414 (MO). Rio Balsa, Duke & Nickerson
14934 (MO). LOS SANTOS: Headwaters, Rio Pedregal, Lewis et al. 2941 (MO, UC, VEN). Between
Tonosi and Macaracas, Oliver et al. 3545 (MO). Panama: Between Capira and Potrero, Dodge &
Hunter 8616 (MO). Cerro Campana, Dressier 3524 (MO). Cerro Jefe, Duke 8005 (MO). Cerro Cam-
pana, Lewis et al. 1922 (MO), 3313 (MO, UC, VEN); Kirkhride & Hayden 282 (MO). Between
Savannas and Rio Yguana, McBride 2634 (F). El Llano-Carti Road, 5 km N of Panamerican Highway,
ca. 300 m. Nee 7880 (MO). Bella Vista, Piper 5338 (F). Cerro Jefe, Tyson 3423 (MO), san blas:
Opposite Ailigandi, Lewis et al. 145 (F, GH, K, MO, NY, PA, UC, US). Ustupo, DArcy 9491 (MO).
VERAGUAS: Sona, Allen 1024 (F). 0.2 mi beyond fork in road at Escuela Alto Piedra, road to Rio
Calovebora, 750 m, Croat & Folsom 33899 (MO). Mouth of Rio Concepcion, Lewis et al. 2841 A

(MO).

57. MITRACARPUS

Mitracarpus Zucc. ex Schultes in R. & S., Syst. Veg. 3. Mant. PI. 210. 1827.
type: Mitracarpus scaher Zucc.

Herbs, annual or perennial. Leaves usually narrow, often subsessile; petioles
united with the stipules to form a setiferous sheath. Inflorescences of terminal
or axillary heads. Flowers minute, 4-merous, the calycine cup short, the lobes
unequal; corolla funnel shaped, the lobes 4(-3), valvate; stamens included or
somewhat exserted, the anthers dorsifixed. Fruits capsular, 2-celled, each cell
1-seeded, transversely or obliquely circumscissile, the calyx persistent; seeds
ellipsoid or ovoid, ventrally 4-lobed and with a cruciform groove.

Mitracarpus occurs in Mexico, Central America, South America, the West
Indies, and in Africa. It is a genus of about 40 species,

a. Stems with numerous hairs; leaves hairy above; nutlets with a single set of diagonally

crossed grooves on ventral face 2. M. hirtus

aa. Stems with few hairs; leaves glabrous or glabrescent above; nutlets with a double set of

diagonally crossed grooves on ventral face 1. M. hreviflorus

1. Mitracarpus breviflorus A. Gray, PI. Wright. 2: 68. 1853. Originally "A//-
tracarpium hreviflorum/'' type: Mexico, Wright 1120, not seen.

Herbs to 1.5 dm, branched, the stems slender, angular, sparsely hairy. Leaves
sessile, usually opposite, occasionally 4-several per node, lanceolate, ovate lan-
ceolate, to 2 cm long, to 0.4 mm wide, the lateral veins few, vague, the blades
stiff, scabrid above, otherwise glabrous, glabrous beneath, the margin revolute.
Inflorescences with heads to 1 cm wide. Flowers white. Fruits to 3 mm long,
glabrous below the line of dehiscence, pubescent above, the horns to 2 mm long,
unequal, pubescent, the seeds oblong, ca. 0.5 mm long, with 2 diagonally crossed
grooves on the ventral surface,

Mitracarpus breviflorus is known from Arizona, Mexico, Costa Rica, and

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

283

Figure 61. Mitnuarpus hirtus (L.) DC— A. Habit (x^/,).— B. Inflorescence (X2V5).— C. Seeds
(x8). [After Duke 3962.]

Panama. Johnston et al. (1949) made several collections of the species on San
Jose Island which were not seen in this study. The double set of grooves on the
face of the nutlet is diagnostic.

CHiRiQUi: Lava flow between El Hato and Bambito, D'Arcy 10041 (MO).

284 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

2, Mitracarpus hirtus (L.) DC, Prodr. 4: 572. 1830.— FiG. 61,

Spermucoce liirta L., Sp. PI., ed. 2. 148. 1762. lectotype: ?Jamaica, Herb. Linnaeus (LINN !25.4,

not seen; microfiche MO).
Mitracarpus brevifolius Gray, PI. Wright. 2: 68. 1853. type: Mexico, Wright 1120, not seen.

Herbs to 0.5 m tall, the stems simple or branched basally, angular, glabrescent
to villose, the hairs white, weak. Leaves lanceolate or oblong, 5(-6) cm long, to
1.5 cm wide, deltoid to acute at the apex, attenuate acute at the base, the costa
prominulous above, subprominent beneath, the lateral veins 3-4, often evanes-
cent, arcuate, strongly ascending, chartaceous, usually concolorous, glabrous to
pubescent above, the hairs short, stiff, scabridulous; petioles short or absent;
stipules connate, adnate to petiole, each part crescentic or hemispherical, to 4
mm long, the terminal margin with 8-10 subulate setas as long as or slightly
longer than the sheath. Inflorescences terminal and axillary, the flowers in sub-
rotund heads, to 0.7 cm in diam., burrlike because of the persistent, stiff calycine
lobes and bracteoles; bracteoles numerous, subulate or linear triangular, erect,
shorter than the capsule. Flowers very small; hypanthium short, the calycine
lobes 4, unequal, lanceolate or subulate, to 1.5 mm long, pellucid, thickly and
scarious marginate, the margin with well spaced, subulate hairs, with several
oblong glands aggregated below the middle; corolla white, the tube cylindrical,
often turgid, ca. 1.8 mm long, the lobes 4, ca. 0.6 mm long, pubescent marginally;
stamens 4, the anthers oblong rotund, ca. 0.45 mm long, glandular apiculate,
sessile or attached near the mouth; stigmas 2, minute, often exserted. Frint a
circumscissile capsule, ca. 1 mm long (exclusive of persistent calyx), dehiscing
uniformly to expose the seeds, the cap hairy, the seeds quadrangular, brown,
pitted,

Mitracarpus hirtus is a common weed throughout tropical America and the
West Indies but has been collected only sporadically in Panama. It is naturalized
in Asia and Malaysia. Steyermark (1972) has an elaborate discussion of the typ-
ification of M. hirtus in his remarks on A/. viUosus (Sw.) C. & S.

BOCAS DEL TORo: Changuinola WaUcy. Dunfap 373 (F). canal zone: A]hrook. Blum 449 (MO).
Summit Gardens, Croat 6761 (MO). Ft. Kobbe, Duke 3962 (MO). Railroad 3 km W of Gamboa, Nee
9505 (MO). CHiRiQUi: Lava flow between El Hato and Bambito, DArcy 10041 (MO), cocle: Boca
del Toabre, confluence of Rio Toabre and Rio Cocle del Norte, Lewis et al. 5487 (MO), dariln: Rm
Morti, 6 mi upstream from Morti Abajo, Duke I0I6I (MO). Rfo Tuquesa, lower Tuquesa mining
company camp called Charco Chiva, Mori 6968 (MO). Panama: Isia Taboga, Allen 1273 (MO). Cerro
Azul, Tyson 2067 (MO), vf.raguas: 0.2 mi beyond fork in road at Escuela Agricola Alto Piedra, road
to Rio Calovebora, 750 m, Croat & Folsom 33868 (MO).

58. MONTAMANS
Montamans Dwyer.*^** type: Panama, M. panamensis Dwyer.

Succulent herbs. Leaves large, the lateral veins numerous, petioles elongate;
stipules free, 2-awned. Inflorescences axillary, cymose, sessile or short pedun-

''^^ Montamans Dwyer, gen. nov. Herhae succulentae. Folia magna, venis lateralibus crebris.

Inflorescentiae axillares, cymosae floribus in caespitem parvum congestis bracteis 2-seriatim dis-
positis. Flores lobis calycinis polymorphis elongatis; corolla tubo cylindrico, lobis 5 utroque lobo

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

285

FrcuRE 62. Montamans panamensis Dwyer. — A. Leaves (x'/z). — B. Inflorescence (x<^). — C
Flower (x6).— D. Flower with calyx removed (x6).— E. Corolla opened to show interior (x 10).— F
Ovary cross section (x 10).— G. Style (x 10). fAfter Dwyer 771 L]

culate, the flowers congested into a head; bracts disposed in 2 series, often galeate
or 2-horned at the apex. Flowers with the calycine lobes elongate, polymorphic
especially at the apex; corolla cylindrical, short, the 5 lobes bearing a galeate
horn at the apex; stamens 4-5, the anthers and filaments short, attached near the
middle of the tube; style slender, the stigmas 2, slender, the ovary 2-ceIleu, the
septum thick, with a single ovule in each cell arising from the base. Fruits rotund,
fleshy, with two plane pyrenes.

Montamans is known only from Panama. The new genus is obviously a mem-
ber of the tribe Psychotriae as is evidenced by its 2-celled ovary with a thick
septum and basally attached ovule in each celK and by its fleshy fruit with 2
pyrenes. It is distinguished from the other genera of the tribe {Psychotria, Cepha-

cornu galeatum ferenlibus; antheris et filamentis brevibus prope medium tubum affixis; ovario bilo-

cellato septo medio crasso ovulis 2 basaliter affixis. Fructus rotundi carnosi pyrenis 2 plano-com-

pressis.

286 ANNALS OF THE: MISSOURI BOTANICAL GARDEN [Vol. 67

elis, Palicourea, etc.) by its elongate, polymorphic and galeate calycine lobes and
by its corolla lobes with a solid obtuse reflexed cylindrical horn below the apex.

1. Montamans panamensis Dwyer.*^^ type: Panama, Dwyer & Lallathin 8792
(MO, holotype). — Fig. 62.

Succulent herbs to 1!4 m tall, the stems terete, dark green, nodose, glabrous,
to 1.5 cm in diam. Leaves oblong rotund to widely oblong, to 35 cm long, to 20
cm wide, triangular deltoid toward the apex, often round or obtuse at the base,
cuneate, sometimes inequilateral, the costa prominulous above, piano com-
pressed beneath, the lateral veins ca. 25, widely arcuate, often with a vein be-
tween a pair of lateral veins, the intervenal areas spreading reticulate, thinly
succulent, chartaceous when dry, somewhat discolorous, glabrate above and be-
neath except for the veins which are sparsely pubescent beneath; petioles to 5
cm long, to 0.3 cm wide, the scars of the petioles hemispherical; stipules tending
to persist at the tip of the stem, free, thick, 2-awned, with the awns spreading,
triangular, equaling the body, auriculate toward the base. Inflorescences sessile,
axillary or on short peduncles, the heads of flowers to 4.5 cm wide, wider than
long; bracts arranged in 2 series, the outermost bracts to 14 mm long, 3-8 mm
wide, wider toward the apex, polymorphic and often triangular galeate or bifid
for ca. 2 mm; bracteoles linear lanceolate, to 9 mm long, galeate. Flowers with
the hypanthium ca. 1 mm long, often yellow, the calyx lobes 5(-6), unequal, often
narrowly lanceolate and falcate, some compressed galeate toward the apex, to 5
mm long, carnose, minutely diffusely pubescent, the galeate projections to 2 mm
long, briefly connate at the base; corolla white, the tube cylindrical, to 6 mm
long, slightly inflated above the middle, a few short, glandular hairs outside, white
villose within the area of filament attachment, the lobes 5, spreading at anthesis,
triangular, to 5 mm long, acute, each lobe with a galeate process just below the
tip, the process cylindrical, solid, obtuse, reflexed, to 4 mm long, at maturity
shorter than the tube; stamens (4-)5, the anthers oblong, to 1 mm long, the
filaments ca. 1 mm long, attached near the middle of the tube; ovary often split
in the middle, to 1 mm in diam., 2-celled, the septum thick, the ovules 2, subplane,
attached basally, the style cylindrical, to 5.5 mm long, the stigmas 2, suberect,
slightly unequal, 0.8-1.3 mm long, slightly wider than the style; ovarian disc
yellow, annular. Fruits white, rotund, to 6 mm in diam., carnose, smooth, gla-
brous with 10 slender longitudinal veins, the pyrenes 2, piano compressed, the
wall thin; seeds 2.

Montamans panamensis is known only from Panama.

CHiRiQUi: Cerro Horqiieta, B/zva^i & Dwyer 2648 (MO). Rio Chiquero, D'Arcy 10985 (MO), Cerro
Horqueta, Dwyer & Hoyden 771 (MO); Dwyer & Lallathin 8972 (MO); Kirkhride 145 (MO).

^^ Montamans panamensis Dwyer, spec. nov. Herbae succulentae. Folia magna, venis latera-
libus ca. 25; petiolis ad 5 cm longis; stipulis liberis 2 setis pracditis. Inflorescentiac axillares sessiles
vel subsessiles floribus in caespitem ad 4.5 cm latum congestis; bracteis 2-seriatim dispositis poly-
morphis ad 14 mm longis. Flores lobis calycis 5(-6) polymorphis, ad 5 mm longis; corolla tubo ad 6
mm longo, lobis 5, ad 5 mm longis, lobo utroque cornu galealum vix infra apicem ferente; antheris
ad 1 mm longis; ovario bilocellato ovulis 2 basaliter affixis. Fructus aibi rotundi ad 6 mm diam,,

carnosi laeves pyrenis 2 plano-compressis.

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

287

Figure 63. Morinda citrifolia L.— A. Habit (xi/2).— B. Flower opened to show interior (x4'/2).
[After Holdridge 6507.]— C. Fruits (xVi). [After Porter et ai 5016,]

288 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

59. MORINDA

Morinda L., Sp. PI. 176. 1753. lectotype: M. rojoc L.

Shrubs or small trees. Leaves opposite, occasionally ternate, petiolate; stip-
ules interpetiolar, connate, persistent, usually bifid. Inflorescences terminal or
axillary, the peduncles present or absent, the flowers in small heads. Flowers
with hypanthium and calyx of adjacent flowers partially or completely united;
calyx truncate, the teeth obscure; corolla infundibuliform or hypocrateriform,
the lobes 4-7, valvate; stamens 4-7, the anther connectives produced at the apex;
ovary 2-4-celled, each with an ascending ovule attached near the base of the cell.
Fruit a syncarp, often large, fleshy and white, each cell with a 2-celled pyrene.

Morinda is well distributed in the tropical areas of the world. It is a genus of
about 80 species, a few of which are a source of dyestuffs. For more detail on
the structure of the fruits and seeds see Hayden & Dwyer (1969).

Literature:

Hayden, M. V., & J. D. Dwyer. 1969. Seed morphology in the tribe Morindeae.
Bull. Torrey Bot. Club 96: 704-710.

a. Leaves 1.7^.0(-5.0) cm wide 3. M. rojoc

aa. Leaves 5-12 cm wide.

b. Largest leaves exceeding 20 cm in length; corolla lobes more than 6 mm long; fruit

9 cm wide or wider 1. M. citrifolla

bb. Largest leaves less than 20 cm long; corolla lobes 2.5-5.0 mm; fruits 1.0-2.5 cm

wide 2. A/, panamensis

1. Morinda citrifolia L., Sp. PI. 176. 1753. type: Herb. Linn, (LINN. 236.1, not
seen; microfiche MO). — Fig. 63.

Trees or shrubs to 25 ft tall, the branchlets 4-angled, glabrous, the nodes well
spaced. Leaves oblong, 14-30 cm long, 12-18 cm wide, obtuse to acute at the
apex, widely cuneate at the base, often slightly inequilateral, the costa grooved
and subimmersed above, prominulous but appearing subplane beneath, to 0.3 cm
wide proximally, the lateral veins ca. 8, prominulous, arcuate, the median veins
to 5 cm apart, chartaceous to papyraceous, concolorous, glabrous or punctate

above, glabrous beneath except the barbate vein axils; petioles to 2 cm long;
glabrate, slightly alate adaxially; stipules oblong or oblong rotund, 10-20 mm
long, 10-20 mm wide at the base, scarious, glabrous, appressed. Inflorescences
axillary, glabrous, drying black, the peduncles 1-4, stout, to 3 cm long, the floral
masses rotund to oblong, to 2 cm in diam. Flowers sessile, the hypanthium hidden
in the floral mass, the calycine cup short, truncate, the margins scarious, the
lobes evanescent, glabrous; corolla white, the tube cylindrical, ca. 6 mm long,
carnose, glabrous outside, villose inside near the mouth, the lobes 5, narrowly
oblong, to 9 mm long, 1.5 mm wide, obtuse, carnose, glabrous; stamens 5, long
exserted at anthesis, the anthers subversatile, linear oblong, to 5.5 mm long,
twisted and crescentic on shedding pollen, the filaments slightly unequal in length,
inserted at the corolla mouth; stigmas 2, spread at anthesis, plumose in appear-
ance, narrowly oblong, to 5.2 mm long, 0.8 mm wide, the margin erose. Fruits
fleshy, succulent, as large as a man's hand, drying black.

1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceac) 289

The range of Morinda cithfolia is India, Malaya, and tropical Australia; it has
become naturalized in the West Indies and occurs sporadically in Central Amer-
ica. The large succulent white fruit bears a resemblance to large edible fruits of
Annona.

The collection Dwyer & Lallathin 8794 has the following notes: "Flowers
with yellow-green corolla tube, the lobes spreading to slightly reflexed, the lobes
brown on the upper surface; fruits at maturity china white with distinct golden
green 'eyes' (ovarian discs and calycine cups), to 0.8 cm in diam.; stipules green,

is used medicinally by the Cuna of Darien.

M. citrifi

BOCAS DEL TORO: Isia Colon, Chiriquf Lagoon, Wedel 2942 (F, G). canal zone: Boyd Roosevelt
Highway, 5 mi W of Sabanita, Croat 14074 (MO). Between Gatiin and Fort Sherman, Croat 15403
(MO). Summit Garden, Croat 16645 (MO). Coco Solo, Dnyer d Duke 7906 (COL, MO, UC, VEN).
Fort Clayton, Dwyer & Lallathin 8794 (MO). Mouth of Rio Chagres, Johnston 1771 (MO). Summit
Garden, Nee 7046 (MO). Fort San Lorenzo, Porter et al. 5016 (MO), colon: Portobelo, Croat 14127
(MO). Man'a Chiquita, Dwyer 5018 (MO). Saliid, Holdridge 6507 (MO). Quebrada Santa Marta, coast
road AVi km SW of Pina, Nee 11701 (MO), san blas: Ailigandi River, Duke & Bristan 333 (MO).

2. Morinda panamensis Seem., Bot. Voy. Herald 136. 1854. type: Panama, Fen-
dler 148 (MO. isotype).

Trees or shrubs to 25 ft tall, the branchlets 4-angled, farinose puberulent or
glabrescent, the nodes well spaced. Leaves oblong, ovate oblong or oblong ro-
tund, 9-20 cm long, 5-15 cm wide, deltoid or rounded at the apex, shortly acu-
minate, the acumen to 1 cm long, basally cuneate, often attenuately so and equi-
lateral, the costa slender above, plane, prominulous or subimmersed above, the
secondary veins 6-9, arcuate, to 3(-4) cm apart, chartaceous, glabrous or farinose
above, pubescent or farinose beneath, rarely totally velutinous pubescent, the
hairs often restricted to the axils and costa; petioles to 2.5 cm long, occasionally
absent, farinose puberulent; stipules deltoid or oblong rotund, 6-10 mm long, 6-
9 mm wide, often wider than long, the peduncles ascending, 1.5-5.5 cm long,
1,5-5.0 mm wide, glabrescent, often farinose, terminated by 3-4 equal or unequal
branches, to 3 cm long, each with a subrotund capitulum, 0.5-1.0 cm wide.
Flowers sessile; hypanthium subrotund, truncate, ca. 1.8 mm long, minutely pu-
berulent, the calycine tube to 1 mm long, the teeth absent or reduced to points;
corolla white, the tube cylindrical, 7-8 mm long, thickly petaloid, minutely pu-
berulent outside, villose inside above the middle, the lobes 4, valvate, thick,
triangular in cross section, narrowly oblong, 2.5-5.0 mm long, tapering narrowly
toward the apex, vaguely to obviously cordate at the base; anthers narrowly
oblong, ca. 2.5 mm long, obtuse, the filaments inserted near the mouth; stigmas
2, digitiform, 3.5-5.0 mm long, erect, the ovary rectangular in cross section, the
wall moderately thick, the septum thin, 2-loculate, the ovules one per cell. Fruits
disposed in a capitulum, to 2.5 cm in diam.

Morinda panamensis ranges from tropical Florida to Mexico, Central Amer-
ica, and the West Indies.

CANAL zone: Fort Lorenzo, Haxden 101 (MO); Porter et al 4985 (MO, SCZ. UC, VEN), 5016
(COL, DUKE, MO, SCZ, UC, VEN), 5017 (COL, DUKE, MO, NY, SCZ, UC, VEN); Tyson &

290 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

Blum 3692 (MO). Gatiin Locks to Fort Sherman, Tyson 3756 (MO). Chagres River Landing, near Fort
San Lorenzo, Tyson & Nee 7310 (MO), cocle: Cerro Pilon, El Valle, Duke & Dwyer 13894 (MO).
colon: Salud, Holdridge 6507 (MO). Salud and Boca de Rio Indio, Howell 84 (MO). Palenque, Piitier
4121 (F). darifn: El Real to Pinogana Duke 5000 (MO). Yaviza near Rio Chucunaqiie, El Punteadero,
Stern el al. 171 (MO). Panama: Chagres, Fendler 148 (MO, US). 16-18 km from the Panamerican
Highway, El Llano-Carti Road, Tyson & Nee 7321 (MO), san blas: Mulatuppo, Duke 8510 (MO).
Mandinga, Duke 8924 (MO).

3. Morinda rojoc L. Sp. PI. 176. 1753. type: not seen.

M. yucatanensis Greenm. Publ. Field Columbian Mus., Bot. Ser. 2. 262. 1907. type: Mexico, Gaumer
362 (F, holotype).

Shrubs, often scandent, to 2 m tall; branchlets angular, smooth, glabrous,
cracking, often drying tan, the nodes well spaced. Leaves oblong, obovate oblong
or narrowly obovate oblong, often falcate, 4.0-15.5 cm long, 1.7-4.0(-5.0) cm
wide, usually 4 times longer than wide, acute or attenuate acute at the base, the
costa slender and prominulous above, prominent beneath, the lateral veins 5-7,
strongly arcuate, prominulous, the intervenal areas smooth, the veinlets usually
evanescent, papyraceous, glabrous above but minutely farinose or punctate, bar-
bate along the costa, often farinose or punctate beneath, the margin slightly in-
volute; petioles to 0.5 cm long, slender, ca. 1 mm wide, glabrous; stipules tri-
angular, to 4 mm long, ca. 5 mm wide at the base, the median awn subulate,
about the same length as the body. Inflorescences axillary, solitary, the peduncle
to 0.3 cm long, stout, puberulent, the terminal head of flowers ca. 0.5 cm in diam.
Flowers with the hypanthium cupuliform, ca. 1 mm long, the calycine cup ca. 0.35
mm long, glabrous, the margin undulate, the lobes 5, indistinct, obtuse, puberu-
lent; corolla white, the tube cylindrical, ca. 5 mm long, ca. 2 mm wide, carnose,
glabrous outside, densely white villose inside above the middle, the lobes as long as
the tube; stamens 5-6, the anthers ca. 2 mm long, obtuse, the filaments slender;

stigmas 2, ca. 1.5 mm long, the style ca. 3 mm long. Fruits rotund, 1-2 cm in
diam.

Morinda rojoc occurs in Florida, Mexico, Central America, the West Indies,
and northern South America, it is interesting that all of the Panama collections
of the species were made on or near beaches.

colon: Guaira (mainland opposite Isla Grande), d^Arcy 4053 (MO), san blas: Mulatuppu (Rfo
IbedO, Duke 8510 (MO). Airport at Irandi, Duke 6511 (MO). Mulatuppu, Duke 8520 (MO). Isla de
Soskatupu, Duke 8966 (MO). Nargana Island, Gentry 1514 (MO). Isla Ustupo, W of Playon Chico,
Gentry 6338 (MO). Isla de Soskatupu, Kirkbride 178, I87A (both MO). Nargana, Tyson & Dwyer
1173 (MO).

60. NERTERA

Nertera Banks & Soland. ex Gaertn., Fruct. Sem. PI. 1: 124. 1788, nom. cons.
type: N. dcpressa Gaertn. = A^. granadensis (L.f.) Druce.

Gomozia Mutis ex L.f., Suppl. PI. 17, 129. 1781, nom, rejic. type: G. granadensis L.f.
Gomosia Lam., Encycl. Meth. Bot. 2: 769. 1788, Gomozia Mutis, ex L.f. orth. mut.
Gomoza Cothen., Disp. Veg. Meth. 5. 1790, Gomozia Mutis, ex L.f. orth. mut.

Creeping, perennial herbs, often forming mats. Leaves small, often subsessile;
stipules minute, adnate to the petioles, the sheath entire or with 2 teeth. Flowers

1980]

DWYER— FLORA OF PANAMA (Family 179. Rubiaceae)

291

Figure 64. Nertcra ^ranadensis (L. f.) Druce.— A. Habit (xVi).— B. Fruit (x6). [After Croat
13778. \ — C, Flower opened to show interior (x 10). — D. Ovary longitudinal section (x9). [After Croat
620, Costa Rica. J

minute, axillary, sessile, usually perfect; calyx truncate or tubular, the teeth 4-
5; corolla tubular or funnel shaped, glabrous within, the lobes 4, valvate; stamens
4, attached basally to the corolla, the anthers apiculate, exserted; ovary 2-celled,
the style long exserted, hairy, the ovules solitary, erect from the base of the cell.
Fruits rotund, succulent, red, with 2 cartilaginous nutlets.

Nertcra is distributed in Mexico through Central America, South America,
and the West Indies, It also occurs in Australia, New Zealand, and the East
Indies, as well as in Hawaii.

292 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

1. Nertera granadensis (L.f.) Druce, Bot. Soc. Exch. Club Brit. Isles. 1916: 637.

1917.— Fig. 64.

Gomozia ^ramidcnsis Mutis ex L.f., Suppl. PI. 129. I78L

Nertera dcpressa Banks & Sol. ex Gaertn., Fruct. Sem. PI. 1: 124. 1788. type: Antartica, Banks (K,

holotype), not seen.

Herbs, epiphytic or creeping, occasionally to 1.3 m tall, the branches short,

glabrous or webby pubescent (mouldy?), often drying deep purplish red. Leaves
ovate rotund, to 1.3 cm long, to 1.25 cm wide, slightly inequilateral, widely deltoid
at the apex, ultimately retuse or minutely apiculate, basally subcordate to trun-
cate, the base deltoid as it passes into the petiole, the costa plane or evanescent
above (then often pubescent), plane to prominulous beneath, the lateral veins 2-
5, often evanescent, the 2 basal veins together with the costa appearing flabellate,
widely arcuate, carnose but drying thin, glabrous or pubescent, the hairs thick,
falcate, appressed, often only the hair bases persistent and the surface then silvery
lepidote, the margin delicately marginate; petioles 0.9 cm long, to 0.3 cm wide,
glabrous, often grooved above; stipules interpetiolar, triangular, to 1 mm long,
to 1 mm wide. Flowers solitary; hypanthium oblong, ca. I mm long, with a few
scattered hairs, the calycine cup short and without lobes; corolla yellow green or
pale yellow, the tube widely funnel shaped, inflated, ca. 0.8 mm long, glabrous,
petaloid; stamens 4, the anthers rotund, ca. 0.3 mm in diam., the filaments ca,
1.5 mm long, inserted at the base of the corolla tube; stigmas 2, reflexed, subulate,
ca. 1 mm long, longer than the style, the ovule 1, large, pendent. Fruits sessile,
scarlet at maturity, oblong, to 2.5 mm long, to 2 mm wide, glabrous, delicately
reticulate and with a few delicate longitudinal lines, the raphides abundant.

Nertera granadensis ranges from Mexico to Chile. Index Kewensis lists the
species as occurring in New Zealand and the Island of Tristan da Cunha.

CHiRiQuf: Cerro Horqueta, Blutn & Dwyer 2613 (MO). Las Cumbres, Cerro Punta. Croat 13778

(MO); Croat <S Porter 16117 (MO). Alto Respinga and above, ca. 2800 m, D'Arcy 9958 (MO). Above

Bajo Chorro, DArcy 1 1092 (MO). Bajo Chorro, Boquele, Davidson 422 (F, MO). Potrero Muleto,
Volcan de Chiriqiii, Boquete, Davidson 1038 (F, MO). Between Quebrada Hondo and divide on

Calderas-Chiriquicito trail, Kirkhride & Duke 930 (MO). Las Nubes, W of Cerro Punta, Liesner 291

(MO). E side of Volcan Baru, 3000 m. Mori & Bolten 7437 (MO). Bajo Grande, ca. 3 km E of town

of Cerro Punl^a, 2200 m, Nee 9977 (MO). Cerro Punta, 6800 ft, Ridgway & Solis 2397 (MO). Rio

Chiriquf Viejo Valley, near Volcan, White 170 (MO). Ri'o Chiriquf, Viejo Valley, White 219 (F). Loma

Larga, Volcan de Chiriquf, Woodson et al. 1032 (F, MO), darien: Cerro Tarcarcuna, W ridge along
Quebrada toward waterfall, 1500-1550 m. Gentry & Mori 14104 (MO).

61. OLDENLANDIA
Oldenlandia L., Sp. PI. 119. 1753; Gen. PI., ed. 5: 55. 1754. lectotype: O.

corynibosa L.

Herbs or rarely subshrubs. Leaves callose mucronate at the apex; stipular
sheath adnate to the petiole. Inflorescences terminal or pseudo axillary, flowers
solitary at the nodes or clustered. Flowers 4-merous, isostylous or heterostylous,
the calycine cup dentate; corolla tube often short, often barbate at the mouth;
stamens inserted at the base of the corolla lobes or well into the tube, the anthers
dorsifixed; style glabrous or pubescent, the stigmas 2, subglobose or elongate,
the ovary 2-celled, the ovules numerous on a peltate placenta. Fruits capsular,

1980]

DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae)

293

c

Figure 65. Oldenlandia corymbosa L. — A. Habit (xVi). [After Lewis et al. 781.] — B. Fruiting
calyx (xAVi). — C. Seeds (x4Vi). [After Fendler W.\ — D. Flower, part of the perianth removed
(x4Vi). — E. Flowering calyx (x4Vi), [After Lewis 41,]

294 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

the wall horny, sometimes produced within the calyx into a beak and there de-
hiscing loculicidally by a cleft; seeds usually numerous, subglobose or angular,
smooth or alveolate.

Oldenlandia occurs throughout warmer areas of the world. It is a genus of
about 300 species.

a. Leaf blades to 0.3 cm long _.- 1. O. calliirichoicles

aa. Leaf blades 1.5-5.5 cm long.

b. Leaves with lateral veins evanescent; flowers 1-5 2. O. corymbosa

bb. Leaves with lateral veins visible; flowers solitary 3. O. lancifolia

1. Oldenlandia callitrichoides Griseb., Mem. Am. Acad. n.s. 8: 506. 1863. type:
Cuba, Wright 268, 1640, not seen.

Prostrate herbs disposed as a mat, or erect to 10 cm tall, the stems smooth,
glabrous, to 0.3 mm wide, branching profusely, the nodes usually well spaced.
Leaves with a spathulate appearance because of elongate petioles, the blades
ovate, rhomboid ovate to oblong, to 3 mm long, to 2 mm wide, acute or obtuse
at the apex, cuneate, obtuse or rounded at the base, the lateral veins delicate, 3-
4, often evanescent, membranous, concolorous, drying green, glabrous; petioles
shorter than or about as long as the blade; stipules adnate to the petiole base,
often with a few dark red glands on the surface or on the margin, often with a
narrowly oblong deciduous appendage to 0.5 mm long. Flowers not seen but
apparently solitary in the axils. Fruits with pedicels to 3 mm long, the body ca.
3 mm long (including calyx), truncate, smooth, glabrous except on teeth, raphides
abundant, the persistent calycine lobes apparently 5, triangular oblong, to 0.6 mm
long,

Oldenlandia callitrichoides occurs sporadically in the tropics of the New
World. This species has the smallest leaves of all the Panamanian Rubiaceae.

colon: Between Garotte and La Guiara, DArcy 9378 (MO). Panama: On stone wall near side-
walk, Calle EstLidiante, Panama City, DArcy I0190' (MO),

2. Oldenlandia corymbose L., Sp. PI. 1 19. 1753. type: not seen; cited by Linnaeus
as in *'Trop. Meridionalis.''— Fig. 65.

Herbs, prostrate or erect, to Vs m tall, sparsely to much branched, the branch-
es lax, glabrous, shiny. Leaves narrowly oblong or linear, 1.5-2.0(-3.0) cm long,
to 0.45 cm wide, deltoid to acute at the apex, cuneate at the base, the costa
evanescent above or barely visible, scarcely prominulous beneath, the lateral
veins evanescent, the margin scabridulous, with some proximal papillae, thinly
carnose, concolorous, glabrate beneath; petioles to 1 mm long, inserted in the
sheath of the connate stipules; stipules cupuliform or obconical, to 2.5 mm long,
the cup with several ascending or deflexed glandular fimbriae of unequal length,
often exceeding the sheath in length. Inflorescences mostly at the nodes, the
flowers solitary or often 2 or 3, infrequently disposed in corymbs of 4-5 flowers,
the peduncles filiform, usually about Vi as long as the leaves, the pedicels fihform,
shorter or longer than the peduncle. Flowers isostylous with the hypanthium
small, the calycine lobes 4, triangular, to 1.1 mm long, scabrid marginate; corolla
lavender or white, the tube ca. 0.6 mm long, the lobes 1-2 mm long; stamens 4,

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 295

the anthers sessile, rotund, inserted on upper V2 of the tube, the pistils glabrous.
Fruits capsular, to 2.5 mm wide, truncate at the apex, subplane; seeds 1 per
carpel, round, ca. 0.5 mm wide, areolate.

Oldenlandia corymhosa is native to Africa, occurring throughout the conti-
nent. It was probably introduced into the New World in the 15th century and has
spread throughout tropical America.

BOCAS DEL TORo: Bocas del Toro Island, Lewis et al. 781 (GH, K, MO, US, UC). Changuinola,
Lewis et aL 796 (GH, MO), 804 (GH, MO, UC, US), canal zone: Frijoles, Croat 6077 (MO). E of
Gatiin Locks, Duke 4297 (MO). Tore Point, Fort Sherman, Duke 4342 (MO). Chagres, Fendler 10
(MO). Madden Dam, Lewis et aL 5365 (COL, MO, SCZ, UC, VEN). Road 3 km SE of bridge,
Gamboa, Nee 9434 (MO). Road 2 km NW of Summit Gardens, 80 m, Nee 9431 (MO). Balboa,
Standley 25660 (MO). Summit, Standley 26970 (MO), barro Colorado island: Croat 6800, 7055
(both MO); Ehinger39 (MO); McDaniel'5024 (MO), cocle: El Valle, Lewis et al 2507 (COL, DUKE,
MO, UC). colon: Mouth of Rio Piedras, Lewis et aL 3170 (COL, DUKE, MO, UC, VEN). Panama:
2 km W of El Llano, Nee 7961 (MO).

3. Oldenlandia lancifolia (Schum.) DC, Prodr. 4: 425. 1830.

Hedyotis lancifolia Schum. in Schum. & Thonn., Beskr. Guin. PI. 72. 1827. type: not seen.

Hedvotis longifolia Schum. in Schum. & Thonn., Bcskr, Guin. PI. 70. 1827. type: not seen.
Oldenlandia longifolia (Schum.) DC, Prodr. 4: 426. 1830.

Herbs, the stems slender, weak, glabrous to puberulent, smooth, the nodes
well spaced, to 4.5 cm apart. Leaves sessile or subsessile, narrowly elliptic, often
slightly inequilateral and curved, 1.5-5.5 cm long, 0.2-0.7 cm wide, attenuate
acute at the apex and the base, the costa immersed or scarcely prominulous
above, prominulous beneath, the lateral veins ca. 4, slender, strongly arcuate
ascending, evanescent, membranous, smooth, minutely and sparsely puberulent
above and beneath, the hairs strigose; stipules subulate and bifid, ca. 3 mm long,
reflexed. Flowers not seen. Fruits subrotund, didymous, to 3 mm long, glabrous,

the raphides numerous, the calycine cup short, the teeth 4, well spaced, acute,
to 0.3 mm long; seeds numerous, ca. 0.2 mm long.

Oldenlandia lancifolia occurs in the tropics of the Old and the New World.
Steyermark (1974) describes the flowers: ''Flowers solitary at the nodes, axillary
or pseudo axillary, rarely 2, pedicellate; pedicels filiform, 0.5-3.0 cm long. Lobes
of the calyx lanceolate or ovate triangular, acute to acuminate, 1.0-1.5 mm long,
glabrous or scabridulous on the margins. Corolla white or rarely tinted with lav-
ender or rose, hypocrateriform; tube 1 mm long, glabrous on the front and on the
back; lobes equal or larger than the tube, 1-2 mm long."

w

BOCAS DEL TORo: Changuinola, banana drainage ditch. D'Arcy II 171 (MO), canal zone: Low-
land E of Gatun Locks, burned about 1957, Duke 4297 (MO). Railroad 2 km NW of Summit Gardens,
80 m. Nee 9431 (MO).

63. PALICOUREA
Palicourea AubL, Hist. PI. Guiane Fr. 1: 172. 1775. type: P. guianensis Aublet.

Palicurea Schultes in R. & S., Syst. Veg. 5: 11, 193. 1819. Palicourea Aubl. orth. mut.

Shrubs or small trees. Leaves mostly opposite, occasionally ternate or qua-
ternate, petiolate; stipules interpetiolar, usually persistent, usually connate, each
fused part of the sheath with 2 lobes or cusps, ascending and often varying in

296 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

length. Inflorescences terminal, rarely axillary, often paniculate, the flowers dis-
posed in racemose, corymbose or thrysoid cymes. Flowers with the perianth and
stamens usually 5-merous; hypanthium short, the teeth reduced or absent; corolla
often orange, yellow or purple, the tube cylindrical, usually curved or dilated
basally, the lobes short, valvate; anthers the same number as the corolla lobes;
ovary 2-celled, the cells 1-ovulate, the ovules arising from the base, the septum
median, thick. Fruits fleshy, composed of 2 pyrenes.

Palicourea, a genus of about 200 species, is common in the tropics of Middle
America and South America. It also occurs in the West Indies. Standley's (1938)
key to the Costa Rican species, which covers 13 of the 25 species in Panama, is
noteworthy.

Literature:

Bremekamp, C. E. B. 1934. Palicourea. Rec. Trav. Bot. Neerl. 31: 283.
Standley, P. C. 1938. Palicourea (Flora Costa Rica). Field Mus. Nat. Hist., Bot.

Ser. 18(4): 1330-1338.
& L. O. Williams. 1975. Palicourea in Flora Guatemala. Fieldiana Bot-

any 24: 130-133.

\. Fruits less than 1 cm long; leaves chartaceous or membranous.
2. Calycine lobes (at flowering) foliosc or imbricate.

3. Calycine lobes foliose 2. P, hella

3\ Calycine lobes imbricate 15. P. ochnoides

T . Calycine lobes (at flowering) not foliose or imbricate.

4. Leaves 3 per node 20. P. triphylla

4'. Leaves 2 per node.

5. Inflorescence corymbiform, usually flat topped, the branches strongly ascending

6. P. fastigiata

5'. Inflorescences not corymbiform or flat topped, the branches not strongly ascend-
ing.

6. Corolla obliquely gibbous at the base 8. P. glhhosa

6'. Corolla not obliquely gibbous at the base.
7, Calycine lobes more than 2.5 mm long.

8. Leaves with lateral veins 15-20.

9. Leaves 4-13 cm long, the lateral veins ca. 20; largest bracts

with a median subulate process at the apex and 2 short lateral

processes 12. P. macrosepala

9'. Leaves 18-21 cm long, the lateral veins 15-18; largest bracts

acute at the apex 9. P. grandibracteata

8'. Leaves with lateral veins 11-15 17. P. pauciflora

7'. Calycine lobes less than 2.5 mm long.

10. Leaves with the lateral veins ca. 30; inflorescence bright yellow

(fresh and dried) 24. P. vestita

10'. Leaves with lateral veins 8-21; inflorescence not bright yellow.
11. Leaves with lateral veins 7-10.

12. Flowers purple, lavender or rose L P. angustifoUa

\T , Flowers green, yellow or white.

13. Leaves pubescent beneath; corolla tube 10-15 mm

long 7. P- galeoUiana

13'. Leaves glabrous beneath; corolla tube to 8 mm long.

14. Flowers white; corolla tube 3-4 mm long

___. 4. P, chiriqidna

14'. Flowers colored; corolla tube to 8 mm long

14. P. montivaga

11 '. Leaves with lateral veins 11-21.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaccae) 297

15. Corolla ca. 40 mm long, webby pubescent outside

_ 18. P. rigidifolia

15'. Corolla to 20 mm long, not webby pubescent outside.
16. Leaves glabrous beneath.

17. Stipular sheath rounded at the apex, scarcely

retuse 22. P. tumidonodosa

17'. Stipular sheath with 2 distinct awns or lobes.

18. Calycine lobes obtuse; corolla 10-20 mm

long, pubescent outside; anthers 3-5 mm

long 10. P. guianensis

18'. Calycine lobes acute to subulate; corolla
4.5-7.0 mm long, glabrous outside; an-
thers less than 3 mm long.
19. Inflorescences with the upper branch-
es spreading; leaves with the lateral
veins ca. 10; corolla green or yellow

14. P. montivaga

19'. Inflorescences with the upper branch-
es contracted; leaves with the lateral
veins I2(-14); corolla red ___

5. P. crocea

16'. Leaves variously pubescent beneath.
20. Stipules with awns.

21. Leaves with lateral veins ca. 20.

22. Flowers yellow 24. P. vestita

22'. Flowers purple, lavender or magenta

_ 16. P. panamcnsis

21'. Leaves with lateral veins 10-15.

23. Fruits turbinate. _ 23. P. veraguensis

23'. Fruits not turbinate.

24. Inflorescence pink; corolla gla-
brous outside 19. P. suhruhra

24'. Inflorescence not pink; corolla

pubescent outside.

25. Leaves narrowly elongate

lanceolate; corolla puber-

ulent, purple, lavender or

L J ^' 1

26. Leaves 6-1 1 cm wide

16. P. panamcnsis

26'. Leaves to 2 cm wide

L P. angiistifolia

25'. Leaves elliptic oblong; co-
rolla hirtellous or villous.
yellow

— W. P. lasiorrachis

20'. Stipules without awns, the lobes ovate-rotund

13. P. montensis

r. Fruits 1.1-1.5 cm long: leaves coriaceous or subcoriaceous

_ 3, />, carpirostrata

27. Leaves acute at apex, drying dark brown, the lateral veins ca. 16 3. P. carpirostrata

27'. Leaves rounded at apex, drying yellowish, the lateral veins 7-10 21. P, tuhtdiflora

1. Palicourea angustifolia H.B.K., Gen. Nov. Sp. PI. 3: 367. 1819. type: Banks
of Orinoco and Rio Negro, Amazonas, Venezuela, not seen.

P. lanceolata Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kj0benhavn 17. 1852. type: Costa
Rica, Oersted 11556 (F, photo of type).

P. intermedia Oerst. ex Polak., Linnaea 41: 571. 1877. type: Costa Rica, Polakowsky 143 (F, photo
of type).

P. moritzii Wernh., J. Bot. 55: 283. 1917. type: Colombia, Moritz 844, not seen.

298 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

Shrubs or trees to 10 m tall, the branchlets terete, smooth, the pith often
conspicuous, white, often constricted below the nodes, the latter well spaced.
Leaves narrowly elliptic, occasionally falcate, 4-20 cm long, 1.5-4.0 cm wide,
acute or obtuse at the apex, acuminate, the acumen to 2.5 cm long, often falcate,
ultimately obtuse or acute, cuneate, occasionally obtuse at the base, the costa
prominulous above and beneath, the lateral veins (7-) 10-14, arcuate, with usually

1 evanescent parallel vein between 2 lateral veins, papyraceous or chartaceous,
glabrous above, puberulent beneath especially on the costa and the veins; petioles
to 2.5(-3.0) cm long, 0.08-0.10 cm wide, glabrous; stipules connate, 3.5-7.5(-9)
mm long, the sheath cylindrical, 2-3 mm long, truncate, glabrous, each part with

2 subulate erect awns, mostly as long as or 2-3 times longer than the sheath.
Inflorescences terminal, solitary, narrowly oblong, narrowly pyramidal or spici-
form, thrysoid paniculate, 6-16 cm long, 2.5-5.5(-8.0) cm wide, the axes purple
or red, the peduncles 2.5-7.0 cm long, to 0.2 cm wide, angular, glabrous, the
lowermost branches well spaced, opposite, alternate or ternate, to 3.5 cm long;
bracteoles minute, subulate, to 2 mm long. Flowers pedicellate; hypanthium and
calycine cup short, ca. 1 mm long, the calycine lobes 5, spreading, to 0.5 mm
long; corolla pale lavender, purple or rose colored, the tube cylindrical, 8-14 mm
long, 0.4 mm wide, petaloid, glabrous outside, glabrous on the upper V2, within,
the lower V3 pubescent, the lobes 5, oblong rotund to oblong, ca. 2.5 mm long,
obtuse; stamens 5, the anthers narrowly oblong, ca. 4 mm long, the filaments
subplane, 1-2 mm long, attached just above the middle of the tube, style as long
as the corolla tube, the stigmas slender, ca. 0.7 mm long. Fruits subrotund, 4-5
mm long, purple black or dark blue, drying black, glabrous, moderately furrowed
longitudinally.

Palicourea angustifolia extends from Costa Rica south to Peru. It is restricted
in Panama to elevations above 2,000 ft. It is frequently confused with P. galeot-
tiana Martens; the latter, however, has a yellow or orange corolla while P. an-
gustifolia has a lavender to purple corolla. Folsom & Collins 1792 has much
reduced leaves, the largest blades to 8 cm long, to 2.5 cm wide.

BOCAS DEL TORo: Robalo Trail, N slope of Cerro Horqueta, 2250-2500 m, Allen 4929 (MO, NY).

CHiRiQUi: Cerro Punta to Rio Caldera, Allen 1438 (MO, NY, US). Cerro Horqueta, /UUU it, wiim <k
Dwver 2663 (MO). Las Nubes, W. of Cerro Punta, 2200 m, Busey 667 (MO). Las Cumbres, Cerro
Punta, Croat 13717, 13764 (both MO). Monte Rey, Boquete, Croat 15852 (MO). Las Cumbres, Cerro
Punta, Croat & Porter 16072 (MO). Bajo Chorro. Boquete, Davidson 111 (MO, US). Cerro Horqueta,
5000-5800 ft, Dwyer et al. 542 (GH, K, MO, UC, US). Boquete, Dwyer 7010 (MO). Cerro Colorado,
1500 m, Folsom & Collins 1792 (MO). Boquete, Hagen & Hagen 2020 (NY). Cerro Horqueta, 6500
ft, Hagen & Hagen 2051 (MO, NY). Boquete, Hagen & Hagen 2056 (NY). Camp Aguacatal, Pittier
3113 (US). Cerro Vaca, Pittier 5332 (US). Upper Rio Chiriquf Viejo, 1300-1900 m, Seibert 299 (MO).
Finca Collins, Boquete, Stern et at. 1094 (MO, US), 2014. 2054 (both MO). Bambito, 1 mi SW of
Cerro Punta, 5600 ft, Tyson 5685 (MO). Cerro Punta, White 29 (MO). Rio Chiriquf Viejo Valley,
White 36 (MO). Finca Lerida, Woodson & Scherv 215 (MO). Quebrada Velo, 1800 m. Woodson &
Schery 266 (MO). Casita Alta to Cerro Cope. 2300-3300 m, Woodson & Schery 369 (MO). Potrero
Muleto, Volcan de Chiriqui, 3500-4000 m, Woodson & Schery 460 (MO). Casita Alta, Volcan de
Chiriquf, Woodson et al. 881 (MO, NY), cocle: Cerro Pilon, 3000 ft, Duke & Lallathin 14998 (MO);
Dnyer 8338 (MO); Kirkhride 1060 (MO); Lallathin 441 (MO), veraguas: 3.9-5 mi N of Santa Fe,
500-1000 m. Gentry 2948 (MO). Alto Piedra, Santa Fe, Lao 516 (MO). Cerro Tute, 750-1000 m, Mori
6249 (MO). N of Santa Fe, Escuela Agricola Alto de Piedra, Mori & Kallunki 2542 (MO). NW of
Santa Fe, 8.8 km from Escuela Agricola Alto de Piedra, Mori et al. 3899 (MO). Cerro Tute, 750-1000
m, Mori et al. 7550 (MO).

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 299

2. Palicourea bella (Standley) Dwyer, comb. nov.

Psychotria bella Standley, J. Wash. Acad. Sci. 18: 185. 1928. type: Panama, Pittier 3250 (US,
holotype; F, isolype).

Shrubs, the twigs 4-angled, stout, golden puberulent, the nodes well spaced
except at the tips. Leaves elliptic, to 15 cm long, to 6.6 cm wide, deltoid at the
base, the costa prominent beneath, the lateral veins ca. 22, arcuate, drying green,
scarcely discolorous, white strigillose above, prominent beneath; petioles mod-
erately slender, to 3 cm long, pilose; stipules connate to 1.5 cm long, bilobate,
the lobes oblong, rotund at the apex, about Vi the length of the body or less.
Inflorescences terminal, to 12 cm long, the cymules densely aggregated at the
ends of the peduncles, the latter about 7 cm long, the primary branches ca, 2,5
mm long, ca. 5-flowered; bracts oblong, to 1.7 cm long, acute, hirtellous. Flowers
with pedicels 4-6 mm long, pilose; hypanthium short, pilose, the calycine cup
short, cupuliform, pubescent outside with short erect hairs and 5 glandular areas
within, each area covered by an albuminous material and containing 6-12 oblong
glands, the calycine lobes foliose, ovate to ovate elliptic, 11-14 mm long, acute,
venose, petaloid, pink when dry; corolla white, cylindrical, 16 mm or longer,
dilated at base, carnose, puberulent outside, villose within except at filament
attachment, the lobes 5, ovate oblong, 4.5-5.0 mm long, obtuse; stamens 5, the
anthers oblong, ca. 3 mm long, the filaments short, 0.5 mm wide, attached above
the middle of the tube; ovarian disc compressed rotund, to 2 mm long, topped by
a doughnut shaped ring, the style slender, 4-10 mm long, the stigmas slender,
fingerlike, ca. 2.5 mm long. Fruits not seen.

Palicourea bella is known only from Panama. The corolla tube being gibbous
at the base with the trichomes restricted basally suggests Palicourea rather than
Psychotria. The foliose calycine lobes also point to Palicourea. Pittier notes that
the peduncle, bracts, and calyx are pink, another feature suggesting placement
in Palicourea. Of great interest are the slimy patches of glands within the calycine
cup.

CHiRiQuf: Alto de Las Palmas, Cerro Horqueta, 2268 m, Pittier 3250 (F, US).

3. Palicourea carpirostrata Dwyer. ^^ type: Panama, Mori 5262 (MO, holotype).

Shrubs to 3 m, the twiglets stiff, drying deep red, glabrous, glandular shiny.
Leaves lanceolate, 7.0-14.5 cm long, 2-4 cm wide, acute, acuminate, the acumen
1.0-1.5 cm long, usually falcate, ultimately obtuse, acute and scarcely inequilat-
eral at the base, the costa plane or immersed above, prominulous distally, prom-
inent proximally beneath, to 1.3 mm wide proximally, rubescent, the lateral veins
strongly arcuate, ca. 16, rubescent, with 1-2 smaller veins between 2 lateral

'^'^ Palicourea carpirostrata Dwyer, spec. nov. Frutices ramulis glabris. Folia lanceolata, 7.0-
14.5 cm longa, 2-4 cm lata, venis lateralibus forte arcuatis coriacea in costa venisque pubescentia;
petiolis ad 3 cm longis. Inflorescentiae ad 15 cm longae, ad 11 cm latae, pedunculo ad 6 cm longo,
jugis 3 ramorum oppositorum praeditae ramis inferioribus ad 6 cm longis, bracteis inferioribus angusto-
oblongis, ad 1.5 cm longis, acutis vix auriculatis basi. Fructus oblongi vel rotundi, 1.0-1.3 cm longi,
in sicco nigri glabri calyce ad 5 mm longo.

300 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

veins, the larger one subparalleling the laterals and zigzagging toward the margin,
the other veinlets pinnatiform, the margin delicately revolute, coriaceous, dis-
colorous, drying dark brown, hirtellous on the costa and the veins beneath, the
intervenal areas puberulent; petioles stiff, divergent or ascending, to 3 cm long,
drying red; stipules not seen. Inflorescences terminal, solitary, pyramidal panic-
ulate, to 15 cm long, to II cm wide, the peduncle to 6 cm long, to 0.2 cm wide,
glabrous, the rachis with 3 pairs of opposite branches, arcuate or angular as-
cending, well separated, the lowermost to 6 cm long, the secondary branches
usually with 3 branches above the middle, these alternate and umbelloid; bracts
persistent, conspicuous, elongate, narrowly oblong, those at the base of the low-
ermost branches to 1.5 cm long, acute, concave, somewhat auriculate at the base,
the margin with 2-3 minute processes, 0,1-0.2 mm long, the bracts of the upper
primary and secondary branches about as long but subplane, to 3 mm long.
Flowers not seen, but judging from the fruits in 2 unilaterally disposed cymes of
2-3 flowers. Fruits oblong or rotund, 1.0-1.3 cm long, black, glabrous, delicately
ribbed, the calycine cup and teeth rostrate, to 5 mm long, the cup scarious,
glabrous, venose, the 5 teeth slightly longer than the cup, triangular, to 3 mm
long, to 0.4 mm wide, glabrous; pedicels slender, glabrous.

Palicourea carpirostrata is known only from Panama. Its specific name is
based on the elongate persistent calyx of the fruit,

VERAGUAS: Cerro Tute, ridge lop, cloud forest, Mori 5262 (MO).

4. Palicourea chiriquina Standley, Contr. U.S. Natl. Herb. 18: 140. 1916. type:
Panama, Pittier 3118 (US, holotype).

Shrubs to 3 m tall, the nodal scars circular, prominent. Leaves elliptic, to 7.2
cm long, to 2.2 cm wide, acute at the apex, the acumen ca. 0.5 cm long, cuneate
or acute at the base, the costa subplane beneath, the lateral veins 8-9, arcuate,
reaching to the margin, drying somewhat red, the margin delicately revolute,
chartaceous, drying red black above, brown beneath, glabrous, minutely punctate
or rugulose above; petioles slender, ca. 0.5 cm long; stipules connate, to 3 mm
long, each part 2-lobed, the lobes linear, shorter than the sheath. Inflorescences
terminal, cymose paniculate, to 6 cm long, ca. 5 cm wide, the peduncle ca. Vi the
length of the inflorescence, terminated by 3 spreading branches terminated by
loose clusters of radiately disposed flowers; bracteoles ca. 1 mm long. Flowers
with the hypanthium short, the calycine cup ca. 1 mm long, the teeth evanescent,
glabrous; corolla white, 5-6 mm long, the tube cylindrical, inflated, the lobes
oblong, ca. 2 mm long, obtuse. Fruits not seen.

Palicourea chiriquina is known only from Panama.

CHiRiQUi: Cuesta de las Palmas, Pittier 31 18 (US). Finca Collins, Boquete, Stern et al. 2014
(US).

5. Palicourea crocea (Swartz) R. & S., Syst. Veg. 5: 193. 1819.

Psychotria crocea Swartz, Prodr. Veg, Ind. Occ. 44. 1788. type: Jamaica, Swartz, not seen

1980) DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 301

Shrubs 1.5-3.5 m tall, the twigs smooth, glabrous, usually drying purple black,
ultimately slender, the nodes well spaced. Leaves lanceolate or occasionally
ovate lanceolate, 6.0-13.5 cm long, 3-7 cm wide, acute or acuminate at the
apex, the acumen short or to 1.2 cm long, the base acute or attenuate acute, the
costa slender, prominulous or plane above, scarcely prominent beneath, the lat-
eral veins 12(-14), arcuate, prominulous above and beneath, not forming a distinct
submarginal vein, evanescent at margin, membranous to chartaceous, glabrous
or minutely puberulent; petioles 0.5-1.5 cm long, slender, 1.0-1.5 mm wide;
stipules connate, the sheath short, 1-2 mm long, slightly swollen, truncate, each
part with 2 subulate, well-spaced awns, as long as or 2-3 times longer than the
sheath. Inflorescences terminal, reddish, contracted paniculate, 7-8(-12) cm
long, 2-4(-5) cm wide, glabrous to minutely puberulent, the peduncle 2.5-
3.5(-7.0) cm long, slender, 0.1 cm wide, the branches few, to 4 cm long, strongly
ascending at anthesis, the lowermost 2 opposite, the upper opposite or alternate;
bracts and bracteoles often persistent in fruit, subulate, to 3 mm long, spreading,
the cymes few flowered. Flowers with pedicels capillaceous, to 1 cm long, con-
spicuously expanded just below the hypanthium; hypanthium oblong, ca. 1 mm
long, glabrous, the calycine cup scarcely measurable, the lobes 5, ovate trian-
gular, 0.3 mm long; corolla red, the tube (in bud) to 4.5 mm long; anthers 2.5-4.0
mm long (Steyermark), glabrous. Fruits rotund, oblong rotund or ovate rotund,
to 3.5 mm long, glabrous, the ribs obtuse.

Palicourea crocea is found in Nicaragua, Panama, in tropical South America,
and in the West Indies.

The Panamanian collections lack mature flowers and the several fruiting col-
lections from the Republic have the fruit much shorter than typical South Amer-
ican material which may reach to 10 mm in length. The corolla in the only flow-
ering material (Panama) seen is glabrous outside. Palicourea crocea bears a
resemblance to Fuchsia paniculata Hemsley (Onagraceae). The latter is imme-
diately distinguished by the numerous seeds which form minute bulges in the wall
of the dried fruit.

CANAi zone: Summit Garden, Croat 10789 (MO). Chiva-Chiva Trail near Miraflores Lake, Tyson
1358 (MO), darien: Confluence of Rio Chucimaque and Ri'o Canclones, Duke 5121 (MO). Panama:
Tocumen, Dwyer 4228 (MO).

6. Palicourea fastigiata Benth. in H.B.K., Nov. Gen. Sp. PI. 3: 368. 1819. type:
Atures, Orinoco, Amazonas, Venezuela, not seen.

Shrubs to 2 m tall, the branchlets angular, glabrate. Leaves elliptic oblong to
ovate oblong, 1.5-1 1.0 cm long, 3.3-5.0 cm wide, acuminate, basally acute, mem-
branous, glabrous; petioles stout, to 0.7 cm long; stipules connate, ca. 5 mm long,
each part with 2 well-spaced awns. Inflorescences terminal, flat topped; peduncle
5.0-10.5 cm long, glabrous, the branches erect or strongly ascending, fastigiate,
the corymbs tripartite; bracts subulate, glabrous. Flowers on slender pedicels,
ca. 1.5 cm long, fastigiate, glabrous; hypanthium and calycine cup small, gla-
brous, the lobes small, ovate or rounded; corolla yellow or orange pink, the tube

302

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol, 67

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Figure 66. Palicourea ^alcottiana Martens. — A. Habit with inflorescence (xVi). — B. Infruc-
tescence (x»/2). — C. Flower, part of perianth removed (xZVi). — D. Ovary, longitudinal section (x5)
[After White 29, \

cylindrical, 8-10 mm long, glabrous outside, hairy within below the middle, the
lobes 5, ovate, somewhat obtuse, reflexed; stamens 5, the anthers linear, exsert-
ed, the filaments slender, attached at the mouth; style slender, about the same
length as the corolla tube. Fruits subglobose or oblong, ca. 5 mm in diam.,
sulcate, glabrous, the persistent calyx umbilicate.

Palicourea fastigiata is known from Costa Rica, Panama, and northern South
America. The determination of this species is uncertain; study of the original
description indicates the Panamanian material cited belongs to F. fastigiata
Benth. Steyermark (1972) in reducing P. fastigiata Benth. (not Cham. &

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 303

Schlecht.) to a synomym of P. crocea var. riparia (Benth.) Griseb. f. riparia,
discusses the Venezuelan collections.

In the New York Botanical Garden an unnumbered Seemann collection from
"Veraguas" was annotated by Hooker in 1849 as P. fastigiata. Its leaves tend
to be subfalcate but in most other respects it matches those of what is here called
P. fastigiata. The inflorescences, while fastigiate, are more patently so than in
the material cited below.

CANAL zone: Madden Dam, Allen 2562 (F, NY). Lion Hill Station, Hayes 667 (F). Colombia:
CHOco: Ri'o Atrato, 2-5 hours below Rio Sucio, above Loma Teguerre, Duke 11012 (MO).

7. Paiicourea galeottiana Martens, Bull. Acad. Bruxelles 11: 136. 1844. type: Not
seen. — Fig. 66.

Psychotria mexicana Willd. ex R. & S., Syst. Veg. 5: 189. 1819. type: not seen.
Paiicourea costaricensis Benth. Vidensk. Meddel. Dansk Naturhist. Foren. Kjpbenhavn 1852: 38.
1853. type: Costa Rica, not seen.

P. mexicana Benth. ex Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kj0benhavn 1852: 37. 1853,

not Psychotria mexicana Willd.

Trees to 6 m tall, or shrubs, the twiglets smooth, glabrous, occasionally con-
tracted below the nodes, the latter well spaced, occasionally rough with the rem-
nants of stipules. Leaves lanceolate to ovate lanceolate, 6-19 cm long, 2-5 cm
wide, acute or deltoid at the apex, acuminate, the acumen deltoid to attenuate
triangular, to 1.5 cm long, the base acute, cuneate or obtuse, the costa promi-
nulous above, prominent beneath, the lateral veins 10-15(-17), widely arcuate,
with 1-2 subparallel, irregular, soon branching smaller veins between 2 lateral
veins, the intervenal areas spreading reticulate, papyraceous, glabrate, often vil-
lose on the costa or also on the veins beneath; petioles to 2 cm long, glabrous;
stipules usually persistent, connate, the sheath cylindrical, occasionally com-
pressed cylindrical, to 4.5 mm long, truncate, each part with 2 erect or somewhat
spreading awns, usually ca. 2 times the length of the sheath, to 11 mm long.
Inflorescences terminal, paniculate, oblong or pyramidal, to 15 cm long, to 15 cm
wide, the peduncle 2-6 cm long, to 0.2 cm wide, red, the branches red, of 7-8
pairs, the lowermost to 7 cm long, usually strongly ascending, occasionally strict;
lowermost bracts subulate, to 0.6 cm long. Flowers pedicellate, the pedicels red;
hypanthium ca. 0.5 mm long, glabrous, the calycine cup to 1 mm long, spreading,
often with 1-2 minute glands at the sinuses, the lobes 5, oblong or compressed
rotund, to 1 mm long, obtuse, usually with a few marginal cilia; corolla yellow
or orange (rarely white), cylindrical, 10-15 mm long, petaloid, glabrous except
ca. 3 mm above the base of the tube within, the lobes 5, oblong, 1.2-2.0 mm
long; stamens 5, the anthers narrowly oblong, ca. 3 mm long, the filaments ca.
1 mm long, attached 3 mm below the mouth; style included or briefly exserted,
5-16 mm long, the shorter style with stigmas to 3.3 mm long, the ovarian disc
compressed rotund, ca. 1 mm long. Fruits ovate oblong, to 6 mm long, conspic-
uously costate, glabrous, the calyx remnants minute.

Paiicourea galeottiana extends from Mexico to Panama. Mori and Kallunki
(4789) note that the tree was visited by hummingbirds.

304 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

CHiRiQUi: Bambito to Cerro Punta, Allen 313 (MO). Paso Ancho to Monte Lirio, Allen 1490
(MO, US). Finca Collins, Boquete, Blum & Dwyer 2571 (MO). Bambito and Cerro Punla, Croat
10579 (MO). N of Audubon Cabin, Croat 13663 (MO). NW side of Cerro Pando, Croat 15941 (MO).
EI Baru above Boquete, 1200-1800 m, D'Arcy 9898 (MO). 9 km by road WNW of Boquete, 1870 m,
Davidse & D'Arcy 10307 (MO). Volcan de Chiriqui, Davidson 878 (MO). Cerro Horqueta, Duke ct
a!. 13635, 13644 (both MO). Finca Collins, Boquete, Dwyer & Hayden 7645. 7691 (both GH, K, MO,
NY, UC, US). Cerro Horqueta, Dwyer & Lallathin 8739 (MO, NY); Hagen cfi Hagen 2055 (NY).
Finca Collins, Boquete, Hayden 161 (MO). Pinola, Kirkhride & Duke 868 (MO, NY). N of San Felix,
36.2 km from Panamerican Highway, Mori & Kallunki 6016 (MO). Monte Lirio, upper Rio Chiriqui
Viejo Valley, Seihert 301 (MO). Foothills of Volcan Baru, NW of El Hato, Stern & Chambers 63
(MO, NY). Finca Collins, Boquete. 5/^^/2 1^/«/. 1 093 (MO). Boquete, Stern etal 1989 (MS). Bambito,
t mi SW of Cerro Punta, Tyson 5690 (MO). 6 mi above Cerro Punta, Boquete trail, 7000 ft, Tyson
7080, 7196 (both MO). Rio Chiriqui Viejo Valley, Lower Cerro Punta trail, White 54 (MO). Bajo
Chorro, Woodson & Schery 655 (MO). Casita Alta, Volcan de Chiriqui, Woodson et al. 909 (MO,
NY). VERAGUAS: 5 mi W of Santa Fe, road past Escuela Agricola Alto Piedra, Croat 23033, 23061
(both MO). NW of Santa Fe, I km from Escuela Agricola Alto de Piedra, Cerro Tute, Mori &.
Kallunki 4789, 4863 (both MO). 6-7 km W of Santa Fe, 2900 ft. Nee 9824 (MO).

8. Palicourea gibbosa Dwyer. ^'* type: Panama, Gentry cfe Mori 14108 (MO, ho-
lotype).

Trees to 3 m tall, the branchlets angular and black when dry, smooth, glabrous,
the nodes well spaced. Leaves elliptic, to 20.5 cm long, to 8.5 cm wide, deltoid,
rounded or obtuse at the apex, acuminate, the acumen to 1.5 cm long, often
falcate, cuneate to acute at the base, the costa slender above, prominulous be-
neath, the lateral veins 12-17, arcuate, not forming an undulate submarginal vein,
with 1-2 smaller veins between a pair of lateral veins and wrinkled on drying,
thinly chartaceous, concolorous when dry, black brown, the costa and veins
golden puberulent; petioles slender, flexuous, 2.5-6.0 cm long, ca. 0.2 cm wide,
minutely puberulent; stipules connate, ca. 0.8 cm long, the sheath to 4 mm long,
black when dry, minutely puberulent, each part not conspicuously truncate, the
awns 2, triangular, to 4 mm long, to 2.5 mm wide at the base, acute, perhaps
scarious toward the apex. Inflorescences red, cymose paniculate, pyramidal, to
17 cm long, to 10 cm wide, the peduncle 1.0-1.5(-2.5) cm long, the branches
mostly opposite, well spaced or crowded on the rachis, occasionally arcuate or
deflexed, slender, the lowermost to 2,5 cm long; flowers few, spreading, usually
at right angles to the axis; lowermost bracts narrowly oblong, to 0.7 cm long;
bracteoles tending to persist, small. Flowers with pedicels slender, to 10 mm
long, to 0.35 mm wide, minutely puberulent; hypanthium ca. 1.5 mm long, gla-
brous, the calycine cup scarcely measurable, the lobes 5, oblong rotund, ca. 1
mm long, rounded at the apex, not contracted basally; corolla yellow, the tube
cylindrical, ca. 10 mm long, ca. 2 mm wide in the middle, gibbous at the base,
the swollen area to 5 mm wide, to 3 mm long, petaloid, glabrous outside, bearded
at the upper part of the swollen area within; stamens 5, the anthers narrowly

" Palicourea gibbosa Dwyer, spec. nov. Arbores ad 3 m alti, ramulis glabris. Folia elliptica, ad
20.5 cm longa, ad 8.5 cm lata, acumine ad 1.5 cm longo. venis lateralibus 12-17 tenui-chartacea in
costa et venis aureo-pubemla; petiolis 2.5-6.0 cm longis; stipulis connatis, ca. 0.8 cm longis, vagina
ad 4 mm longa, utraque parte setis 2 praedita. Inflorescentiae cymoso-paniculatae, ad 1 cm longae,
ad 10 cm latae, pedunculo 1.0-I.5(-2.5) cm longo. Flores in pedicellis ad 10 mm longis; hypanthio
glabro lobis calycis ca. I mm longis, apice rotundis; corolla lutea tubo ca. 10 mm longo, extus glabro;
antheris ca. 3 mm longis. Fructus compresso-rotundi vel subturbinati in sicco lutei glabri costis
inconspicuis.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 305

oblong, to 3 mm long, obtuse, the filaments slender, ca. 2 mm long, affixed near
the middle of the tube; ovarian disc compressed rotund, bilobed, to 0.8 mm long,
wider than long; style linear, to 13 mm long, drying yellow, with an S-shaped
bend toward the base. Fruits compressed rotund to subturbinate, to 7 mm long,
often as wide as long, subtruncate at the apex, drying dull yellow, glabrous, the
ribs not conspicuous, obtuse, each with a delicate median longitudinal ridge.
Fruits not seen.

Palicourea gibbosa is known only from Panama. It is marked by several

diagnostic features: inflorescence red; flowers usually at right angles to the axis

to which they are attached; corolla tube usually conspicuously gibbous and asym-
metrical at the base.

darien: West Ridge trail from summit camp to waterfall, E of camp, Cerro Tacarcuna, 1550-
1700 m. Gentry & Mori 14108 (MO).

9. Palicourea grandibracteata Dwyer.^** type: Panama, Mori & Kallunki 5262
(MO, holotype).

Shrubs to 3 m tall, the branchlets moderately slender, obtusely angular, pu-
berulent, dark red when dry, the nodes well spaced. Leaves elliptic, sometimes
obovate elliptic, 4-13 cm long, 1.5-4.5 cm wide, deltoid or rounded at the apex,
acuminate, the acumen to 1.5 cm long, ca. 0.2 mm wide in the middle, basally
acute or cuneate, often inequilateral, the costa lightly prominulous or plane
above, prominulous beneath, drying dull red, the lateral veins ca. 20, arcuate,
rigidly papyraceous, discolorous, green or brown when dry, villosulose on the
costa and the veins beneath; petioles 2-3 cm long, often curved near the lamina,
rigid, ca. 0.15 cm wide, glabrous along most of their length; stipules persistent,
strongly ascending, the sheath inflated, to 5 mm long, scarious, glabrous, each
part with 2 triangular lobes, to 8 mm long, to 5 mm wide. Inflorescences terminal,
paniculate, to 15 cm long, to 1 1 cm wide, the peduncle 5-6 cm long, often some-
what reflexed, glabrous, the branches opposite, usually disposed as 4-6 pairs;
bracts elongate, narrowly lanceolate or ovate lanceolate, the lowermost to L5 cm
long, to 0.3 cm wide, acute, keeled, closely appressed to the stem, glabrous,
venose. Flowers (only partially seen) pedicellate, the pedicels slender, to 1.1(-1.4)
cm long, ca. 0.5 mm wide; hypanthium to I mm long, glabrous, the calycine cup
inflated, to 1 mm long, the lobes 5, ovate, to 5 mm long, unequal in width, to 3
mm wide basally, acute, glabrous, venose. Fruits subrotund or turbinate, to 1.6

cm long including the persistent calyx, truncate, 10-ribbed, glabrous, black when
dry.

Palicourea grandibracteata is known only from Panama. It is distinguished

"^"^ Palicourea ^grandibracteata Dwyer, spec. nov. Frutices ad 3 m alti ramulis obtuso-angulatis
puberulis. Folia elliptica, 4-13 cm longa, 1.5-4.5 cm lata, apice deltoidea vel rotundata, acuminata
acumine ad 1.5 cm longo, basi acuta vel cuneata venis lateralibus ca. 20 rigido-papyracea; petiolis ad
2 cm longis; stipulis forte connatis vagina inflata, ad 5 mm longa, scariosa glabra parte utraque lobis
2 triangularibus, ad 8 mm longis, ad 5 mm latis. Inflorescentiae paniculatae, ad 15 cm longae, ad 11
cm latae, pedunculo 5-6 cm longo; bracteis elongatis angusto-lanceolatis vel ovato-lanceolatis, ad 1.5
cm longis, ad 0.3 cm latis. Flores calyce lobis 5 ovatis, ad 5 mm longis. Fructus ad 1.6 cm longi,

11

glabri.

306 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

by its inflated stipules, each part of which has 2 triangular lobes, by its conspic-
uous bracts and elongate calycine teeth.

VERAGUAS: NW of Santa Fe, 2 km from Escuela Agricola Alto de Piedra, Mori & Kallunki 5262
(MO).

10. Palicourea guianensis Aubl., Hist. PL Guiane 1: 173. tab, 66. 1775, type:
French Guiana, Aiiblet, not seen.

Shrubs or trees, 4(-7) m in height, the branchlets terete, ultimately angular,
smooth, glabrous, usually drying black, often constricted below the nodes, mostly
6-8 cm apart. Leaves oblong, often widely so, occasionally ovate oblong or
elliptic, rarely oblong rotund, 9.5-32.0 cm long, 4-17 cm wide, deltoid to occa-
sionally obtuse or acute at the apex, often acuminate, the acumen mostly 1-2 cm
long, often falcate, the base cuneate, acute, obtuse or rounded, often slightly
inequilateral, the costa prominulous above, prominulous to prominent beneath,
to 2.2 mm wide proximally, the lateral veins (11-)15-18, widely arcuate, the
intermediate veins, if present, l-2(-3) short, soon branching, subparallel to the
laterals, the intervenal areas smooth, occasionally minutely punctate beneath;
petioles 1.5-3.0 cm long, glabrous; stipules connate, to 10 mm long, the sheath
short, glabrous, each part 2-lobed, the lobes oblong, often closely applied to the
stem, usually 3-4 times the length of the sheath. Inflorescences terminal, usually
solitary, rarely 2-3, paniculate, glabrous or occasionally puberulent, oblong, elon-
gate dome shaped, hemispherical or triangular in outline, 15-22 cm long, 5-9 cm
wide; peduncles erect, stiff, 8-14 cm long, angular, often lignose, 0.2-0.3 cm
wide, the branches numerous and stout. Flowers pedicellate; hypanthium ca. 1
mm long, the calyx obsolete, the cup acetabuliform, scarcely measurable, the
teeth scarcely measurable; corolla yellow, the tube cylindrical, somewhat con-
stricted below the middle, expanded basally, 1-2 cm long, stiffly petaloid, pu-
berulent outside, occasionally the hairs granulose, glabrous within, the lobes 5,
triangular, often somewhat unequal in length, to 2 mm long, acute; stamens 5,
the anthers oblong, 3-5 mm long, the filaments plane, glabrous, shorter than the
anthers, attached near the middle of the tube; style slender, often as long as the
tube, the stigmatic area to 6.5 mm long, the stigmas oblong, to 5 mm long, the
ovarian disc compressed rotund, to 0.75 mm long. Fruits ovoid, to 5 mm long,
conspicuously costate, glabrous, drying black, the persistent calycine cup and

ovarian disc ca. 1 mm long.

Palicourea guianensis extends from Guatemala south through Central Amer-
ica and the Guianas of northern South America. It also occurs in the West Indies.

BOCAS DEL TORo: Mile 10, railroad, Croat & Porter 16353 (MO). Chiriquicito, Lewis et al. 2045
(GH. K, MO, UC, US). Almirante, McDaniel 5053 (MO). Isia Colon, Wedel63 (MO). Water Valley,
Wedel897, 1524, 1612, 1840 (all MO). Chiriqui Lagoon, WeJel 1355 (MO). Old Bank Island, Wedel
2057, 2168 (both MO). Fish Creek Mountains, Chiriqui Lagoon, Wedel 2355 (MO). Fish Creek low-
lands, Wedel 2384 (MO), canal zone: Mouth of Ri'o Chagres, Allen 896 (MO). Cruces Trail and
Madden Dam Road, Bartlett & Lasser 16345 (MO). Gamboa and Pedro Miguel, Bartleti & Lasser
16509 (MO). Albrook, Blum 568 (MO). Cerro Viejo on K 16 C, Blum 1223 (MO). 2 mi from Gatun
Locks, Blum 1489 (MO). Gaillard Highway, 1 mi NW of Summit, Blum 2344 (MO). Pipeline Road,
Gamboa, Clewell & Tyson 3316 (MO). Albrook, Correa 33 (MO). Road K-6 E of Arraijan, Croat
15028 (MO). BetweenLocks and Fort Sherman, Croat 15362 (MO). Camp Pina, Duke 4423 (MO).

1980J DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 307

Hill C-6, Fort Sherman, Duke 4424 (MO). Road K-10, 1-2 mi from Kobbe, Duke 11695 (MO). Fort
Sherman, Duke & Mussell 6635 (MO). K-6 Road, Dwyer 2850 (MO). Albrook, Dwyer 6748 (MO).
Coco Solo, Dwyer & Duke 7884 (GH, MO). Gamboa, Navy Pipeline Road, Hoines 545 (MO). Ancon
Hill, Killip 39950 (MO). Fort Giilick, Lazor & Blum 5394 (MO). Gatun Lake, Lewis et at. 1832 (GH,
MO, US). Gaillard Highway at Rio Masambi Grande, Nee 7066 (MO). Albrook Air Force Base,
Stimson 5069 (MO). Fort Sherman, Road S-2, Stimson 5099 (MO). Albrook Air Force Base, Stimson
et al. 5210 (MO). Farfan Beach, Sullivan 569 (MO). 6 mi N of Gamboa, Tyson 1494 (MO). Gatun
Railroad Station, Tyson 2281 (MO), barro Colorado island: Croat 4608, 4898. 5408, 5568, 5836,
5957, 6184, 6364, 6380, 12426 (all MO); Ehinger 17 (MO); Hayden 2, 35 (both MO); Shattuck 954
(MO); Starry 6, 135 (both MO); Tyson 4208 (MO), colon: Santa Rita Ridge Trail, 17-35 km from
Boyd-Roosevelt Highway, 400-800'm, Mori & Crosby 6322 (MO), darien: Cerro Pirre, Bristan 505
(MO). El Real, Croat & Porter 15465 (MO). Rio Pirre. Croat & Porter 15536 (MO). 2-3 mi SE of El
Real. Duke 4833 (MO). Rfo Pinas, Duke 10547 (MO). Upper Rio Membrillo and Camp 7, Construction
Road to San Bias, Duke 10893 (MO). Loma Cuasi, behind Manene, Duke 13616 (MO). Rio Mortf,
Camp Morti Hydro, Duke 14119 (MO). Uruti River, Duke & Bristan 229 (MO). Cerro Pirre, Duke
& Elias 13667, 13677 (both COL, MO, UC. VEN). Manene to mouth of Ri'o Cuasi, Kirkhride &
Bristan 1396 (MO). 0-2 mi E of Tres Bocas, Kirkhride & Duke 1156 (MO). N slope of Cerro Pirre,
300-700 m, Mori & Kallunki 5376 (MO). N slope of Cerro Pirre lower montane rain forest, 700-950
m, Mori & Kallunki 5469 (MO). LOS santos: Loma Prieta, Duke 11862 (MO); Lewis et al. 2216 (GH,
MO, US). Panama: Altos de Pacora, Busey 805 (MO). Cerro Jefe, Duke 9420 (MO); Dwyer et al.
5033 (MO, SCZ, UC); Kirkhride & Crehhs 23 (MO). SE slopes Cerro Trinidad, Kirkhride & Duke
1671 (MO). Cerro Azul, Tyson 2140 (MO). Chepo. Tyson 6774 (MO). Cerro Jefe, Tyson et al. 3331,
3430, 4368, 4400 (all MO). Arraijan, Woodson et al. 1365 (MO), san blas: SE of Puerto Obaldfa,
Croat 16735 (MO). Cuna Reservation, Rio Morti, 6 mi upstream from Mortf Arriba, Duke 8449 (MO).
Mulatuppu (Rio IbedO, Duke 8484, 8486 (both MO). E of Cangandi'-Mandinga Airport. 2-5 mi S of
Mandinga, Duke 14775 (MO). Ri'o Mulutupu, Kirkhride 222 (MO). Opposite Ailigandi, Lewis et al.
210 (GH, K, MO, NY, UC, US), veraguas: Rio de Jesus, Dwyer 1333 (MO).

11. Palicourea lasiorrachis Oerst., Vidensk. Meddel. Dansk Naturhist. Foren.
Kj0benhavn 39. 1852. type: Costa Rica, Oersted 11555 (F, photo, NY, pho-
to, US, isotype).

Shrubs to 3 m tall, the branchlets terete, smooth, glabrous, ultimately pilose
or villosulose, contorted, angular, the pith porate. Leaves elliptic oblong to ovate
oblong, 6.5-16.0 cm long, 2-6 cm wide, the costa prominulous above, prominent
beneath, the lateral veins ca. 16, arcuate, with 1-2 subparallel irregular veins,
evanescent toward the margin, the intervenal areas spreading reticulate, stiffly
chartaceous, glabrous above, villosulose on the costa and the veins beneath;
petioles to 2 cm long, to 0.1 cm wide, the stipules connate, the sheath 3-5 mm
long, truncate, scarious, glabrous or with a few stiffly subulate, deciduous hairs,
each part with 2 subulate awns, ca. 3 mm long. Inflorescences terminal, solitary,
thrysiform paniculate, to 14 cm long, to 7 cm wide; peduncle 4-8 cm long, ca.
0.8 cm wide, webby pubescent, the hairs of varying lengths, the branches well
spaced, to 3.5 cm long; bracts and bracteoles subulate. Flowers pedicellate; hy-
panthium rotund to oblong, 0.3-0.8 mm long, glabrate to puberulent, granulose,
the calycine cup scarcely measurable, eglandular within but with occasional sub-
ulate glands at the sinuses, the lobes 5, triangular or rotund, 0.5-0.7 mm long,
densely villosulose to glabrate; corolla green to yellow, the tube slender, 7-10
mm long, glabrous outside except occasionally granulose and pubescent toward
the apex, densely villose within 2-3 mm above base; anthers 5, narrowly oblong,
subsessile; style slender, to 4.5 mm long, the stigmas digitiform, ca. 2 mm long.
Fruits not seen.

Palicourea lasiorrachis is known only from Costa Rica and Panama. The

308 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

flowers are small for a Palicourea and the indumentum of the calyx and the
corolla is quite variable. Kirkbride & Duke {994) note that the corolla is purple
with the inflorescence green to yellow. Antonio Molina has annotated a sheet,
Williams et al. 28241 (F), from El Canon, S of Cartago, Talamanca Range, Costa
Rica, 2,500 m as P. lasiorrachis. The fruit is turbinate, ca. 5 mm long, conspic-
uously ribbed, on pedicels shorter than the fruit.

CHiRiQUi: Bajo Chorro, Boquete, 6000 ft. Davidson 171 (MO). Chiriqiii Trail, Elfin Forest at
Divide, Kirkbride & Duke 994 (MO).

12. Palicourea macrosepala Krause, Bot. Jahrb. Syst. 54: Beibl. 119: 41. 1916.
type: Costa Rica, Tonduz 7704 (Photo, F),

Trees to 5.6 m tall, the bark of the twiglets somewhat smooth, minutely pus-
tulate, glabrescent. Leaves strongly ascending, oblong elliptic, 18-21 cm long,
7.5-8.0 cm wide, deltoid or acute at the apex, attenuate acute at the base, the
costa plane or sulcate above, prominent beneath, angular, drying red, to 1.5 mm
wide, the lateral veins 15-18, arcuate, drying red beneath, the intervenal areas
spreading reticulate, the blade glabrous above, pubescent beneath, especially on
the veins, the costa sparsely ciliate, the intervenal areas minutely pubescent;
petioles to 3.5(-4.5) cm long, glabrescent, cracking, sometimes twisted, the blade
slightly decurrent; stipules connate, intrapetiolar, the sheath appressed to the
stem, cylindrical, ca. 0.7 cm long, stiffly petaloid, each part with 2 triangular,
coriaceous lobes shorter than the sheath, the hairs few. Inflorescences terminal,
solitary, cymose paniculate, to 23 cm long, plane, the lowermost branches op-
posite, strongly ascending, to 9 cm long; bracts linear subulate, to 1 cm long, the

longer ones with a median subulate process and 2 short processes. Flowers with
the hypanthium compressed cylindrical, ca. 1 mm long, glabrous, the calycine

cup ca. 0.8 mm long, the lobes 5, oblong, 1.5-2.0 mm long, scarcely puberulent;

corolla pale purple, the tube cylindrical, uniformly wide, ca. 5 mm long, thinly
carnose, glabrous within, the lobes 5, 1-3 mm long, thick, glabrous, cucullate;

stamens 4-5, the anthers narrowly oblong, ca. 3 mm long, obtuse, sagittate ba-
sally, the filaments ca. 0.3 mm long, attached near the mouth or at the base of
the tube; ovarian disc compressed ovate rotund, ca. 0.5 mm long, the style slen-
der, 3.2-4.5 mm long, the stigmas ca. 0.5 mm long. Fruits (immature) narrowly
oblong, ca. 5 mm long, ca. 1.5 mm wide, obtuse, conspicuously reflexed, the
calycine lobes persistent, much wider than at anthesis.

Palicourea macrosepala is known from Costa Rica and Panama. In the type
the calyx lobes are longer than in the Panamanian collection, being 2.5-3.0 mm
long. The inflorescence appears different in having few pairs of widely spread
branches rather than strongly ascending numerous branches. The Costa Rican P.
brenesii Standley has markedly elongate calycine lobes in the flowering stage.
The leaf blades of this species, however, tend to be obtuse or rounded at the
apex and the lateral veins (ca. 24) are more numerous than in P. macrosepala,

CHiRiQuf: 5.4 km from Hato de Volcan on the road to Los Lagunas, Correa & Lazor 1478 (MO).

1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 309

13. Palicourea montensis Dwyer/^ type: Panama, Mori & Kallunki 5880 (MO,
holotype).

Trees to 5 m tall, the branchlets terete, ultimately angular or plane, often
twisted, smooth, black when dry, golden puberulent, the nodes well spaced.
Leaves often moderately ascending, ovate oblong or oblong, 4.5-10,0 cm long,
2.5-5.0 cm wide, deltoid or widely deltoid toward the apex, acuminate, the acu-
men to 1 cm long, obtuse to truncate at the base, sometimes inequilateral, the
costa subplane above, prominulous beneath, the lateral veins 10-11, arcuate, the
submarginal vein close to the margin, puberulent on both sides, especially on the
costa and the veins, rigidly papyraceous, discolorous; petioles rigid, to 2 cm long,
ca. 0.15 cm wide; stipules small, connate, compressed rotund, to 0.4 mm long,
each part of the sheath with 2 ovate rotund lobes Vi-H the length of the sheath,
obtuse, puberulent. Inflorescences terminal or pseudo terminal, pyramidal panic-
ulate, to 16 cm long, to 23 cm wide, the peduncle to 7 cm long, to 0.35 cm wide,
puberulent, the lowermost branches 2, opposite, to 7 cm long, often arcuate, the
upper branches disposed as 4-5 well spaced pairs, mostly 1-4 cm apart; bracts
subulate, 3.5 mm long. Flowers sessile or subsessile; hypanthium short, the ca-
lycine cup small, patulous, to 0.2 mm long, the lobes 5, subrotund, to 0.3 mm
long, obtuse; corolla white, the tube cylindrical, to 4 mm long, to 2.3 mm wide,
petaloid, glabrous outside, within densely villose near the middle, the lobes 5,
oblong, to 2 mm long, obtuse; stamens 5, the anthers narrowly oblong, ca. 1.5
mm long, obtuse, deeply sagittate below, the filaments linear, ca, 2 mm long,
attached below the middle of the tube; ovarian disc oblong rotund, to 0.6 mm
long; style ca. 2 mm long, the stigmas ca. 1.3 mm long, thick. Fruits not seen.

Palicourea montensis is known only from Panama. It is closely related to P,
tumidonodosa Dwyer, known only from one collection in Darien Province, which
has a somewhat immature inflorescence. It differs in having a much more patulous
inflorescence, in having smaller and less turgid stipules with each part of the
sheath bilobed, in its thin chartaceous leaves with fewer lateral veins, and in
being puberulent. The nodes of P, tumidonodosa, unlike those of P. montensis,
are conspicuously swollen and turgid. The bracts of P. montensis, subtending
the lowermost branches of the inflorescence, seem shorter. The fact that the
flowers are white is noteworthy.

CHiRiQui: N of San Felix, on Bocas del Toro border, 5000-5500 ft, Mori & Kallunki 5880 (MO).

^^ Palicourea montensis Dwyer, spec. nov. Arbores ad 5 m altae ramulis aureo-puberulis. Folia
ovalo-oblonga vel oblonga, 4.5-10.0 cm longa, 2.5-5.0 cm lata, acumine ad I cm longo, subobtusa
basi vel truncata venis lateralibus 10-11, rigido-papyracea discoloria minuto-puberula utrinque; pe-
tiolis ad 2 cm longis; stipulis connatis vagina ad 0.4 mm longa, lobis 2 ovato-rotundis, 0.2-0.3 mm
longis, obtusis puberulis. Inflorescentiae terminales vel pseudoterminales, ad 16 cm longae, ad 23 cm
latae, pedunculo ad 7 cm longo, jugis ramorum inferiorium ad 7 cm longis. Flores hypanthio bre-
vissimo cupula calycis ad 0.2 mm longa, lobis subrotundis, ad 0.3 mm longis; corolla alba tubo ad 4
mm longo extus glabro intus proxime medium denso-villoso, lobis ad 2 mm longis; antheris ad 1.5
mm longis; stylo ca. 2 mm longo; disco ovarii ca. 0.6 mm longo.

310 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

14. Palicourea montivaga Standley, J. Wash. Acad. Sci. 18: 279. 1928. type:
Costa Rica, Standley & Valeria 49850 (US, holotype; F, isotype).

P. salicifolia Standley, J. Wash. Acad. Sci. 18: 280. 1928. type: Costa Rica, Tonduz 11828 (F,
holotype).

Shrubs or subshrubs to 2.5 m tall (fide Standley), the branchlets smooth,
glabrous, the pith porate, the nodes crowded or not, occasionally swollen. Leaves
elliptic or obovate elliptic, 4-10 cm long, 1.0-3.2(-4.0) cm wide, deltoid or obtuse
at the apex, often abruptly acuminate, the acumen to 0.7 cm long, acute, usually
vaguely falcate, basally cuneate to acute, the costa prominulous above and be-
neath, the lateral veins 10-13, scarcely prominulous above, prominulous beneath,
with 1^ subparallel intermediate veins between a pair of lateral veins, these soon
evanescent, the intervenal areas patulous reticulate, stiffly chartaceous, mem-
branous ?, glabrous; petioles to 1.5 cm long, 0.1-0.15 cm wide; stipules connate,
the sheath ca. 2 mm long, each with 2 awns about the same length as the body
or longer, to 2.5 mm long, scarious, glabrous. Inflorescences (in fruit) cymose
paniculate, oblong, to 9 cm long, to 5 cm wide, rounded at the apex; peduncles
to 3 cm long, the lowermost branches paired, strongly ascending, to 4 cm long,
the upper branches several, well spaced, spreading, paired or ternate, glabrous
or puberulent. Flowers sessile or on pedicels to 2.5 mm long; hypanthium ca. I
mm long, the calycine cup ca. 0.6 mm long, the teeth triangular, ca. 0.3 mm long,
obtuse or acute; corolla green or yellow, the tube ca. 7 mm long, ca. 2.5 mm
wide, the lobes triangular, obtuse, ca. 1.5 mm long. Fruits obovate to obovate
rotund, ca. 4.5 mm long, glabrous, each pyrene 5-costate, often one seed aborted
and then the fruit subfalcate.

Palicourea montivaga is known from Costa Rica and Panama. The leaves of
the type collection are much narrower and have a more salicoid appearance than

the single Panama collection.

CHlRiQuf: Bajo Chorro, Davidson 255 (MO, US).

15. Palicourea ochnoides Dwyer.^^ type: Panama, Witherspoon et al, 8872 (MO,
holotype).

Trees to 3 m tall, the branchlets slender, glabrous, drying red, glabrous, the
nodes well spaced, swollen. Leaves lanceolate, 8-14 cm long, 2.5-4.0 cm wide,
acute, caudate acuminate, the acumen to 1.5 cm long, the costa slender, plane
or immersed above, prominulous beneath, to 1.5 mm wide proximally beneath,
the lateral veins ca. 22, arcuate, plane above, prominulous and drying red be-

''^ Palicourea ochnoides Dwyer, spec. nov. Arbores ramulis gracilibus, glabris. Folia lanceolata,
ad 14 cm longa, ad 4 cm lata, venis lateralibus ca. 22, rigido-chartacea supra glabra sublus puberula;
petiolis ad 2.5 cm longis; stipulis connatis vagina turgida scariosa glabra in sicco rubra parte utraque
lobis triangularibus duobus, ad 4 mm latis basi. Inflorescentiae terminales cymoso-paniculatae, pyr-
amidales, ad 12 cm longae, ramis oppositis jugis 3-4, bracteis inferioribus ad 1.8 cm longis concavis
brevi-auriculatis basi; bracteola utraque florem unum occulante foliosa oblonga, ad 11 mm longa, ad
5 mm lata, subplana, glabra. Flores pedicellati; hypanthio glabro cupula calycis brevissima lobis 5
foliosis imbricatis ovato-oblongis, ad 5 mm longis, ad 2.5 mm latis subconcavis glabris in sicco luteo;
corolla alba tubo intus villoso.

1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 3 1 1

neath, not forming a distinct undulate, submarginal vein, with 1-2 smaller veins
subparallel to and located between adjacent lateral veins, these evanescent to-
ward the middle of the blade, the smaller veins pinnatiform, the margin scarcely
thickened, stiffly chartaceous, concolorous, glabrous and smooth above, puber-
ulent beneath; petioles 0.5-2.5 cm long, ca. I mm wide, stiff, glabrous; stipules
persistent toward the tips of the twiglets, connate, the sheath turgid, cylindrical,
scarious, drying red, each part with 2 triangular lobes, about as long as the sheath,
to 4 mm wide at the base, ciliolate. Inflorescences terminal, solitary, cymose
pyramidal paniculate, to 12 cm long, to 6.5 cm wide, the peduncle to 6.5 cm long,
to 0.18 cm wide, smooth, glabrous, the branches opposite, disposed as 3-4 well-
spaced pairs, the lowermost to 3 cm long, the lowermost bracts oblong, to 1.8
cm long, concave and often investing much of the branch, acuminate, scarious,
drying red, glabrous, shiny, venose, with an auricle on each side at the base, the
lobe to 1 mm long, obtuse, ciliolate, each flower apparently partially hidden by
a bracteole, the latter oblong, to 11 mm long, to 5 mm wide, foliose, acuminate,
venose, glabrous, the same general texture as a calycine lobe. Flowers pedicel-
late, the pedicels to 1.5 mm long, glabrous, the hypanthium 0.7-1.0 mm long,
drying red, glabrous; calycine cup short, expanded, the lobes 5, imbricate, foliose,
ovate oblong, to 6 mm long, to 2.5 mm wide, obtuse, concave, glabrous inside
and out, venose, drying yellow; corolla white, carnose, glabrous outside, villose
within, the lobes slightly cucullate; anthers 5, oblong, to 1 mm long. Fruits not
seen.

Palicourea ochnoides is known only from Panama. It is so called because the
inflorescence and the flower buds have the appearance of an Ouratea of the
Ochnaceae. The new species has several diagnostic characters: the lowermost
bracts as long as the branches of the inflorescence and auriculate at the base;
bracts subtending the individual flowers being foliaceous; calycine lobes being
imbricate and reaching a length of 6 mm. Unfortunately only immature corollas,
stamens, etc. were seen. While the collectors described the flowers as white, it
is likely that they were describing the bracteoles and/or the calycine lobes rather
than the corolla.

VERAGUAS: Trail E side of Cerro Tute, ca. 1200 m, Withcrspoon et al. 8872 (MO).

16. Palicourea panamensis Standley, Ann. Missouri Bot. Gard. 25: 839. 1938.
type: Panama, White & White 14 (MO, holotype; F, isotype).

Shrubs to 3 m tall, the twigs terete, glabrous or pubescent, usually golden
villose above, the twiglets hollow, 4-angled, bright purple, the hairs deciduous.
Leaves elliptic oblong or ovate oblong, 10-23 cm long, 6-11 cm wide, rounded
or obtuse at the apex, abruptly acuminate, the acumen widely triangular, 1.0-1.5
cm long, acute, widely cuneate and often slightly inequilateral at the base, the
costa prominulous above, subplane to prominent beneath, the lateral veins 12-
21, widely arcuate, the intermediate veins several, branched and evanescent,
often short, the tertiary veins parallel and somewhat perpendicular, papyraceous
or chartaceous, discolorous, glabrate above, puberulent on costa and veins be-
neath; petioles 1.5-4.5 cm long, 0.15-0.25 cm wide, stiff or flexuous, glabrate;
stipules persistent, connate, the sheath 0.8-1.5 cm long, turgid, cylindrical, trun-

312 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

cate, each part with 2 lanceolate, subulate or ovate subulate equal lobes, coria-
ceous, puberulent. Inflorescences terminal, solitary, thrysoid paniculate, ovate
oblong to pyramidal, to 26 cm long, to 14 cm wide, the peduncles rigid, 3-7 cm
long, 0.2-0,4 cm wide, angular, puberulent. Flowers with the hypanthium oblong,
ca. 1 mm long, puberulent, the calycine cup ca. 0.3 mm long, the lobes 5, unequal,
1.2 mm long, acute, puberulent or ferrugineous villose, the margin ciliolate, re-
flexed after fall of corolla tube; corolla purple, lavender, magenta, or yellow?,
the tube to 15 mm long, glabrate or pubescent. Fruits (immature) subrotund or
turbinate, to 3 mm long, truncate, the persistent calyx to 2 mm long, the lobes
ciliolate, triangular, longer than the calycine cup, 10-ribbed, glabrous, drying

black.

I have reservations about placing the new species in Palicourea rather than in
Psychotria. The fact that the corolla tube is gibbous at the base, the anthers blue,
the calyx lobes lavender (fide Dwyer 5034) and the cymules lavender green seem
to favor Palicourea. The corolla, on the other hand, is white which is more sugges-
tive of Psychotria. The stipules are so reduced in size as to be difficult to discern
and often appear to be torn by the exaggerated nodal scars.

PANAMA: Cerro Jefe, Dwyer et al. 5034 (MO); Mori & KaUimki 6082, 6493, 6505\ Mori 7127 (all
MO).

Palicourea panamensis is known only from Panama. Collections from Chiriqui
have flowers ranging in color from purple, lavender, to magenta. Duke 6575 from
Darien notes that the flowers are yellow and the pedicels red.

BOCAS DEL TORO: Divide on Chiriquicito-Caldera Trail, Elfin Forest, Kirkhride & Duke 994 (MO).
CHiRiQuf: Slopes of Volcan de Chiriqui Viejo, along Ri'o Chiriqui Viejo, 1200 m, Allen 986 (MO).
NW side of Cerro Pando, Croat 15939 (MO). Foothills of Volcan Baru, N W of El Halo, 4500 ft. Stern
& Chambers 61 (MO). Las Lagunas, 2 m SW of El Volcan, Tyson 870 (MO). Valley of upper Rio
Chiriqui Viejo, 1300-1900 m, White & White 14 (MO), darien: Cerro Pirre, 3700 ft, Duke 6575 (MO).
Cerro Pirre, 2500-4500 ft, Duke & Elias E 13729 (MO). Cuasi-Cana trail between Cerro Campamiento
and La Esclarera, E of Tres Bocas, Kirkhride & Duke 1331 (MO), veraguas: Santa Fe, 2 km from
Escuela Agricola, Alto de Piedra, Mori & Kallunki 5262 (MO).

17. Palicourea pauciflora Standley, J. Wash. Acad. Sci. 18: 278. 1928. type:
Panama, Pittier 3237 (US, holotype).

Shrubs, the twigs branching freely, terete, ultimately angular, the nodes well
spaced. Leaves elliptic to obovate elliptic, to 7.5 cm long, to 2.5 cm wide, nar-
rowly acuminate, the acumen to 1.5 cm long, ultimately acute, cuneate at the
base, the lateral veins ca. 11, arcuate, stiffly chartaceous, concolorous; petioles
ca. I cm long; stipules persistent, connate, turgid, ca. 0.5 cm long, each part with
2 triangular lobes, acute to obtuse. Inflorescences terminal, to 5 cm long, to 5
cm wide; peduncle to 3 cm long, the branches few, slender, opposite, few flow-
ered, the bracts narrowly oblong, to 1 cm long; bracteoles narrowly oblong.
Flowers with pedicels to 1 cm long; hypanthium short, the calycine cup and lobes
to 0.8 cm long, the lobes longer than the cup, glabrous, 5.0-6.5 mm long; corolla
greenish purple, large, 1.0-1.5 cm long, the tube about twice the length of the
lobes. Fruits not seen.

Palicourea pauciflora is known only from the type collection.

chiriqui: Cerro Horqueta, 2100-2268 m, Pittier 3237 (F, US).

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 313

18. Palicourea rigidifolia (Dwyer & Hayden) Dwyer, comb. nov.

Cephaelis rigiJifolia Dwyer & Hayden, Ann. Missouri Bot. Gard. 55: 37. 1968. type: Dwyer 8075 A
(MO, holotype).

Shrubs or small trees to 5 m tall, the branchlets terete, the internodes drying
contracted and angular. Leaves oblong, 10-21 cm long, 3.5-8.0 cm wide, acute
at the apex, ultimately acuminate, the acumen to 1.3 cm long, cuneate or slightly
obtuse at the base, the costa prominulous above, prominent beneath, convex, the
lateral veins 12-15, arcuate, prominent, stiffly subcoriaceous to rigidly charta-
ceous, slightly discolorous, brown when dry, appressed white ciliate above, gold-
en pilose beneath with some hairs hook shaped and rigid; petioles 1.0-2.5 cm
long, lignose, pilose; stipules connate, the sheath urceolate or briefly cylindrical,
rigid, turgid, to 0.7 cm long, golden puberulent, each part with 2 subulate awns,
to 0.4 cm long, rigid, well spaced and erect. Inflorescences terminal, to 9 cm
long, the peduncle absent or 1-4 cm long, the flowers often crowded above the
middle of the rachis, the latter densely villosulose, to 0.3 cm wide; the lowermost
bracts subulate, to 4 mm long, the flowers large, arranged in a loose subrotund
head, to 8 cm wide, sometimes resembling a fan. Flowers with the hypanthium
subrotund, to 1 mm long, white villose, the teeth 5-6, ca. 0.8 mm long; corolla
violet or purple, the tube conspicuously elongate, to 4 cm long, to 0.6 cm wide
toward the apex, narrower below the middle, to 1.5 mm wide, petaloid outside
with patches of webby, flat, triangularly subulate hairs to 1.5 mm long, the cells
of which are bubblelike, glabrous within, the lobes 5, short, oblong, to 3 mm
long, thicker than the tube, moderately hooded; stamens 5, the anthers oblong,
3.0-3.5 mm long, the style elongate, 30-40 mm long, ca. 0.35 mm wide, plane
compressed, the stigmas 2, fingerlike, 2.0-3.3 mm long. Fruits sessile, immature,
didymous, dirty yellow, drying red, subrotund, to 8 mm in diam., minutely golden
puberulent.

Palicourea rigidifolia is known only from Panama. It is distinguished by its
stiff and usually subcoriaceous leaves and its corolla having a cobwebby mass of
hairs outside.

PANAMA: Cerro Jefe, Busey 787 (MO); Dwyer 8075 A (MO); Folsoin et al. 2010 (MO); Gentry
762 (MO); Tyson et al. 4339 (MO).

19. Palicourea subrubra Polak., Linnaea 4: 571. 1877. type: Costa Rica, Pola-
kowsky 200 A (Photo, F).

Small trees to 3 m tall, the branchlets slender, ultimately 4-angled, glabrous,
the nodes well spaced. Leaves ascending, narrowly lanceolate, 10-15 cm long,
2.5-3,6 cm wide, attenuate toward the apex, often contracted, acuminate, the
acumen to 2.5 cm long, cuneate at the base, the costa plane or immersed above,
scarcely prominent beneath, the lateral veins 12-14(-20), arcuate, slender, prom-
inulous, the submarginal vein absent, rigidly membranous, glabrate above, pu-
berulent beneath, somewhat hirsute on the costa, the hairs stiff, parallel to the
blade; petioles slender, 2.5-2.8 cm long, to 0.1 cm wide; stipules connate, per-
sistent, the sheath somewhat turgid, to 0.3 cm long, to 0.3 cm wide, each part
with 2 awns, the latter to 0.25 cm long, erect, glabrous. Inflorescences terminal.

314

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 67

Figure 67. PaUcourea triphylla (MueU.-Arg.) DC— A. Habit (xVi).— B. Flower (x2>/2).— C
Flower, opened to show interior (xZVi).— D. Fruit (xV/i). [After Nee 8178.]

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 315

paniculate, to II cm long, to 9 cm wide, pink; peduncle to 3 cm long, glabrous,
the branches disposed as 7-10 pairs, opposite or subopposite, spreading or as-
cending, the lowermost to 6 cm long; bracts often persistent, subulate, to 0.5 cm
long. Flowers with pedicels 0.3-0.5 cm long, slender, with each bracteole minute,
oblong or subulate, to 1.5 mm long, erosulose at the apex; hypanthium ca. 1 mm
long, the calycine teeth 5, triangular, ca. 0.35 mm long; corolla yellow, red,
purple, violet, the tube narrowly cylindrical, 10-15 mm long, glabrous outside,'
villose within, only 1-2 mm wide at the base, the lobes 5, triangular, ca. 1 mm
long; stamens 5, the anthers narrowly oblong, to 3.2 mm long, obtuse, the fila-
ments ca. 2 mm long, attached ca. 3 mm below the mouth; ovarian disc subrotund,
to 0.7 mm long, the style slender, to 4.5 mm long, the stigmas 2, linear oblong,
ca. 2 mm long. Fruits with pedicels shorter or longer than the fruit proper, oblong
rotund or oblong, to 0.8 cm long, ribbed, the persistent calyx to '/a the length of
the body, glabrous.

Palicourea subruhra is known from Costa Rica and Panama. It is a high
mountain species distinguished from the other Panamanian species by its reduced
stipules and pinkish inflorescence. The variability of corolla colors is noteworthy.

CHiRiQui: Bajo Chorro, Davidson III (F). Cerro Horqueta. 6500 ft, Hagen & Ho^^en 2069 (F).
darien: Cerro Tacarcuna, S slope, ridge top forest, well below summit, premontane wet forest 1250-
1450 m, Gentry & Mori 13913 (MO).

20. Palicourea triphylla (Muell.-Arg.) DC, Prodr. 4: 526. 1830.— Fig. 67.

Psychotria triphylla Muell.-Arg. in Mart.. Fl. Bras. 6(6): 233. 1881. type: British Guiana, Patris, not
seen.

Shrubs or trees to 7 m tall, the branchlets terete, smooth, glabrous, usually
constricted below the nodes. Leaves 2 or 3 at a node, lanceolate, often falcate,
often 4 times longer than wide, to 26 cm long, to 8 cm wide, acute, usually
cuspidate, the cusp to 1.5 cm long, acute or cuneate at the base, usually slightly
inequilateral, the costa prominulous above, prominent beneath, the lateral veins

12-16, arcuate, prominulous or immersed above, prominulous beneath, the in-
tervenal areas spreading reticulate, papyraceous, usually concolorous, puberulent
above especially on the costa, often farinose above from persistent hair bases,
villosulose beneath especially on the veins, often farinose in the intervenal areas;
petioles almost absent or to 1.5 cm long, often curved; stipules connate, the
sheath short, to 2 mm long, the awns 2, erect, subulate, to 12 mm long, stiff,
marginally puberulent. Inflorescences terminal, solitary, exceeding the upper-
most leaves, to 30 cm long, the peduncle rarely absent, to 15 cm long, often as
long as the floriferous portion, stout, puberulent, thrysoid paniculate, the branch-
es numerous, stiffly ascending, the flowers numerous. Flowers with the hypan-
thium subrotund to cuneate, ca. 1 mm long, puberulent, the calycine cup ca. 1
mm long, glabrous within, the teeth reduced to 5 irregular erosulose points; co-
rolla orange or yellow, the tube narrowly cylindrical, subturgid at the base, ca.
12.5 mm long, densely puberulent outside, villose within at the mouth and with
a few hairs at the base of the tube, the lobes 5, short, ca. 1.5 mm long, obtuse,
not cucullate; stamens 5, the anthers narrowly oblong, ca. 3 mm long, obtuse,
dorsifixed, the filaments short, ca. 1.5 mm long, attached near upper V3 of the

316 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

tube; style linear, the ovary thick walled. Fruits pedicellate, subrotund, ovate
rotund, ca. 4 mm in diam., the calycine cup coroniform, each pyrene markedly
4-costate, puberulent, the raphides numerous.

This species is widely distributed throughout tropical America. The fact that
the species has ternate leaves permits ready recognition.

BOCAS DEL TORo: Macaw Hills, S of Bocas, Wedel 521 (MO). Chiriqui Lagoon, Wedel 1143
(MO) Water Valley, Wedel 1904 (MO), canal zone: Ancon Hill, Bartlett & Lasser 16313 (MO).
Ancon, Gervais 132 (US). Corozal, Standley 27374 (US). Rio Tapia, Standley 28172, 28205 (both
US) Atbrook Air Force Base, Tyson 2033 (MO), chiriqui: San Felix to Cerro Flor, Allen 1939 (MO).
San Felix, Pittier 5254 (US), cocle: El Valle, Allen 1892 (MO, NY), 2477 (MO), 3603 (NY). Penono-
me, Dnyer 2018 (NY). La Pintada, Hunter cS Allen 523 (MO). El Valle, Woodson et al. 1303 (MO,
NY), darien: Gold mine at Cana, 500-600 m, Croat 37644 (MO). Cana, Goldman 1906 (US). La
Palma, Pittier 6698 (US). Cana, Stern et al. 506 (MO). Periaque Camp, Tyson et al. 4766 (MO).
HERRERa: Las Minas and Pese, Duke 12334 (MO). Ocu, Lenis et al. 1648 (MO). La Peiia, Stern et
al. 1762 (MO). Panama: Rio Tocumen, Bartlett & Lasser 16478 (MO). Panama and Chepo, Dodge
et al. 16641 (MO). Cerro Azul, Dw\er 4001 (MO). Cerro Campana, Dnyer et al. 4715 (MO). Pacora
and Chepo, Dwver et al. 5122 (MO). Cerro Azul, Dwyer & Hayden 8026 (MO). Goofy Lake, Gentry
6122 (MO, NY). Ri'o Tocumen, Hunter & Allen 245 (MO). Sabanas, Bro. Paul 607 (US). Taboga
Island, Pittier 3597 (US). Rio Tocumen, Standley 26641 (US). Miraflores Lake, White 249 (MO). Isla
Taboga, Woodson et al. 1473 (MO, NY), veraguas: San Jose, Santiago-Santa Fe Road, Nee 8178

(MO).

Mori

Trees 5-8 m tall, the trunk 10 cm or more dbh, the twiglets much branched,
angular, glabrous, smooth, the internodes 1.5-5 cm apart, the nodes knotty,
turgid, the petiole scars conspicuous. Leaves oblong rotund or oblong, 6-9 cm
long, 3.2-6 cm wide, regularly less than twice as long as wide, rounded or obtuse
at the apex, rarely deltoid, the acumen absent or to 3 mm long, rounded or widely
cuneate at base, the costa slender, prominulous beneath, the submarginal undu-
late vein 1-3 mm from margin, the interval areas indistinct above, reticulate
beneath, the margin plane or scarcely revolute, thickly coriaceous, drying dull
yellow green, glabrous, lustrous; petioles 1.5-2.5 cm long, rigid, glabrous; stip-
ules connate, the sheath less than 1 mm long, truncate, coriaceous, glabrous, the
awns scarcely measurable, widely spaced. Inflorescences terminal, exceeding the
uppermost leaves, cymose paniculate, 8-12.5 cm long, 3-7 cm wide, the peduncle
4-7 cm long, to 0.27 cm wide, angular, drying black, glabrous, the lowermost
branches opposite, rigid, angular-ascending, the upper branches few, usually
alternate, the bracteoles minute, triangular subulate, to 2 mm long, the cymes
few flowered, often delicately lavender-green. Flowers sessile, the hypanthium
ca. 1 mm long, drying black, glabrous, the calyx cup scarcely measurable, the

" Palicoiirea tuhiiHflora Dwyer spec. nov. Arhores 5-8 m altae, ramulis angulatis glabris nodis
crassis. Folia oblongo-rotunda vel oblonga, 6-9 cm longa, 3.2-6 cm lata, apice rotundata vel obtusa
venis lateralibus 7-10, valde coriacea glabra; petiolis 1.5-2.5 cm longis, glabris; stipulis connatis vix
2 mm longis truncatis glabris. Jnflorescentiae terminales cymoso-paniculatae, 8-12.5 cm longae 3-7
cm latae, pedunculo 4-7 cm longo ramis floribusque paucis. Florihus lobis calycis rotundis ad ovato-
rotundis ad 1 mm longis; corolla alba tubiforme extus glabra untus barbata lobis ad 4 mm longis; an-
theris lineari-oblongis, ad 2 mm longis, versatilibus. Fructus oblongi, ad 1 cm long!, vix costati, glabri.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhkiceae) 317

lobes often lavender, compressed-rotund to ovate-rotund, less than 1 mm long,
obtuse, the corolla occasionally falcate in bud, the tube white, yellow at the mouth,'
tubular, to 6.5 mm long, gibbous at base, constricted in middle, glabrous outside'
bearded above the middle within, the lobes 5, oblong, to 4 mm long, often bearded
at base; anthers linear-oblong, to 2 mm long, versatile, the filaments longer than
the tube, attached near the mouth; style filamentous, as long as the corolla tube,
the stigmas linear, to 2.2 mm long; ovarian disc mound-shaped, exceeding the
calyx. Fruits oblong, to 1 cm long, to 0.7 cm wide, often falcate (1 seed aborted?),
delicately costate, glabrous, drying black.

lotype).

Mori 13787 (MO

Trees to 5 m tall, the twigs smooth, glabrous, black when dry, the nodes
crowded, turgid (persistent stipules). Leaves elliptic, 6-13 cm long, 4.5-5.5 cm
wide, deltoid toward the apex, acuminate, the acumen scarcely evident, ulti-
mately obtuse, acute or attenuate acute toward the base, the costa prominulous
above, subprominent beneath, the lateral veins ca. 13, arcuate, not forming an
undulate marginal vein, the margin thinly callose, stiffly chartaceous, discolorous,
glabrous; petioles stiff, ascending, to 2 cm long, ca. 0.1 cm wide, glabrous; stip-
ules perhaps connate toward the base, compressed cup shaped, turgid, ca. 0.5
cm long, subtruncate or rounded at the apex, scarious, glabrous, the margin entire
or minutely retuse and ciliolate. Inflorescences terminal, solitary, paniculate, to
13 cm long, to 5 cm wide; peduncle 1.0-3.5 cm long, the branches opposite,
ascending, the lowermost well spaced, to 4 cm long; bracts and bracteoles ovate
triangular or oblong, to 3 mm long, acute or obtuse, glabrescent. Flowers (in bud)
sessile; hypanthium inconspicuous, 0.3 mm long, the calycine cup short, 0.5 mm
long, scarcely turgid, the lobes 5, oblong, 1.5 mm long, stiffly petaloid, the hairs
short, ascending, white; corolla green (in bud), the tube cylindrical, puberulent
outside, within with elongate, ascending hairs at base, the wall stiffly carnose,
the lobes cucullate; anthers oblong, 1.3 mm long, obtuse. Fruits not seen.

Palicourea tumidonodosa is known only from Panama. It is distinguished by
its swollen stipules which are without awns. Noteworthy is the fact that the leaves
are glabrous.

Panama: Cerro Mali, base camp near Colombian border, lower montane wet forest 1500 m

Gentry & Mori 13787 (MO).

''* Palicourea tumidonodosa Dwycr, spec. nov. Arhores ad 5 m altae ramulis glabris. Fotia
elliptica, 6-13 cm longa, 4.5-5.5 cm lata, versus apicem deltoidea venis lateralibus ca. 13, rigido-
chartacea glabra; petiolis ad 2 cm iongis; stipulis fortasse versus basin connatis compresso-cupuli-
formibus turgidis ca. 0.5 cm Iongis. apice subtruncatis vel rotundatis. Inftorescentiac paniculatae ad
13 cm longae, ad 5 cm iatae. pedunculo 1.0-3.5 cm longo; bracteis bracteolisque ovato-triangularibus
vel oblongis, ad 3 mm Iongis. Flores sessiles; cupula calycis dentibus 5 obiongis, ca. 1.5 mm longa;
corolla tubo cylindrico; antheris ca. 1.3 mm Iongis. Fructus non visi.

3 J8 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

23

Mori & Dressier 7796 (MO

holotype).

Trees 2-5 m tall, the branchlets glabrous or with dense, irregular, stiff, curled
hairs, often drying brown, the nodes moderately well spaced or ultimately crowd-
ed. Leaves oblong, 3-10 cm long, 0.8-4.0 cm wide, obtuse or acute at the apex,
acuminate, the acumen to 1.5 cm long, acute, cuneate to obtuse at the base, the
costa slender, subplane above, prominulous to subprominent beneath, often
drying red, the lateral veins 1 1-13, arcuate, prominulous beneath, with 1-2 veins
of comparable prominence between 2 lateral veins, the intervenal areas reticulate,
papyraceous, somewhat discolorous, drying a deep brown above, with a few
scattered stiff hairs beneath on the costa and the veins, the margin occasionally
lightly revolute; petioles to 1 cm long, glabrous to densely pubescent, often de-
flexed; stipules persistent, connate, turgid, 5.5-1.0 cm long, the sheath bowl
shaped, each part with 2 subulate awns, as long as or longer than the sheath.

Infi

15 cm long, 4.5-10.0

cm wide; peduncle 2.3-5.5 cm long, ca. 0.25 cm wide, densely golden villose,
yellow, often drying dark red, the branches few, the lowermost 2-4 cm long;
lowermost bracts oblong, to 1.5 cm long, 0.1 cm wide, densely villose. Flowers
with pedicels to 3 mm long, puberulent, the calycine cup 0.8 mm long, glabrous,
the teeth 5, widely triangular, 0.3-0.5 mm long; corolla with the tube cylindrical,
perhaps not dilated at the base, to 6.5 mm long, ca. 1.7 mm wide, petaloid,
glabrous outside, villose below the middle, the lobes 5, oblong, to 2.5 mm long,
apically deltoid, glabrous; anthers subsessile, 1.7 mm long, attached immediately
below the mouth; ovarian disc compressed oblong, 0.8 mm long; style linear, 9
mm long, stigmas wider than the style, ca. 0.5 mm long. Fruits (immature) ped-
icellate, the pedicels shorter than the fruit, turbinate, to 1.2 cm long, truncate at
the apex, cuneate basally, drying black, costate, scattered tuberculate, with a
few scattered hairs, the persistent calycine lobes withered, perhaps to 2 mm long.

Palicoiirea veraguensis is known only from Panama. It is readily distinguished
by its small leaves with pubescence scattered beneath, the hairs being stiff and
brown. The overall pubescence of the inflorescence is striking. The mature fruit
is large for a Palicourea and is probably more elongate than the 1.2 cm noted,
as the coUectors describe the living fruit as "green." The conspicuously turbinate
fruit is diagnostic.

CHiRiQUi: Cerro Colorado, 50 km N of San Felix, continental divide, cloud forest, 1200-1500 m,
Mori & Dressier 7796 (MO). N of San Felix at Chiriqui'-Bocas del Toro border. Cerro Colorado
Copper Mine Road, continental divide, 5000-5500 ft, Mori & Kallunki 5867 (MO), veraguas: Cerro
Tute, Mori 6767 (MO).

24. Palicourea vestita Standley, J. Wash. Acad. Sci. 18: 277. 1928. type: Costa
Rica, Standley 42909 (US, holotype; F, photo).

T' Palicourea veraguensis Dwyer, spec. nov. Arhores 2-5 m altae ramulis glabris vel dense
villosis. Folia oblonga, 3-10 cm longa, 0.8-4.0 cm lata, apice obtusa vel acuta acuminata venis
lateralibus 11-13. papyracea supra glabra, subtus diffuso-villosa; petiolis ad 1 cm longis; stipulis ad
1 cm longis setis subulatis. InftoresceiUiae terminales cymoso-paniculatae, 5-15 cm longae, dense
villosae, ramis paucis; bracteis inferioribus oblongis, ad 1.5 cm longis. Friictiis turbinati. ad 1.2 cm
longi. truncati basi cuneati.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 319

Shrubs 1-2 m tall, the twiglets terete, becoming angular, the hairs webby,
somewhat bulbous based, the nodes well spaced, somewhat turgid with remains
of stipules, the pith white. Leaves ascending, elliptic, 7-22 cm long, 2.5-10.0 cm
wide, obtuse to deltoid at the apex, acuminate, the acumen 1.0-1.5 cm long,
ultimately obtuse or acute, obtuse to cuneate at the base, the costa prominulous
above, prominent beneath, the lateral veins of larger blades to 30, prominulous
beneath, widely arcuate, not forming a distinct submarginal vein, usually drying
yellow, with 1-2 short and soon branching smaller veins parallel to the laterals,
rigidly chartaceous, often concolorous, often drying green above, usually golden
hirsute on the costa and veins above, more hairy beneath including the intervenal
areas; petioles to 2 cm long, rigid, webby pubescent; stipules persistent, connate,
densely pubescent, the sheath turgid, bowl shaped or barrel shaped, with a me-
dian longitudinal groove or a plane glabrate area, giving a buttocks effect to each
part, to 2.5 cm long, truncate, each part with 2 subulate, acute, often curved
awns, these equal to or exceeding the body. Inflorescences yellow, usually drying
yellow, terminal, paniculate, 8-18 cm long, 3.0-8.5 cm wide, the hairs dense,
short, stiff; peduncle 3-9 cm long, 0.1-0.2 cm wide, the lowermost branches
opposite or whorled, spreading, the uppermost branches relatively well spaced;
bracts linear subulate or linear oblong, to 0.8 cm long, pubescent, curled. Flowers
with pedicels ca. 0.5 cm long; hypanthium somewhat turbinate, ca. 1 mm long,
densely golden pubescent, the calycine cup apparently absent, the lobes 4, tri-
angular or ovate, ca. 0.5 mm long, somewhat spreading, densely pubescent; co-
rolla yellow, the tube to 13 mm long, slightly expanded at the apex and base,
petaloid, minutely pubescent outside, densely villose within below the middle,
the lobes 5, triangular, ca. 1.5 mm long; anthers linear oblong, ca. 2 mm long,
the filaments ca. 1 mm long, attached ca. 2 mm below the mouth; style to 1.5
mm long, barely exserted, the stigmas scarcely wider than the style. Fruits glo-
bose, ca. 5 mm in diam., compressed laterally, pubescent, the 2 pyrenes each
with 3 obtuse ribs.

dley 42909 [F, US]) and Panama.

[MO]

cocle: Near summit of El Petroso, ca. 7 km N of El Cope, ca. 800 m, D'Arcy 11261 (MO). Cerro
Pilon, 2900 ft, Dwyer & Lallathin 8668 A (MO, NY), 5960 (MO), veraguas: Beyond Rio Tres Brazos,
9 km from Santa Fe, Croat 25598 (MO). Valley of Rio Tercero Braso beyond Escuela Agricola Alto
Piedra above Santa Fe, primary forest along road, Croat 27341 (MO). Cerro Tute, Folsom & Edwards
3357 (MO). NW of Santa Fe, 8.8 km from Escuela Agricola Alto de Piedra, Mori & Kallunki 6173
(MO).

64. PENTAGONIA

Pentagonia Benth., Bot. Voy. Sulph. 105. 1844. Nomen conserv. type: P. mac-
rophylla Benth.

Megaphyllum Spruce ex Baill., Hist. PI. 7: 456. 1880. type: Brazil, Spruce herb., 6230 (K, not
seen).

Seemannia Hook., J. Bot. 7: 567. 1848. Nomen rejic. type: Panama, Seemann (K, not seen).
Watsonamra Kuntze, Rev. Gen. Pi. 302. 1891. Nomen rejic. type: W. macrophylla (Benth.) Kuntze.

Shrubs or small trees, rarely large trees, usually with stems simple, the stems
often succulent. Leaves usually large, entire or pinnatifid lobed, lineolate under

320 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

magnification; petioles occasionally bearing auricles, occasionally absent; stipules
free, large, acute. Inflorescences axillary, cymose corymbose; peduncle usually
present, bracteate. Flowers with the hypanthium turbinate or campanulate, the
calycine cup coriaceous, the wall replete with tough fibers, often with glands
inside, spathaceous or 5-6-lobate; corolla funnelform or tubular, coriaceous, gla-
brous or villose within, the lobes 5-6, short, valvate; stamens 5-6, the anthers
dorsifixed, included, the filaments equal or unequal in length, usually villose at
the base, attached to the tube; ovarian disc annular or cupuliform, the stigmas
narrow, obtuse, the ovary 2-celled, the ovules numerous, the placentas elongate,
swollen, adnate to the septum. Fruits baccate, nutlike when dry, globular, often
lenticellate; seeds numerous, obtusely angular, the testa thin.

Pentagonia occurs in Central America and in most countries of tropical South
America. It is a genus of about 25 species.

Excluded Species:

Pentagonia magnifica Krause, Bot. Jahrb. Syst. 40: 325. 1908. type: Colombia. Lchmann 8886 (B,
not seen; NY, isotype; photo, MO).

Watsonamra magnifica (Krause) Standley, Contr. U.S. Natl. Herb. 17: 441. 1914.

When Standley effected the new combination in 1914, he remarked of Pittier
4148 (US) from the Rio Culebra, above Santa Isabel, Province of Colon, Panama:
''It is impossible to be certain that the Panama specimen is of this species, since
it is in fruit, and even the persistent calyces are mutilated so that their form
cannot be determined definitely." I have examined Pittier's collection and agree
with Standley. The solitary leaf of the Pittier collection measures 78 cm x 29 cm,
is not lobed, and is sessile. The lack of a petiole precludes it from being P,
macrophylla. The inflorescence is much reduced and branches in a deliquescent
fashion. The fruit is oblong, measuring 2 cm in width and about 3 cm in length;
the persistent calycine lobes are about 1 cm long.

a. Leaves entire, not pinnatifid lobed.

b. Leaves sessile 10. P wendlanJii

bb. Leaves obviously petiolate.

c. Subshrubs 1.0-1.5 m tall: leaves small, to 22 cm long; hypanthium and calyx to

6 mm long 9. P. veraguensis

cc. Shrubs or trees; leaves large, 25-45(-100) cm long.

d. Shrubs or small trees, to 7 m tall; fruits to 2.8 cm wide.

e. Corolla lobes 5.5-7.0 mm long; anthers dimorphic, the larger to 8.5

mm long 5. P. macrophylla

ee. Corolla lobes 10-12 mm long; anthers all alike ca. 4 mm long

__ 2. P. bocataurensis

dd. Trees to 17 m tall; fruits ca. 4 cm wide 6. P. nuciformis

aa. Leaves pinnatifid lobed.

f. Leaves subsessile (the blades often appearing petiolate as they narrow to a slender

wing on the costa) 8. P. tinajita

ff. Leaves obviously petiolate.

g. Petioles with conspicuous auricles.

h. Auricles ca. !4 the length of the petiole, not drying red, the margin un-
dulate; calycine lobes ca. 3 mm long 3. P. hrachyotis

hh. Auricles occupying ca. V2 the length of the petiole, usually drying red, the

margin erosulose or crisp, often with subulate processes; calycine lobes
4-7 mm long 7. P. pinnatifida

Petioles lacking auricles.

i. Petioles ca. 15 cm long; corolla tube ca. 45 mm long, the lobes ca. 0.6 mm

long L P. alba

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 321

ii. Petioles (3.0-)5.0-8.5 cm long; corolla tube ca. 20 mm long, the lobes ca,

10 mm long 4. P, gymnopoda

1. Pentagonia alba Dwyer.^^ type: Panama, Bristan 1224 (MO, holotype; isotype

NY).

Small shrubs, the stems not seen. Leaves (only one seen) pinnately lobed,
about 8 lobes on each side, the rachis between the lobes 1-3 cm wide, to 24 cm
long, to 7 cm wide, acute toward the apex, scarcely acuminate, the lateral veins
of each lobe ca. 8, arcuate, stiffly papyraceous, glabrous above except for the
costa, puberulent beneath; petioles to 15 cm long, 0.5 cm wide in the middle, the
base swollen, to 1.3 cm wide; stipules not seen. Inflorescences axillary, the few
flowers on a reduced peduncle. Flowers sessile; hypanthium ca. 0.5 mm long,
glabrous, the calycine cup expanded, ca. 0.7 cm long, glabrous with oblong glands
within arranged in rows, the lobes 5, oblong lanceolate, ca. 10 mm long; corolla
white, the tube funnel shaped, 4.5 cm long, ca. 0.3 cm wide at the base, 1.7 cm
wide at the apex, thinly carnose, glabrous outside, villose within at the point of
filament attachment, the lobes 5, oblong rotund to subtriangular, 0.6 mm long;
stamens 5, the anthers oblong, 2.5-3.0 mm long, the filaments 11-13 mm long,
unequal, 2 short, 2 intermediate in length, and one elongate; style (in bud) slender,
the stigmatic area oblong, the lobes 2. Fruits not known.

Pentagonia alba is known only from Panama. It is immediately distinguished
from all other Pentagonia of the Republic by the combination of pinnatifid lobed
leaf blades with a winged ''rachis" in the Interlobular areas and white flowers.

darien: Cerro Pirre, Bristan 1224 (MO, NY).

2. Pentagonia bocataurensis Dwyer,^^ type: Panama, Cooper & Slater 53 (F,
holotype).

IShrub, branches slender. Leaves entire, to 1 m long. Flowers with the hy-
panthium short, the calycine cup oblong, ca. 1.2 cm long, coriaceous, golden
puberulent outside, glabrous within and with clusters of glands within, the lobes
perhaps 5, unequal, triangular oblong to widely subulate, 0.4-0.8 mm long, acute,
ciliolate; corolla with the tube cylindrical, 3.5-5.0 cm long, puberulent outside,
glabrous within, coriaceous, the lobes 5, narrowly oblong, 10-12 mm long, often
acute; stamens 5, the anthers oblong, ca. 4 mm long, the filaments 1.8-2.0 mm
long, villose, especially below the middle; style linear, 30 mm long or longer, the
stigmas 2, oblong, ca. 5.5 mm long, ca. 2 mm wide, obtuse. Fruits not seen.

The new species is known only from Panama. The holotype was annotated

^^ Pentagonia alba Dwyer, spec. nov. Frutices parvi. Folia pinnatiformi-lobata, lobis angusto-
lanceolatis, ad 24 cm longis, ad 7 cm latis, lobis terminalibus 3, inferioribus 2 aequalibus apicem versus
acutis venis lateralibus lobi utriusque ca. 8; petiolis supra leviter angusto-alatis, ad 15 cm longis.
Flores sessiles, cupula calycis ca. 0.7 mm longa, lobis ca. 10 mm longis; corolla alba, ca. 4.5 cm
longa, lobis 5, ca. 0.6 cm longis; antheris 2.5-3.0 mm longis, filamentis inaequalibus.

^^ Pentagonia bocataurensis Dwyer, spec. nov. Folia ad 1 m longa. Flores cupula calycis ca.

1.2 cm longa lobis fortasse 5, 0.4-0.8 mm longis in margine ciliolata; corolla tubo 3.5-5.0 cm longo,
extus aureo-puberulo, lobis 5 angusto-oblongis, 10-12 mm longis acutis; staminibus 5, antheris ob-
longis, ca. 4 mm longis, filamentis brevibus, villosis. Fructus non visi.

322 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

by Standley many years ago as Pentagonia magnifica Lehmann, a species orig-
inally described from Lehmann 8886 from the Rio Timbique, Colombia. P. mag-
nifica, however, has 5 unequal calycine lobes which are obtuse at the apex while
in the new species the lobes are acute at the apex; P. magnifica has a much
shorter corolla, has shorter staminal filaments, linear anthers, and a much shorter
style.

BOCAS DEL TORo: Changuinola, Cooper & Slater 53 (F).

3. Pentagonia brachyotis (Standley) Standley, J. Wash. Acad. Sci. 17: 171. 1927.

Watsonamra brachyotis Standley, Contr. U.S. Natl. Herb. 17: 443. 1914. type: Panama, Williams
999 (F, holotype).

Shrubs to small trees, 0.75-3.00 m tall, the stems glabrous, lenticellate. Leaves
pinnatifid lobed, the lobes 4-6 per side, the entire blade rhombic, 33-70 cm long,
to 42 cm wide, cuneate at the apex, the lateral lobes 4-9 cm wide, cuneate and
ultimately acute or obtuse at the apex, the blades subtruncate to broadly cuneate
at the base, the interlobular areas or sinuses 0.2-4.0 cm, thinly coriaceous or
papyraceous, glabrous to puberulent; petioles to 14 cm long, conspicuously au-
riculate at the base, the auricles rounded, 4-7 cm long, membranous, the margin
undulate, red; stipules ovate oblong, to 3.5 cm long, to 1.2 cm wide, acute,
appressed puberulent on the outside. Inflorescences axillary, ca. 4 cm wide, the
flowers congested, the peduncle much reduced. Flowers 5-merous, sessile; hy-
panthium 0.5 cm long, appressed pubescent, the calycine cup subcampulate, to
2 cm long, 1 cm wide, appressed pubescent outside, glabrous within, with rows
of oblong glands below the sinuses, the lobes widely triangular or oblong, to 3
mm long, ciliolate; corolla tube cylindrical, to 3 cm long, 0.5 cm wide, glabrate,
coriaceous, the lobes triangular, to 0,3 cm long; anthers subsessile, oblong, 3.5
mm long, attached near the mouth, the filaments pilose beneath; style 1.5-2.0 cm
long, the stigmas oblong rotund, ca. 3 mm long. Fruits sessile, ovate to ovate
rotund, to 3 cm long, to 2 cm wide, truncate, drying red brown, conspicuously
lenticellate, minutely rugulose, the calycine cup barrel shaped, to 0.8 cm long,
the 5 narrowly oblong lobes slightly longer than the tube, glabrous to puberulent
outside.

Pentagonia brachyotis is known only from Panama. Elias 1785 (MO) from the

Rio Mulatupo, San Bias, is probably P. brachyotis. It is in fruit. The fruits are
sessile, rotund, to 2 cm in diam., lignose, lenticellate, and the persistent calyx is
1.8 cm long. On this collection the petioles are not intact in the specimen and no
auricular appendages are visible.

colon: Santa Rita Ridge Trail, 17-35 km from the Boyd-Roosevelt Highway, 400-800 m, Mori
& Kalhmki6349 (MO), darien: Summit of Cerro Pirre, 1000-1400 m. Gentry & Clewell 7046 A (MO).
Cana-Cuasi Trail (Camp 2), Chepigana, Terry & Terry 1426 (F, MO). Marraganti, Williams 999 (F).
Panama: Stream flowing out of Serrania de Maje, Folsom & Collins 1714 (MO), san blas: Rio Chico,
Yaviza, Bnrch et al. 11241 (F, MO, US). Perme, Cooper 252 (F, US).

4. Pentagonia gymnopoda (Standley) Standley, J. Wash. Acad. Sci. 17: 171. 1927.

Watsonamra gymnopoda Standley, Contr. U.S. Natl. Herb. 17: 444. 1914. type: Panama, Pittier
3858 (US, holotype; NY, isotype).

1980]

DWYER— FLORA OF PANAMA (Family 179, Ruhiaceoe)

323

B

Figure 68. Penta^onia macrophylla Benth. — A. Leaves (x'/4). — B. Leaf venation (x IVi). — C.
Inflorescence (xVi). — D. Flower, part of perianth removed (xlVi). — E. Ovary, longitudinal section
(x3). [After Croat 16542.]— V. Fruit [xVi). [After Duke 4270.]

324 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

Shrubs or trees to 4 m tall, the branches lignose, puberulent, the leaves crowd-
ed at the tips. Leaves in bud bursting out of the terminal stipules, oblong, to 1
m long or longer, to 70 cm wide or wider, deeply pinnatifid lobed, the lobes 6-

13, the terminal portion 3-lobed, the median lobe either ovate oblong or obovate
oblong, as long as the 2 lateral lobes, the latter oblong, usually narrowly so, 5-

25(-35) cm long, 2-8(-9) cm wide, cuneate at the apex, 2-8 cm apart, the sinuses

subacute or rounded, extending almost to the rachis, the rachis and the winglike

portion of the blade 0.5-3.0 cm wide, the costa prominulous above, prominent

beneath, to 0,25 mm wide, minutely pubescent, the secondary veins 6-9, usually
one for each lobe, the branches of the secondary veins ca. 6 per lobe, thickly

herbaceous to stiffly chartaceous; petioles (3.0-)5.0-8.5 cm long, subterete, lack-
ing auricles, dilated proximally; stipules ovate lanceolate, to 6.5 cm long, to 2.5
cm wide, acute, coriaceous, strigillose orglabrate outside. Inflorescences densely
flowered, the peduncle to 1 cm long; bracts oblong, to 3 cm long, strigillose and
ciliate. Flowers sessile; hypanthium ca. 1 cm long, the hairs appressed, dense,

the calycine cup somewhat expanded, 0.5-0.7 cm long, coriaceous, tuberculate
outside, minutely puberulent outside, glabrous within, with a few oblong glands

at the base, the lobes 5, oblong rotund, somewhat unequal, 8-10 mm long, to 10
mm wide, glabrous within, scarious margined, ciliolate; corolla white, yellow or
pale green, the tube narrowly cylindrical, ca. 20 mm long, coriaceous, puberulent
outside, densely bearded within at the point of filament attachment, the lobes 5,
oblong, ca. 10 mm long, glabrous within; stamens 5, the anthers oblong, ca. 3
mm long, the filaments unequal, 8-10 mm long, 2 short, 1 intermediate, and 2
long, bearded basally, attached ca. 8 mm from the base of the tube; style 9 mm

long, the stigmatic area ca. 4 mm long, the stigmatic lobes unequal, concave, 1
narrow, ca. 0.8 mm wide, the other ca. 1.5 mm wide, both pubescent adaxially.
Fruits sessile, ovate, pear shaped or subrotund, to 2 cm long, usually truncate
at the apex, densely lenticellate, the persistent calyx to 2 cm long, the base of
the calycine tube set within a delicate ringlike scar.

Nee

1385 report the fruit as purplish brown.

colon: Loma de Gloria, Fato, PiUier 3858 (BM, NY, US), darien: Airstrip at Cana gold mine,
Croat 37986 (MO). Panama: El Llano-Carti Road near Gorgas Mosquito Control Project at km 12,
Croat 26026 (MO). Cerro Jefe, Duke 9354 (MO). Cerro Azul, Dwyer 1385 (MO, US), 7072 (MO).
Cerro Jefe, 3100 ft. Gentry 3508 (MO). Cerro Azul, 2-3 mi S of Goofy Lake, Lewis et aL 292 (GH,
MO, US). Cerro Jefe and La Eneida, Luteyn & Foster 1130 (MO). El Llano-Carti Road, Mori &
Kallunki 2856 (MO). EI Llano-Carti Road, ca. 10 km from the Panamerican Highway, 350 m, Mori
& Kallunki 3535 (MO); Nee 10528 (MO).

5, Pentagonia macrophylla Benth., Bot. Voy. Sulph. 105. tab, 39, 1845, type:
Panama, Bentham (K, holotype, not seen). — Fig. 68.

Watsonamra macrophylla (Benth.) Kuntze, Rev, Gen. PI. 302. 1891.

W. pubescens Standley, Contr. U.S. Natl. Herb. 17: 441. 1914. type: Panama, Pittier 3822 (US,
holotype; NY, isotype).

Trees to 7 m ( or to 16 m?), the nodes crowded at the apex of the stem. Leaves
ovate rotund, oblong, or rhombic rotund, to 45 cm long, to 24 cm wide, widely

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 325

deltoid at the apex, basally cuneate, rounded or occasionally truncate, often
inequilateral, the costa prominent above and beneath, the lateral veins 12-15,
coriaceous, glabrous above, minutely sericeous on the veins beneath; petioles to
14 cm long, 0.3-0.5 cm wide in the middle, not strongly dilated basally; stipules
free, triangular, 3.5-4.0 cm long, ca. 1.5 cm wide at the base, coriaceous, glabrous
or golden puberulent. Inflorescences axillary, sessile or pedunculate, the cymes
with few to many flowers aggregated usually into a rotund mass shorter than the
petiole; bracts red. Flowers yellow; hypanthium obovate rotund, 0.6-0.8 cm long,
coriaceous, glabrous, the calycine lobes 5, erect, oblong, 0.6-0.9 cm long, obtuse,
concave, coriaceous, moderately pubescent, with a cluster of minute, oblong
glands within at the sinuses; corolla with the tube cylindrical, 3,5-4.0 cm long,
to 1 cm wide, puberulent outside, densely golden villose within near filament
attachment, the lobes 5, spreading or scarcely reflexed, ovate oblong, 5.5-7.0
mm long, acute, villose marginally, smooth; stamens 5, included in the tube, the
anthers perhaps dimorphic, the smaller saucer shaped, ca. 3.5 mm long, the pollen
aborted?, the larger to 8.5 mm long, polleniferous, the filaments linear subulate,
1.2-2.0 cm long, differing slightly in length, ca. 1.3 mm wide at the base, attached
below the middle of the tube, densely villose basally on the adaxial side; style
linear, 2.0-2.5 cm long, ca. 0.8 mm wide, the stigmas 2, spread at anthesis, oblong
rotund, 3-4 mm long, obtuse. Fruits rotund, to 2.8 cm in diam., nutlike, villo-
sulose, the calycine cup 1.5-2.0 cm long, the lobes lanceolate, narrow oblong or
subrotund, to 1.3 cm long, equal to or exceeding the cup in length, the wall of
the fruit ca. 0.35 cm in diam.

Pentagonia macrophylla is known only from Panama, the Dept. Choco, Co-
lombia, and Ecuador. ''Hoja de Murcielago." Folsom & Robinson 2381 has ex-
ceptionally small leaves measuring only 19-25 cm in length.

BOCAS DEL TORo: Qucbrada Huron, Kirkhhde & Duke 425 (MO). Isla Colon, Lao 93 (MO);
Wedel 481 (MO). Chiriqui Lagoon, Wedel 1561 (F, MO). Old Bank Island, Wedel 1931 (F, MO).
Chiriqui Lagoon, Wedel 2535, 2873 (both MO, US), canal zone: Frijoles, Allen 926 (MO). Rio
Chagres, Allen 874 (MO). Juan Mina, Bartlen & Lasser 16555 (MO). Without other locality, Blum
& Tyson 2322 (MO). Pipeline Road, Croat 12364 (MO). Road between Gatiin and Pina, 3 km S of
Pina, Croat 36931 (MO). Coco Solo, Duke 4270 (MO). Galena Point, Dwyer & Robyns 161 (MO).
Gatiin Lake, NW shore, Dwyer 1816 (GH, MO, US). Summit and Gamboa, Greenman & Greenman
5217 (MO). Pipeline Road, Hayden 69 (MO). Rio Indio, Lao & Holdrid^e 54 (MO). Pipeline Road,
Lewis et al. 5458 (MO, SCZ, UC). Fort Sherman, Maxon & Valentine 6987 (US). Railroad, 6 km W
of Gamboa, Nee 7206 (MO). 2 km W of Gamboa, Nee 7531 (MO). Pipeline Road, V/i km NW of
Gamboa, Nee 9584 (MO). Gatuncillo, Piper 5602 (US). Tabernilla, Pittier 3822 (F, US). Agua Clara,
Pittier 3992 (NY, US). Frijoles, Standley 27421, 27497. 27565 (all US). Gamboa, Standley 28422
(US). Fort Randolph, Standley 28691 (MO, US). Fort Sherman, Standley 31034 (US). Obispo, Stan-
dley 31696 (US). Fort Sherman, Tyson 3809 (MO); Tyson & Blum 1667 (MO), barro Colorado
island: Bailey cfe Bailey 135 (F); Ban^ham 440 (F); Brown 135 (F); Croat 4128, 4646, 4685, 5453,
5821, 6220, 6244, 8167, 10355, 14580, 16542 (all MO); Ehinger 58 (MO, US); Foster 1052 (F); Havden
11, 32 (both MO); Kenoyer 545 (US); Knight 69-3 (MO); Luteyn 1350 (F); Maxon et al. 680 (MO);
Oppenheimer 6609-14-1305 (MO); Robyns 65-15 (MO); Shattuck 1049 (MO); Standley 31328, 31455,
40789 (all US); Starry^ 21 (MO), 142 (F, MO); Woodson & Scherv 963 (MO), cocle: El Valle, Allen
2801 (F, MO), 3736 (F). Alto de Calvario, 7 km N of El Cope, 700 m, Folsom 4097 (MO); Folsom
& Robinson 2381 (MO). Boca del Toabre, Lewis et al. 5542 (BM, MO), colon: Camp Pifia, Allen

3580 (F). Rio Indio, road from Portobelo to Nombre de Dios, Croat 33633 (MO). Icacal between
Salud and Boca, Howell 234 (MO). Pina, Kirkbride <S Hayden 318 (MO). Fort San Lorenzo, Quist-

gaard 26 (MO). Road to Ri'o Indios, 3 km S of Piiia, Sullivan 130 (MO), darien: Cerro Pirre, Bristan
557 (MO); Duke 4842 (MO). Panama: Chagres, Fendler 208 (MO). Cerro Trinidad, Kirkbride & Duke
1678 (MO). 5-10 km NE of Altos de Pacora, ca. 750 m, Mori & Kallunki 4978 (MO). El Llano Carti

326 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

Road, 5 km N of Panamerican Highway, Nee 7935 (MO). Arenoso, Seihert 599 (MO), san blas:
Headwaters of Rio Cuadi, Camp Diablo, Duke 3649 (MO, UC).

6. Pentagonia nuciformis Dwyer.®^ xype: Panama, Croat 23093 (MO, holotype).

Trees to 17 m tall, the twigs terete, ultimately angular, the nodes moderately
crowded; petiole scars oblong or obovate rotund, to 1 cm long. Leaves oblong,
entire, 15-30 cm long, 5-16 cm wide, widely rounded or widely deltoid toward
the apex, scarcely acuminate or with the acumen to 1 cm long, the costa prom-
inulous above and ridgelike, prominent beneath, the lateral veins 12-14, at first
strict, then arcuate toward the margin, the intervenal areas lineolate, thinly co-
riaceous, glabrous; petioles 2-4 cm long, slightly dilated at the base, glabrous;
stipules free, deltoid oblong, to 4 cm long, to 2 cm wide, acute, glabrous, cori-
aceous, black when dry, with 2 well-spaced proximal keels converging toward
the apex. Flowers not seen. Fruits ca. 15, axillary, oblong or subrotund, often
angular, to 4 cm long, to 3 cm wide, nutlike, lignose, glabrous but with minute,
brown scales, deciduous, the persistent calycine cup 0.3-0.5 cm long with a few
glands within, the glands to 5 mm long, the lobes 5, often withered, oblong, equal
to or longer than the tube, the pericarp lignose, to 4 mm thick, smooth and lightly
reticulate within; seeds abundant, disposed as 2 oblong rotund clusters when dry,
to 2 cm long, the individual seeds oblong, ca. 7 mm long, angular, shiny, minutely
honeycombed in appearance.

Pentagonia nuciformis is known only from Panama. The height of the tree
and the large fruits immediately distinguish it from all other Central American
species of the genus. Croat (pers. comm.) considers it strikingly different from
the several species of Pentagonia with which he is familiar in the field; he points
out that the tree branches freely. This is noteworthy as Pentagonia usually has
a simple trunk.

VERAGUAs: 5 mi W of Santa Fe on the road past Escuela Agricola Alto Piedra, Pacific side of
divide, 800-1200 ft, Croat 23093 (MO).

7. Pentagonia pinnatifida Seem., J. Bot. (Hooker) 7: 566. 1848. type: Panama,
Seemann 1019 (BM, not seen, photo MO).

Pentagonia pittieri (Standley) Standley, J. Wash. Acad. Sci. 17: 170. 1927.

Watsonamra pittieri Standley, Contr. U.S. Natl. Herb. 17: 443. 1914. type: Panama, Pittier 4298
(US, holotype; F, NY, isotypes).

Shrubs to 3 m tall, the stems terete, lignose, glabrous. Leaves crowded ter-
minally, oblong, to 1 m or more in length, deeply lobed and pinnatifid, the indi-
vidual lobes narrowly oblong, to 45 cm long, to 10 cm wide, tapering deltoidly.

^'^ Pentagonia nuciformis Dwyer, spec. nov. Arbores ad 17 m altae. Folia oblonga, 15-30 cm

longa, 5-16 cm lata, versus apicem lato-rotunda vel late deltoidea vix acuminata vel acumine ad 1 cm

longo praedita, venis lateralibus 12-14, tenui-coriacea glabra; petiolis 2-4 cm longis; stipulis liberis

deltoideo-oblongis, ad 4 cm longis, basi 2 cm latis. Fructus ca. 15 in subrotundum fasciculum infra
verticillos terminales foliorum congesti oblongi vel subrotundi, ad 4 cm longi, ad 3 cm lati, nuciformes

lignosi glabri sed squamis brunneis minutis deciduis praediti cupula calycis persistente ca. 0.5 cm
longa, intus glandulis, ad 0.5 mm longis praedita, lobis 5 saepe marcidis tubo aequalibus vel quam longi-

oribus, pariete fructus ad 4 cm crasso; seminibus multis in massas duas oblongo-rotundas aggregatis.

1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 327

the apex of the blade 3-lobed, the median of these ovate oblong, ca. 3 times
longer than the lateral pair of apical lobes, the costa prominent above and be-
neath, convex beneath, to 0.6 cm wide, the lateral veins usually 1 per lobe, the
secondary veins simulating lateral veins in each lobe, ca. 7, arcuate, prominulous,
the finest veinlets lineolate, thinly coriaceous, glabrate above, puberulent on the
veins beneath; petioles to 17 cm long, probably longer in the larger leaves, lignose,
dilated proximally, glabrescent, with large foliaceous basal auricles, subrotund,
to 7.5 cm long, ca. 8 cm wide basally, fleshy and red, the margin erosulose or
crisp, often with marginally subulate processes, to 2 cm long, ca. 0.1 cm wide,
minutely puberulent; stipules free, lanceolate to triangular subulate, to 5.5 cm
long, sericeous outside. Inflorescences axillary, to 6 cm long, the cymules densely
flowered. Flowers with the hypanthium ca. 5 mm long, sericeous, the calycine
cup subcampanulate, 3-4 times the length of the hypanthium, constricted basally,
thinly coriaceous, minutely sericeous outside, glabrous within, the lobes oblong,
4-7 mm long, obtuse; corolla tube ca. 2.5 cm long (fide Standley), appressed
pilose. Fruits sessile, orange, rotund, ca. 2 cm in diam., minutely pubescent to
glabrate, minutely rugulose but not scaly, the persistent calyx as long as or longer
than the fruit.

Pentagonia pinnatifida is known only from Panama and Colombia. It is easily
distinguished from other Pentagonia of Panama except P, brachyotis by its pin-
natifid leaves and petioles bearing large foliaceous auricles. Among the Choco
the fruits are eaten, and the leaves and fruits are ''taken in a tea as a blood
purified' (fide Duke). '^Biruchichiho/^

darien: Rio Chico, from Yaviza at Rio Chucunaque to ca. I hour by outboard from junction,
Burch et al. 1124 (F, GH, K, MO, NY, UC, US). Rio Pirre, Croat & Porter 15501 (MO). 2-3 mi SE
of El Real, Duke 4842 (MO). 10 mi S of El Real on Rio Pirre, Duke 5468 (MO). Quebrada Bidoto
(Peccary Creek) off Rio Areti, Duke 13599 (MO), Quebrada '^Camachimuricate," Casa de Bartolo,
Kennedy 2857 (MO). Manene to Rio Cuasi, Kirkbride & Bristan 1401 (MO). From Pinogana to Yaviza,
Stern et al. 136 (MO, US). Panama: Rio Maje, above first waterfall, 50 m, Croat 34419 (MO). Trail
between Ri'o Maje and Quebrada Brava, 2 mi upstream from waterfalls, Bayano Lake, Croat 34740
(MO). SAN BLAS: Hill SE of Puerto Obaldia, Croat 16733 (MO). Mainland opposite Ailigandi, Lewis
et al. 187 (GH, MO, US). Puerto Obaldia, Pittler 4298 (F, US). Colombia: choco: Region Guayabal,

5 hours by foot to SE of Acandi, Forero et al. 971 (MO).

8. Pentagonia tinajita Seem., Bot. Voy. Herald 134. 1854. type: Panama, David,
Seemann 1595 (BM, not seen, photo MO).

Watsonamra tinajita (Seem.) Kuntze, Rev. Gen. PI. 302. 1891.

Shrubs to 3(-4) m tall. Leaves subsessile, pinnatifid lobed, 0.75-1.00 m tall,
to 0.35 m wide, acute or rounded at the apex of the blade, cordate at the base,
the sinuses not rounded, coriaceous, glabrous to pubescent; petioles short, the
lamina decurrent (Standley: 'lower portion of the blade petiolelike 8-16 cm
long''); stipules ovate, to 4 cm long, acute. Inflorescences with the cymules
densely flowered, the bracts triangular, elongate, acute. Flowers with the hypan-
thium rotund, ca. 5 mm long, the calycine cup turbinate, to 15 mm long, the
calycine lobes oblong, shorter or about as long as the cup, obtuse; corolla green,
cylindrical or infundibuliform, the tube ca. 2.5 cm long, the lobes 5, triangular or
ovate, 6-7 mm long. Fruits rotund, 1.0-1.7 cm in diam., lenticellate; seeds to 3
mm long, smooth.

328 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

Pentagonia tinajita is known only from Panama. As Hooker (1849) points out
concerning the specific name: "... (it) is taken from the fruit which resembles
the water jars (tinajas) used in the country." There is a detailed figure accom-
panying Seemann's original diagnosis.

CHiRiQui: David, Pittlcr 3369 (US). San Felix, Pittier 5214 (US).

9. Pentagonia veraguensis Dwyer.^^ type: Panama, Croat 27504 (MO, holotype).

Suhshruhs to 1.5 m tall, unbranched, the stem terete, glabrous, the nodes well
spaced, the petiole scars hemispherical or oblong, 0.5-1.2 cm long. Leaves rhom-
boid, 16-33 cm long, 6-16 cm wide, acute or deltoid toward the apex, acuminate,
the acumen scarcely developed, cuneate or acute toward the base, the costa
prominulous above, prominent beneath, the lateral veins 12-14, rigid, not arcuate,
forking near the margin or with 1 or 3 small veins near the margin, the submarginal
undulate vein lacking, the intervenal areas minutely lineolate, glabrate above,
puberulent on the costa and the veins beneath, rigidly papyraceous, slightly dis-
colorous; petioles 2.5-6.0 cm long, those in the middle ca. 0.3 cm wide, puber-
ulent and twisted; stipules ovate lanceolate, to 3 cm long, to 1.1 cm wide at the
base, subcoriaceous, glabrous, with 2 keels on the back toward the base, well
spaced but converging toward the apex, glanduliferous within toward the base,
the small glands oblong, to 1.5 mm long. Inflorescences axillary, the flowers few,
the peduncle shorter than the flowers and with short scales; bracts red, crowded,
deciduous, numerous, imbricate, perhaps binate, narrowly oblong, to 1.8 cm long,
to 0.8 cm wide, scarious on the back, puberulent or glabrate, ciliate, glabrous
within; bracteoles beneath the hypanthium oblong rotund, 8-12 mm long, oth-
erwise like the bracts. Flowers with the hypanthium ca. 2.5 mm long, white
puberulent, the calycine cup cylindrical, 3.5-10.0 mm long, densely minutely
pubescent or glabrate within below the sinuses, the lobes 5, ovate oblong, ca. 8
mm long or longer in fruit, obtuse, glabrous within, coriaceous; corolla white or
light yellow; stamens 5, the anthers oblong, to 2.5 mm long, the filaments slender,
9.0-1 1.5 mm long, villose at the base, arising ca. 7 mm from the base of the tube;
style slender, ca. 15 mm long, the stigmas widely oblong, ca. 2.5 mm long, piano
compressed, obtuse. Fruits sessile, oblong to rotund, to 3.5 cm long, yellow
toward the base, at first rugulose then moderately smooth, the calyx red, at first
persistent, to 2.5 cm long, ultimately wrinkled except for the calycine cup.

Pentagonia veraguensis is known only from Panama. It is characterized by
its small size and its small rhombic leaves, although a recent indeterminate col-
lection by Mori & Kallunki (6184) from Escuela Agricola Alto de Piedra, Vera-
guas, resembles this species. It is in fruit and is described by the collectors as a
"slender unbranched tree, 4 m tall, 4 cm dbh." Its leaves measure to 40.5 cm
and are deltoid at the apex. Unfortunately flowers were not collected.

^Pentagonia veraguensis Dwyer, spec. nov. Siiffriitices ad L5 m, ramis deficientibus. Folia

subrhomboidea, 16-33 cm longa, 6-16 cm lata, apicem versus acuta vel deltoidea venis lateralibus

12-14, subtus in costa venisque puberula; petiolis 2.5-6.0 cm longis; stipulis ovato-ianceolatis. ad 3

cm longis, extus bicarinatis. Inftorescentiae floribus breviores. Flores cupula calycis 2.5-10.0 mm

longa, lobis ca. 8 mm longis; corolla alba vel lutea; antheris ca. 2.5 mm longis. Fructiis oblongi, ad
3.5 cm longi.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhimeae) 329

VERAGUAS: Valley of Ri'o Dos Bocas, 11 km from Escuela Agricola Alto Piedra above Santa Fe
on road to Calovebora, 450 m, Croat 27504 (MO); Mori et al. 3878 (MO). 16 km NW of Santa Fe,
road to Calovebora, 300-500 m, Mori & KaUimki 6128 (MO).

10. Pentagonia wendlandii Hook., Bot. Mag. pL 5230, 1861. type: not known.

Watsonamra wendlandi (Hook.) Kuntze, Rev. Gen. PI. 320. 1891.

Shrubs to IVi m tall, the stem unbranched, smooth. Leaves sessile, entire,
oblong obovate, to 1 m long, to 50 cm wide, rounded or widely deltoid at the
apex, tapering gradually to base, ultimately auriculate, each auricle ca. 2.5 cm
wide, the costa prominulous above, prominulous to prominent beneath, the lateral
veins ca. 18, prominulous above and beneath, coriaceous, glabrous to puberulent
beneath, purple beneath; stipules ovate oblong or triangular, 5.0-6.5 cm long,
puberulent outside. Inflorescences axillary, few flowered; bracts ovate lanceo-
late, to 1 .2 cm long, obtuse, pubescent. Flowers sessile or subsessile, the pedicels
to 1 cm long; hypanthium short, the calycine cup campanulate, ca. 1 cm long,
purplish red, stiffly petaloid, pubescent outside, rugulose within, usually with
numerous oblong glands within the cup near the sinuses, occasionally with an
additional gland at the sinus, the calycine lobes 5, erect, somewhat unequal, most
about V2 the length of the cup, the longest almost as long as the cup, often 1 lobe
conspicuously short and acute; corolla yellow, the tube narrowly cylindrical, to
2.5 cm long, thickly carnose, glabrous to puberulent outside toward the apex,
villose within only near point of filament attachment, the lobes triangular to ob-
long, 4-7 mm long, 2.2-2.6 mm wide; stamens 5, the anthers oblong, 4 mm long,
obtuse, the filaments unequal, to 16 mm long, to 0.8 mm wide, densely villose at
the base; style ca. 18 mm long, glabrous, the stigmas oblong, 3.5 mm long, obtuse,
plane, the ovarian disc compressed cylindrical, ca. 2 mm long, thin walled, trun-
cate, marginally undulate. Fruits oblong rotund, to 4.5 cm long, drying black,
puberulent, the persistent calyx to 1.5 cm long.

This species has been reported only from Panama and Costa Rica. It is ques-
tionable whether the type exists; the species was ''described from cultivated
plants of Central American origin.'' The species is probably limited to Panama
and Costa Rica. The flowers are described as yellowish-green, red, and yellow;
the calyx is described as purplish-red or red-pink. Dressier 4362 notes that the
leaves are pink-red beneath. Croat 33632 (MO) has a closeup (photograph) of the
leaves and fruits attached to the herbarium sheet.

BOCAS DEL TORO: Changuinola Valley, Dunlap 187 (F, US). Qucbrada Huron, Rapid Streams,
Kirkhride & Duke 425 (MO). Chiriquicito, 5 m S along Rio Guaramo, Lewis 2101 (GH, MO, US).
Old Bank Island, Wedel 2018 (MO), colon: Rio Indio, road from Portobelo to Nombre de Dios. 50
m, Croat 33632 (MO). Road to Portobelo, 2 km NE of Puerto Pilon, Nee & Tyson 10918 (MO).
PANAMA: El Llano-Carti Highway, 12-15 km N of El Llano, Dressier 4362 (MO). EI Llano-Carti
Road, 9.6 km from Panamencan Highway, ca. 350 m, Mori & Kallunki 5602 (MO).

65. POGONOPUS

Pogonopus Klotzsch, Monatsber. Konigl. Preuss. Acad. Wiss. Berlin 1853: 500
1853. type: P. ottonis.

Chrysoxylon Wedd., Monog. Cinch. 100. 1849, not Chrysoxylon Casar. 1843. type: C. fehrifu^um

Wedd.
Howardia Wedd., Ann. Sci. Nat. (Paris) Ser, 4. 1: 166. 1854, not Klotzsch, I860, type; Not specified

330

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 67

B

Figure 69. Po^onopus speciosus (Jacq.) Schum. — A. Habit with flowers (x!/2). [After Croat
12465.]— B, Fruit (x2). [After Croat 8959.]

Trees or shrubs. Leaves with the stipules small, cuspidate. Inflorescences
terminal, paniculate, the cymes small. Flowers pedicellate, the pedicels bracteo-
late; hypanthium turbinate, the calycine cup short, 5-dentate, one of the lobes
disposed as a large stalked foliaceous blade; corolla tubular, villous within, the
lobes 5, valvate; stamens 5, the anthers slender, versatile, exserted,the filaments
attached to the wall of the corolla tube; ovarian disc thick, annular, the style
slender, the stigmas 2, obtuse, the ovary 2-celled, the ovules numerous, longi-

1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 331

tudinally adnate to the septum. Fruits capsular, woody, loculicidally dehiscent
into 2, many-seeded valves; seeds horizontal, the testa thick.

Pogonopus occurs in Central America and tropical South America. It is a
genus of about 3 species.

1. Pogonopus speciosus (Jacq.) Schum., Mart. Fl. Bras. 6(6): 265. 1889. — Fig. 69.

Macrocnemum speciosus Jacq., Hort. Schoenbr. 1: 19. 1797. type: Not given.

Pogonopus ottonis Klotzsch, Monatsber. Konigl. Preuss. Acad. Wiss. Berlin 500. 1853. type: Otto
901 (B).

Howardia grandiflora Wedd., Ann. Sci. Nat. (Paris) Ser. 4, 1: 70. 1854. type: Schlim 755 (P).

Howardia caracasensis Wedd., Ann. Sci. Nat. (Paris) Ser. 4. 1: 71. 1854. type: Funck 463 (P).

Pogonopus caracasensis (Wedd.) Nichols, Diet. Gard. 3: 175. 1886.

Carmenocania porphyrantha Wernh., J. Bot, 50: 241. 1912. type: Schlim 755 (BM).

Trees to 10 m tall, the branchlets terete, smooth, glabrous to glabrescent,
often with fusiform white lenticels. Leaves oblong, rarely obovate oblong, 15-
21(-30) cm long, 6.0-11.5 cm wide, 2 to 3 times longer than wide, usually acute,
shortly acuminate, the acumen 1.5-2.0 cm long, ultimately subobtuse to acute,
basally attenuate acute to cuneate, equilateral, the costa slender, subplane to
prominulous above, usually prominent beneath, the lateral veins 10-12, the me-
dian veins 1.5-2.0 cm apart, arcuate, papyraceous, slightly discolorous; petioles
minute or 1-3 cm long, wiry, to 0. 1 cm wide, often curved, glabrous to puberulent;

stipules compressed rectangular, much wider than long, to 3 mm long, the apic-
ulum ca. 1 mm long, densely golden pubescent. Inflorescences terminal and ax-
illary, usually exceeding the uppermost leaves, thrysoid cymose, spreading, gla-
brous, to 25 cm long, to 15 cm wide, either sessile with 2 or 3 branches from the
stem tip or with peduncles 5-10(-13) cm long, wiry, often plane and twisted, the
primary branches few, the pairs 2-4, opposite, 2-3 cm apart in the larger inflo-
rescences, strict or arcuate, strongly ascending, the cymules terminal, flabellately
disposed. Flowers with the hypanthium oblong or turbinate, ca. 3 mm long,
puberulent, constricted below the calycine cup, the latter ca. 1 mm long, turgid,
eglandular within, 4 of the calycine teeth subulate, to 1 mm long, the 5th expanded
as a flaring foliose bract-like structure, ovate, oblong rotund or trapeziform ro-
tund, 1-5 cm long, to 4 cm wide, acute to deltoid at the apex, occasionally vaguely
acuminate, rounded or cuneate at the base with 5-7 conspicuous veins, glabrous
or puberulent, the stalk slender, 1.5-2.0 cm long; corolla red or pink, the tube
narrowly cylindrical, ca. 25 mm long at anthesis, constricted medially, stiffly
petaloid above, subcoriaceous on the lower third, puberulent outside and in ex-
cept villose inside at the filament attachment point, the lobes 5, triangular, ca. 4
mm long, erect; stamens 5, the anthers oblong, 1.5-2.5 mm long, the filaments
linear, to 28 mm long, plane, often exserted to 3 mm above the mouth, attached
ca. 8 mm from the base of the tube, the style linear, usually slightly exceeding
the tube, puberulent, the stigmas coiled, ca. 2 mm long. Fruits capsular, oblong
rotund, 5-7 mm long, to 6 mm wide, usually truncate, drying black, glabrous,
conspicuously lenticellate, the furrow usually conspicuous; seeds round or ob-
long, ca. 0.4 mm in diam.

Pogonopus speciosus ranges from Mexico to Colombia and Venezuela.
^'Chorcha de Gallo' ; ^'Madroiio \ '^Chibigui.^'

332

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 67

B

J ^^/LtMM-

Figure 70. Posoqueria latifolki (Rudge) R. & S.— A. Habit (x!/2).— B. Flower (x 1/2).— C. Sta-
mens (x2>/2).— D. Ovary, cross section (x3).— D. Fruit (xVi). [After Allen 955.]

CANAL zone: Fort Kobbe, Allen 2017 (MO). Cerro Viejo on K16C, Blum 1260 (MO). 1 mi S of
Madden Dam, Blum 2629 (MO). Madden Forest, Road 2, Croat 8959 (MO). Road S-ll, NW of
EscobaK Croat 12465 (MO). Gaillard Cut NE of Paraiso, Croat 12511 (MO). 1 mi N of Paraiso, Croat
12652 (MO). Near Salamanca Hydrographic Station, Dod^e et al. 16977 (MO). Madden Dam area,
Dnyer 8372 (MO). Barro Colorado Island, Foster Mil (MO). Madden Dam, Lewis et al. I (GH,
MO, UC, US). Forest Preserve Road, 1 km from Madden Dam, Mori 421 (MO). Road C-29, 6 km
ESE of Gamboa, Nee 8846 (MO). Barro Colorado Island, Slmttuck 688 (MO). Road K-15 near Huile,
Smith et al. 3282 (MO), colon: 3 mi E of Transisthmian Highway, road to Salamanca, Gentry 6720

1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 333

(MO). 4 km ENE of Buena Vista, road to Sardinilla, A'^^^ 9125 (MO). Darien: Cabecero del Rfo
Pirre, Brlstan 1330 (MO). Rio Aruza, Bristan 1346 (MO). Rio Yape, Bristan 1425 (MO). Pucro to Rio
Pucro, Duke 5361 (MO). Rfo Sambii, 0.5 mi above Rio Venado, Duke 9290 (MO). Rio Tuira between
Rfo Punusa and Rfo Mangle, Duke 14597 (MO). Rfo Pirre, Duke & Bristan 8264 (MO). Rfo Pucro
between Cerro Mali and Cerro Tacarcuna, Gentry & Mori 13874 (MO). Rfo Piedras, Stern et al. 724
(MO). Natural Bridge, Madden Lake, Croat 12400 (MO). Panama: Rfo Chagres, 1 mi above Madden
Lake, Duke 4465 (MO). Panamerican Highway, 28 km E of Bayano Bridge, Folsom 1387 (MO).
Stream flowing out of Serrania de Maje, Folsom & Collins 1729 (MO). Ipeti, base of Serrania de
Maje, Folsom & Lantz 1931 (MO).

66. POSOQUERIA

longift

Solena Willd., Sp. PL 1: 961. 1797. type: 5. longiflora (Aubl.) Willd. = Posoqueria hmgiftora Aubl.
Stannia Karst., Ausw. neuer schonbl. Gew. Venez. 27. 1848. type: S. grandifiora Karst., S.formosa
Karst., or S. metensis Karst.

Trees or shrubs. Leaves usually large, coriaceous; stipules free, subulate to
broad. Inflorescences cymose corymbose to umbellate; bracts minute. Flowers
zygomorphic, 5-merous, the buds curved, the calyx 5-lobed, the lobes somewhat
imbricate, auriculate at the base, glandular on the back; corolla infundibuliform
or hippocrateriform, the tube elongate, glandular papillose outside, the lobes
unequal, ascending, spreading or reflexed, imbricate; stamens unequal, inserted

at the mouth, the anthers slender, apiculate, sagittate at the base; ovary 2-celled
or incompletely 1-celled, the ovules numerous, the style elongate. Fruits baccoid,
2-celled; seeds numerous, large, embedded in a gelatinous pulp.

Posoqueria is found in Mexico, Central America, the West Indies, and South
America. It is a genus of about 15 species.

Literature:

Steyermark, J. 1967. Posoqueria in Botany of the Guayana Highlands — Part VII
Mem. New York Bot. Gard. 17: 322-333.

1. Posoqueria latifoiia (Rudge) R. & S., Syst. Veg. 5: 227. 1819.— Fig, 70,

Solena latifoiia Rudge, Hist. PI. Guiane 1: 26. 1805. type: French Guiana, not seen.

Tocoyena longifolia H.B.K., Nov. Gen. Sp. PI. 3: 411. 1819. type: Not seen.

Posoqueria decora DC, Prodr. 4: 375. 1830. type: Not seen.

Stannia panamensis Walp. & Duchass., Linnaea23: 755. 1850. type: ''Isthmo panamensi," not seen.
Posoqueria panamensis (Walp. & Duchass.) Walp., Ann. Bot. Syst. 2: 797. 1852.

Stannia metensis Karst., Linnaea 28: 441. 1857. type: Colombia, Karsten, not seen.

Trees to 9 m tall, the branchlets terete, ultimately angular, glabrous, the nodes
3-15 cm apart. Leaves oblong, ovate oblong, rarely rotund, 8-32 cm long, 3.5-
22 cm wide, rounded or widely deltoid at the apex, obtuse, cuneate, acute, rarely
truncate at the base, the costa plane or grooved above, prominent beneath, the
lateral veins 6-8, the intervenal areas delicately reticulate, the margin usually
revolute, subcoriaceous, shiny and glabrous above, glabrous to occasionally fine-
ly puberulent beneath, the petioles lignose, 0.3-2.5 cm long, glabrous, the stipules
triangular, 1.0-1.3 cm long, obtuse, coriaceous, glabrous, somewhat dilated near
attachment. Inflorescences terminal, glabrous, drying black, the elongate flowers
in cymules disposed radially at the end of a short peduncle, the latter 0.5-2,2 cm

334 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

long. Flowers sessile or subsessile, the hypanthium oblong or turbinate, ca. 0.8
cm long, smooth, glabrous, drying black, the calycine cup shorter than the hy-
panthium, thickly carnose, the teeth evanescent or 5, triangular to subrotund, to
0.5 mm long; corolla white, elongate, slender, in bud having a clavate appearance,
the connate lobes oblong or ovate oblong, distinctly flexed, to 1 .5 cm long, obtuse,
the tube 4.0-12.5(-16.0) cm long, 0.2-0.4 cm wide, glabrous outside, puberulent
at the base and at the mouth within, thickly petaloid, the lobes 4, often reflexed
at maturity, narrowly oblong, 1.2-1.7 cm long, puberulent proximally; stamens
4, the anthers exserted, oblong, crescentic, ca. 4 mm long, acute at the apex and
base, the abaxial face puberulent, the filaments attached at the mouth, shorter
than the corolla lobes, the style filamentous, as long as the corolla tube, the
stigmas digitiform or narrowly oblong, ca. 2 mm long. Fruits resembling an or-
ange, globose, 4-5(-6.7) cm in diam., maturing yellow, smooth, glabrous, the
seeds white to greenish, numerous in a greenish pulp.

The range of Posoqueria latifolia is southern Mexico south to Panama; in
South America it occurs in Colombia, Venezuela, in the Guianas, in Amazonian
Brazil, and in Peru. The pulp of the fruit is edible. Steyermark's maintenance of
P, panamensis, in view of my examination of over 80 collections in Panama, is
untenable. The characters used in keying out P. panamensis and P. latifolia
(Steyermark, 1967, pp. 322-323) overlap to a great degree. Croat (1979) notes
that the flowers are pollinated by hawkmoths. ''Boca (de) Vieja''; ''Borajo'';
''Fruta de Mono''; ''Huevo de Mono"; 'Truta de Murcielago''; ''Mariquito de
Marea"; ''Monkey Apple"; "Tuli Viejo"; "Sui Tirmas" {Stier 96),

BOCAS DEL TORo: Finca 4 (Delta), 10 mi NW of Almirante, DArcy 1 1203 (MO). Changuinola
River, Lewis et al. 884 (F, GH, K, MO, NY, PMA, UC). Chiriquicito, Lewis et aL 2048 (MO). Nances
Cay Island, Wedel 579 (MO). Water Valley, Wedel 997 (MO). Chiriqui Lagoon, Wedel 1187 (MO).
Water Valley, Wedel 1614 (MO). Chiriqui Lagoon, Wedel 2379 (MO), canal zone: Chiva-Chiva
Trail, Allen 955 (MO). Fort Sherman, Blum 1499 (MO). Summit Garden, Croat 13134, 16559 (both
MO). Fort Kobbe, Duke 4258 (MO). Fort Sherman, Duke 9245 (MO). Farfan Beach, Duke 11734
(MO). Coco Solo, Dwyer & Duke 7920A (MO). Albrook Air Force Base, Hayden 63 (MO). DeviPs
Beach near Limon Bay, Johnston 1565 (MO). Fort Sherman, Johnston 1684 (MO). Trail along Rio
Petitpie; road to Fort Sherman from Gatiin Locks, Mori & Kallunki 2699 (MO). Howard Air Force
Base, Oliver & MacBryde 1888 (GH, MO). Rio Cocoli, Road K-9, Stern et aL 317 (MO). Albrook
Air Force Base, Stimson 5373 (MO). Chiva-Chiva Trail, Tyson 1337 (MO). Fort Clayton, Tyson 3470
(MO). BARRO COLORADO ISLAND: Aviles 7 A (MO); Croat 6057, 6737, 12633, 15062 (all MO); Shattuck
302, 1067 (both MO); Weaver & Foster 1478 (MO), chiriqui: 12.4 mi N of David, Lewis et al. 714
(MO), cocle: El Valle, Alston & Allen 1861 (MO). La Pintada, Escobar 9 (MO), colon: Santa Rita
Ridge, Correa & Dressier 923 (MO). San Miguel de la Borda, Croat 9882 (MO). Punta Pedro, Icacal,
Howell 38 (MO). Piiia, Lewis et al, 1845 (GH, MO, UC, US). Along Caribbean at Quebrada Santa
Marta, AV2 km SW of Pina, Nee 11695 (MO), darien: Cerro Pirre, Bristan 583 (MO). Quebrado
Venado and Peje Swamp, Bristan 1016 (MO). Pico Piraque, Duke 8142 (MO). Rio Balsa, Duke 8778
(MO). Pifias, Duke 10594 (MO). Santa Fe, Duke 14275 (MO). Canasas and Sabalo, Duke 14474 (MO).
Rio Uruseco, AV2 km S of El Real, Mori & Kallunki 5517 (MO). Santa Fe, Tyson et al. 4812 (MO).
LOS Santos: Loma Prieta, Duke 11839 (MO); Lewis et al. 2220 (COL, MO, UC, VEN). Macaracas
and Quebrada Bejuco, Tyson et aL 2903 (MO). Panama: Cerro Campana, Castillo 21 (MO). El

Llano, Croat 14502 (MO). Cerro Campana, Croat 14680 (MO). Chepo, Dodge 10722 (MO). Quebrada
La Palma, Dodge <5 Allen 17357. 17367 (both MO). Rio Pita, Duke 4763 (MO). Ri'o Teribe, Duke
5673 (MO). Isla del Rey, near San Miguel, Duke 10418 (MO). Cermeno, Dwyer 6631 (MO). Goofy
Lake, Cerro Azul, Dwyer 7129 (MO). Chagres, Fendler 293 (MO). El Llano, Gentry & Tyson 1726
(MO). Lidice, Jaramillo 11 (MO). Chilibre, Lao & Holdridge 7 (MO). Falls of La Chorrera, Lewis
et aL 5207 (MO, SCZ, UC, VEN). Gorgas Memorial Labs, 5-10 km NE of Altos de Pacora, 600 m,
Mori & Kallunki 3338 (MO). El Llano-Carti Road, 12 km N of Panamerican Highway, Nee & Dressier
9382 (MO). El Llano-Carti Road, A^^^ & Dwyer 9236 (MO). Fort San Lorenzo, Oustgaard 56 (MO).
Villa Rosario, Saldana 40 (MO). Saboga Island, Tvson & Loftin 5148 (MO). Isla Taboga, Woodson

1980J DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 335

et al. 1434 (MO), san blas: Puerto Obaldfa, Croat 16949 (MO). Soskatupu, Duke 8960 (MO). Isla
Pifia, near Mulatupo, Elias 1703 (GH, MO, US). Opposite Achituppu, Lewis et al 114 (GH, MO,
UC, US). Opposite Ailigandi, Lewis et al. 192 (GH, K, MO, NY, UC, US), veraguas: Islade Coiba,
Dwyer 1621, 2393 (both MO). Mouth of Rio Concepcion, Lewis et al, 2821 (COL, DUKE, MICH,
MO, UC). NW of Santa Fe, 11 km from Escuela Agricola de Piedra, 450-550 m, Mori & Kallunki
3894 (MO). S of Santa Fe, 450 m. Nee 8058 (MO). 14 km of Santiago, Bridge Santiago-Santa Fe
Highway, Nee 9919 (MO).

67. PSYCHOTRIA

Psychotria L., Syst. Nat., ed. 10. 929, 1122, 1364. 1759. Nomen conserv. type
P. asiatica L.

Psychotrophum P. Br., Civ. Nat. Hist. Jam. 160. 1756. Nomen rejic.
Myristiphyllum P. Br., Civ. Nat. Hist. Jam. 152. 1756. Nomen rejic.
Nonatelia Aubl., Hist. PI. Guiane 182. 1775. type: yV. officinalis Aubl.
Mapouria Aubl., Hist. PI. Guiane I: 175. 1775, type: M. guianensis Aubl.
Ronahea Aubl., Hist. PI. Guiane 1: 154. 1775. type: R. latifolia Aubl.

Siibshrubs, shrubs or trees, rarely scandent, occasionally epiphytic. Leaves
rarely sessile, opposite; stipules free or connate, usually caducous. Inflorescences
usually terminal, often paniculate, occasionally aggregated into heads; bracts
usually persistent, occasionally involucrate; bracteoles usually present, often de-
ciduous. Flowers mostly small, (4-)5(-8)-merous, bisexual, often heterostylous,
the hypanthium small, the calycine cup usually short, occasionally with small
glands within, the lobes or teeth absent or present, usually 5; corolla usually

white or yellow, the tube short, the lobes valvate in the bud, occasionally galeate;
stamens included or exserted, the anthers usually narrowly oblong, the filaments
variously inserted in the corolla tube; ovarian disc usually cushion shaped, the
style solitary, slender, the stigmas 2(4-5), the ovary 2(4-5)-celled, the septum
thick, the ovules 1 per locule, attached basally. Fruits fleshy, the flesh usually
soon deciduous exposing 2 pyrenes, the endocarp of the pyrene ventrally flat or
grooved, dorsally costate or ecostate, the testa thin, the albumen hard, sometimes
ruminate.

Psychotria is well distributed throughout the warmer parts of the world. It is
the largest genus of the Rubiaceae with about 800 species. It is the largest genus

of the family in Panama.

The Panamanian species with few exceptions sort into 2 genera without dif-
ficulty. Although from a worldwide viewpoint there is much to support the in-
corporation of Cephaelis into Psychotria as was done by Steyermark (1974), I

maintain them as separate genera.

The frequent absence of stipules on individual herbarium sheets represents a
major difficulty in preparing and using a key. Stipules are of great diagnostic
importance in the Rubiaceae as a whole, and are morphologically of greater im-
portance than the style, stigmas, or anther. This is perhaps the reason why Paul
Standley, the doyen of students of the Rubiaceae of the New World until his
death in 1967, usually gave little attention to the internal parts of the flowers in
his descriptions. The differences between many species of Psychotria are difficult
to put into words. Standley (1940) in describing P, allenii summed up the problem:
''Like the majority of Psychotria species, this one has no outstanding distinctive
characters. It is, however, unlike any other known from Central America.""

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1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 343

1, Psychotria acuminata Benth., Bot. Voy. Sulphur 107. 1845. type: Isle of Gor-
gona and Colombian Coast. Not seen.

la. Psychotria acuminata ssp. acuminata.

Shrubs to 3 m tall, the branchlets terete, smooth, glabrate, rimose, ultimately
angular; nodes well spaced; pith white. Leaves widely oblong or lanceolate, 6-
18 cm long, 2.5-8.5 cm wide, deltoid to acute apically, long acuminate, the acu-
men to 2 cm long, acute, often falcate, basally obtuse to cuneate, occasionally
inequilateral; the costa prominulous above, prominulous to prominent beneath,
the lateral veins 11-12, strict then arcuate, usually with 1-2 smaller and soon
evanescent veins between 2 lateral veins, the tertiary veins irregular pinnatiform,
the intervenal areas spreading reticulate, chartaceous, usually concolorous,
drying lustrous green above, glabrous or occasionally villosulose on the costa
and veins beneath; petioles 1.2-2.0 cm long, glabrous; stipules connate, cupuli-
form, usually ca. 3 mm long, often ciliate, truncate and with 2 erect awns, scarcely
longer or almost twice as long as the body, puberulent. Inflorescences shorter
than terminal leaves, terminal or pseudo terminal, paniculate, often subhemi-
spherical or oblong, 1.0-6.5 cm long, 0.8-2.0 cm wide, puberulent; peduncles to
3.5 cm long, 0.7-LO mm wide, the lowermost branches opposite or alternate, the
upper branches few, opposite or alternate; bracts minute, the bracteoles absent.
Flowers 4-5-merous, the hypanthium short, cupuliform, to 0.5 mm long, some-
what truncate or with minute irregular teeth, puberulent; corolla white, green or
yellow, the tube cylindrical, to 6 mm long, glabrous outside with a few hairs
below the filament attachment within, the lobes 4-5, oblong or ovate, to 2.5 mm
long, glabrous or minutely puberulent outside; stamens 4-5, included in long-
styled flowers, the anthers narrowly oblong, ca. 1 mm long, the filaments attached
near the middle of the tube; styles long or short, the long styles to 7 mm long,
exserted 2-3 mm; ovarian disc bilobed, to 0.5 mm deep, the stigmas 1.0-1.8 mm
long. Fruits usually didymous, the pyrenes 2, rarely 5, 4 in epiphytic species,
rounded, 5-7 mm in diam., glabrous, the ventral face of the endocarp with a

groove, the dorsal surface ribless, yellow, turning gray blue at maturity.

This well-collected species ranges from Mexico south to Peru. The Panama-
nian material has long been misidentified as Psychotria cuspidata Bredemeyer ex
Romer & Schultes, a species confined to Venezuela (cf. Steyermark, 1972). Croat
15077 notes that the stamens may extend 3 mm above the throat and Croat 14981
notes that the stamens are in a row on one side of the corolla.

CANAL zone: Without other locality, Blum 569 (MO). Albrook, Dnyer 6708 (MO, US), 6747
(MO). Las Cruces Trail, Dnyer & Hayden 5 (MO). Pipeline Road, Gentry 1947 (MO). Pavon Road,
NW part of Canal Zone, Johnston 1627 (MO). Pipeline Road, N of Gamboa, Luteyn & Foster 1526
(MO, US). Howard Air Force Base, Oliver & Machryde 1895 (GH, MO, UC, US). Navy Pipeline
Reserve Area, ca. 3 mi NW of Gamboa, Stimson 5342 (MO, NY). 2 mi N of Cocoli, Tyson 1638
(MO). Fort Sherman, Tyson 1709 (MO); Tyson & Blum 1668 (MO). Fort Clayton, Tyson 1802 (US).
BARRO COLORADO ISLAND! AvUes 46 (MO); Croat 4092, 4525, 4607, 4891, 5195A, 5877, 6112, 6181,
6217, 6549. 6773, 7301, 8020, 9542, 11179, 11788, 14981, 15077 (all MO); Ebinger 4 {U0)\ Hayden 5,
24, 53, 134, 148, 149 (all MO); Kennedy 1846 (MO); Killip 39958 (MO); Knight 69-25 (MO); V^elsh
19831 (MO); Woodworth & Vestal 423 (MO), colon: Rfo Piedras, on road to Portobelo, Blum et aL
2520 (MO). Achiote, Tyson et al. 4512 (MO), darien: La Boca de Pirre, Bristan 1270 (MO). Rfo
Tucuti, Duke 5256 (MO). Between Rfo Balsa and Rfo Aretf, peak 300 ft high, Duke 8740 (MO). Cativo
Swamp near Tusijuanda, up Rfo Balsa, Duke 13526 (MO). Piria-Canasas Trail near Piria, 100 m, Duke

344 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

14316 (MO). Cuasi-Cana Trail on Cerro Campamiento, E of Tres Bocas, Klrkhride & Duke 1225
(MO). Rio Tuquesa at mining camp, ''Charco Chiva/' Mori 6946 (MO). Campamento Buena Vista,
Rio Chucunaque, above confluence with Rfo Tuquesa, Stern et al. 862 (MO). Cana-Cuasi Trail (Camp
2), Chepigana, Terry & Terry 1488 (MO). Periaque Camp at River, Tyson et al. 4745 (MO), herrrra:
12.5 mi S of Ocu, 1200 ft, Lewis et al. 1629 (GH, MO, US). Los santos: Loma Prieta, Cerro Grande,
2400-2800 ft, Lewis et aL 2192 (MO, UC, VEN). Panama: Nuevo Emperador, Baker 16 (MO). Cerro
Jefe, Croat 14440 (MO). 6 mi E of Chepo, Panamerican Highway, Duke 4087 (MO). Rio Pita, 1-3 mi
above Rfo Maestra, Duke 4745, 4768 (both MO). Near confluence of Rio Pacora and Rio Corso, ca.
450 m, Duke 11964 (MO). Fort San Lorenzo, Hayden 109 (MO). Cerro Campana, Kirkhride &
Hayden 309 (MO). Altos del Rio Pacora, Lewis et al. 2344 (MO, UC). 5-10 km NE of Altos de
Pacora, Mori & Kallunki 6055 (MO). El Llano-Carti Road, 11-12 km from Interamerican Highway,
300^00 m, Mori et al. 6893 (MO). Rios Pacora and Corso, Oliver 2376 (MO, UC, VEN). Cerro Azul
to Cerro Jefe, ca. 2000 ft, Tyson et al. 4317 (MO). Near Arraijan, Woodson et at. 773. 1346 (both
MO). VERAGUAs: Isla de Coiba, Dwyer 2381 (MO). 5 mi NW of Santa Fe, on Caribbean slope above
Rfo Primero Brazo, 700-1200 m, Liesner 830 (MO). 5 mi W of Santa Fe, road past Escuela Agricola
Alto Piedra, on Pacific side, 800-1200 m, Liesner 882 (MO).

lb. Psychotria acuminata ssp. boraginoides Dwyer.*^^ type: Panama, Liesner 1 187
(MO).

Psychotria acuminata subsp. boraginoides is known only from Panama and
Colombia (Choco). It is easily recognized by the reflexed branches of the inflo-
rescences which are curved and have the flowers secundly disposed.

Panama: El Llano-Carti Road, 19 mi N of El Llano, 500 m, Busey 897 (MO). 2.4 mi beyond
Cerro Jefe toward Altos de Pacora, 800-1000 m, Croat 22660 (MO). Campo Tres, 3 mi NE of Alto
de Pacora, Croat 22692 (MO). 20 km above Panamerican Highway. El Llano to Cartf, 200-400 m,
Kennedy 2567 (MO). Campo Tres, 3 mi NE of Altos de Pacora, Liesner 526 (MO). 12 mi above
Panamerican Highway, Liesner 1187 (MO). 5 mi NE of Altos de Pacora, Mori & Kallunki 6055 (MO).

2. Psychotria aggregata Standley, Contr. U.S. Natl. Herb. 18: 128. 1916. type:
Panama, Pittier 3264 (US, holotype).

Fleshy shrubs to 1 m tall, the stem round, the branchlets angular, puberulent,
longitudinally striate. Leaves oblong or oblanceolate, 12-20 cm long, 3-5 cm
wide, deltoid at the apex, often vaguely contracted, the acumen to 0.5 cm long,
acute to attenuate acute at the base, often oblique, the costa prominulous above,
prominent beneath, to 0.18 cm wide proximally, the lateral veins 10-16, arcuate,
usually with one subparallel vein between a pair of lateral veins, thinly coriaceous
to papyraceous, puberulent, often farinose (deciduous hair bases?); petioles to 3
cm long, longitudinally striate; stipules connate, apparently persisting only at the
apex of the stem or branchlets, presumably oblong rotund, to 1 cm long or longer
with 2 divergent triangular lobes shorter than the body of the stipule and the base
of the body with a shallow auricle on each side, crassate, glabrous. Inflorescences
axillary, paniculate, subsessile or on peduncles to 3 cm long, the flowers aggre-
gated into few-flowered heads, these borne on short branches or subsessile; bracts
at the base of the branches subulate, shorter or longer than the branches; floral
bracts and bracteoles narrowly ovate or oblong. Flowers sessile or subsessile;
hypanthium ca. 0.7 mm long, pubescent, the calycine cup 1.2 mm long, petaloid,

**'* Psychotria acuminata ssp. boraginoides Dwyer spec. nov. Differt a P. acuminata ssp. acumi-
nata ramis inflorescentiae primo simplicibus, 5-10 mm longis, tunc dichotome ramificantibus cymulis
sessilibus 1-3-floriferis floribus in eminentiis minutis secundis 1-3 mm distantibus dispositis.

I980J DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 345

pubescent, eglandular within, the raphides numerous, the teeth 5, somewhat un-
equal, subulate or narrowly oblong, 0.6-1.0 mm long; corolla white, the tube
cylindrical, to 2.5 mm long, petaloid, pubescent outside, ?glabrous within, the
lobes 5, in bud spreading, galeate, the anthers fitting in snugly, the hood solid,
at anthesis the lobes elongating, the galeate area pulling away from the anthers
with much of the lobe plane; stamens 5, the anthers oblong, ca. 1 mm long,
obtuse, the filaments short; ovarian disc ca. 0.6 mm long, the style slender, ca.
2 mm long, the stigmas 2, slightly wider than the style. Fruits (immature?) sessile,
oblong, ca. 5 mm long, 3.5-4.5 mm wide, white, glabrous, the raphides numerous,
the persistent calyx to 2 mm long, the teeth evident.

Psychotria aggregata is known only from Panama. The stipules are quite
variable in shape. The galeate tips of the corolla lobes are diagnostic. Dwyer &
Hayden 722 notes that the leaves are "velvetv ereen."

BOCAS DEL TORO: Chiriqui Trail near Chiriqui border, Kirkhride & Duke 995 (GH, MO, US).
CHiR[Quf: Boquete, Cerro Horqueta, 5000 ft, Dwyer & Hayden 722 (MO). Chiriqui Trail between
Pinola and David, Kirkhride & Duke 859 (MO). Cerro Pando, Panama-Costa Rica border, 16 km NW
of El Hato de Volcan, 2000-2480 m, Mori & BoUen 7274 (MO). N of San Felix at Chiriqui'-Bocas del
Toro border, road to Cerro Colorado copper mine. 5000-5500 ft, Mori & Kallunki 5387, 5941, 5953
(all MO). Alto de Las Palmas, Cerro Horqueta, Pittier 3264 (US). Bajo Chorro, Skog & D'Arcy 4092
(US), cocle: Cerro Pilon, Croat 14352 (MO); Dwyer & Lallathin 8957 (MO). Cerro Gaital Caracoral,
Dnyer & Correa 8977 (MO). Panama: Road to Altos de Pacora. vicinity of schoolhouse, Mori &
Kallunki 2366 (MO).

3. Psychotria alboviridula Krause, Notizbl. Konigl. Bot. Gard. Berlin 6: 208.
1914. type: Brazil, Ule 9846. Not seen.

Shrubs or trees to 7 m tall, the branchlets terete, smooth, glabrate, drying red
or black, the nodes well spaced. Leaves oblong or lanceolate, 7.5-25.0 cm long,
2.5-8.5 cm wide, often distinctly inequilateral, tapering obtusely or deltoidly at
the apex, acuminate, the acumen to 1.8 cm long, often falcate, cuneate or atten-
uate acute, often slightly inequilateral at the base, the costa prominulous above,
often grooved, prominent beneath, to 3 mm wide proximally, often the lower Vi
of the blade with solitary domatia in the vein axils, these to 6 mm long, the pore
1-3 mm wide, the lateral veins 7-12, arcuate, often subrigid, subcoriaceous to
papyraceous, often drying red brown, especially younger leaves, glabrous; peti-
oles to 2 cm long, 1.5-2.2 mm wide; stipules caducous, triangular or ovate, 4-8
mm long, acute, glabrous, except setose within. Inflorescences terminal, solitary,
at the end of short branches or axillary, paniculate, glabrous, 15-20 cm long, the
peduncle 5.0-7.5 cm long, flattened, the branches few, the flowers few, borne in
spikelike fashion, spreading or strict, well spaced, often deflexed, opposite and
monostichous above, the lowermost 3-4 often verticillate, tumid above junction
with rachis; bracts ovate or triangular, to 2.5 mm long, acute. Flowers drying
red; hypanthium short, 0.8 mm long, the calycine cup 0.3-0.5 mm long, turgid,
eglandulose within or with minute red glands, glabrous except minutely ciliolate
on the teeth, the teeth 5, widely triangular, to 0.3 mm long; corolla often shorter
than the hypanthium and the calyx, to 2.5 mm long, the tube short, the lobes 5,
oblong, ca. 0.8 mm long, glabrous; stamens 5, the anthers subrotund, ca. 0.8 mm
long, the filaments short; stigmas and style ca. 0.5 mm long, the stigmas perhaps

346 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

longer and wider than the style. Fruits sessile or subsessile, oblong, ca. 5 mm
long, usually obtuse at the apex and the base, glabrous, drying red black, sulcate,
the persistent calyx scarcely measurable.

Psychotria alhoviridula is known from Costa Rica, Panama, Colombia, and
Peru. The spikelike and few-flowered branches of the spreading inflorescence
coupled with an overall deep coloration when dry make it one of the easiest of
the Panamanian Psychotria to recognize. Nee's collection 10937 is the only one
having fruit. Occasional fruits of this collection are teratological, measuring to
1.5 cm long and appearing like a small fruit of Ficus. These swollen fruits may
be stimulated by insects. P. alhoviridula has the largest domatia encountered
among the Panamanian species. Steyermark (1972) reduced P. alhoviridula to the
synonymy of P. remota Bentham.

darien: Peak between Rio Balsa and Rio Areli, ca. 300 ft, Duke 8745 (MO). Puerto St. Dorothea,
D\vyer 3218 (MO). Panama: Between Cerro Jefe and La Eneida, 2100-2900 ft, Dwyer 8210 (MO).
Banks of Rfo Ipeti, Folsom & Collins 1664 (MO). Stream flowing out of Serrania de Maje, Folsom
& Collins 1715 (MO). Road from El Llano to Carti-Tupile, continental divide to 1 mi from divide,
300-500 m, Liesner 1280 (MO). El Llano-Carti Road, 16-I8V'2 km by road N of Panamerican Highway,
400-450 m, Nee & Tyson 19037 (MO). Cerro Azul to Cerro Jefe, Tyson et al. 4237 (MO). 16-18 km
from Panamerican Highway on El Llano-Carti Road, Tyson & Nee 7318 (MO), san blas: Trail from
E of Cangandi-Mandinga airport road, 2-5 mi E of Mandinga Airport, Duke 14774 (MO). Colombia:
CHOCo: Between Rio Curiche and Alto Curiche, Duke 9581 (MO). Madurex Logging Camp, Duke
9933 (MO). Between Rio Nercua and Quebrada Barrial and Quebrada Ambrosio near Camp Curiche,
10-294 m, Duke 11469 (MO). Rio Salaqui, Hydro Camp, 1 100 ft, Duke 15777 (MO). N of Alto Curiche,
ca. 300 m, Duke & Idroho 11223 (MO).

4. Psychotria allenii Standley, Ann. Missouri Bot. Card. 27: 342. 1940. type:
Panama, Allen 1796 (F, holotype; NY, isotype).

Trees 3-10 m tall, the branchlets slender, smooth, ultimately angular, the
nodes well spaced. Leaves lance oblong to oblong, occasionally oblong rotund
or obovate rotund, 5.5-16.0 cm long, 2.3-6.0 cm wide, acute, obtuse or occa-
sionally rounded, the acumen 1.0-1.5 cm long, often falcate, widely cuneate and
often inequilateral at the base, the costa slender, immersed, plane or prominulous
above, prominulous to subprominent beneath, the lateral veins 6-8(-9), arcuate,
prominulous beneath, often dull red, the intervenal areas smooth, the reticulations
occasionally evanescent, chartaceous, concolorous, glabrate above and beneath;
petioles 1.0-2.5(-3.0) cm long; stipules deciduous, perhaps connate, each part of
the sheath subrotund, 1-5 mm long with 2 thickly subulate or triangular lobes,
usually shorter than the sheath. Inflorescences terminal, solitary, cymose panic-
ulate, exceeding the uppermost leaves, 10-17 cm long, 5-10 cm wide; peduncle
4.5-8.5 cm long, to 0.4 cm wide, stiff, glabrate to puberulent, the branches few;
peduncle either terminating in 4-5 radiately disposed branches or the lowermost
branches 2-4 from which a rachis extends bearing 2 well-spaced sets of 4-5
verticillately disposed branches, the lowermost branches to 6.5 cm long, these
terminated by several few-flowered cymes. Flowers small (in bud), sessile, drying
red, the hypanthium ca. 0.5 mm long, glabrous, the calycine cup saucer shaped,
shorter than the hypanthium, the teeth small or obsolete, triangular, to 0.3 mm
long; corolla white, to 4 mm long, the lobes as long as the tube, glabrous outside,
the tube densely villose within; stamens 5, the anthers oblong, ca. 1.5 mm long,

1980J DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 347

the filaments to 0.8 mm long. Fruits large, to \.5 cm in length, carnose, blue,
rotund, the fleshy pericarp persisting, the pyrenes hemispherical, smooth except
minutely rugulose microscopically, not costate, blue {Mori & Kallunki 3174).

Psychotria allenii is known only from Panama. Standley stated: "Like the
majority o{ Psychotria species, this one has no outstanding distinctive characters.
It is, however, unlike any other known from Central America." Unfortunately
Standley did not see fruits, which tend to retain the fleshy portion of the pericarp,
unusual in Psychotria. The fruits are large, exceeded among Panamanian Psy-
chotria only by P. carnosocarpa, P. chiapensis, P. calophylla, and P. olgae. The
relatively smooth blue pyrenes are diagnostic.

cocle: EI Valle, Allen 1796 (MO, NY). Cerro Pilon to El Valle, ca. 2500 ft, Duke & Dwyer 13903
(MO). Cerro Pilon, ca. 2900 ft, Dwyer & Lallathin 8629 (MO). La Mesa above El Valle, Gentry 7425
(MO). Panama: Cerro Campana, Croat 14685 (MO). Cerro Campana, Kirkhride & Hayden 113 (MO).
El Llano-Carti Road 17.5 km from Interamerican Highway, 350 m, Mori & Kallunki 4617 (MO). 5-
10 km NE of Altos de Pacora, Mori & Kallunki 6050 (MO), veraguas: 3.9-5.0 mi S of Santa Fe,
Gentry 2972 (MO). 7 mi W of Santa Fe, Nee 11155 (MO).

5. Psychotria bakeri Dwyer. ^^ type: Panama, Lewis et al. 2249 (MO, holotype).

Shrubs to 1.5 m tall, much branched, the branchlets slender, often flexuous,
smooth, glabrous except densely ciliolate within the stipules, ultimately dark red,
the nodes turgid, well spaced. Leaves elliptic salicoid or elliptic rhomboid, 1.5-
6.5(-9.0) cm long, 0.6-1.5(-3.0) cm wide, deltoid toward the apex, acuminate or
not, the acumen vague, attenuate acute toward the base, the costa prominulous
above, sometimes sulcate, scarcely prominent beneath, to 0.3 mm wide, the lat-
eral veins 8-10, moderately conspicuous beneath, inconspicuous above, the mar-
gin slightly callose, somewhat wrinkled, stiffly chartaceous, glabrous, smooth,
concolorous, dark red; petioles absent or to 3 mm long; stipules free, often per-
sistent terminally, glabrous, red when dry, somewhat triangular oblong, rhomboid
rotund or somewhat rectangular, truncate at the apex, with 2 subulate horns,
often unequal in length, sometimes longer than the body. Inflorescences terminal,
solitary, glabrous except for the bracts and bracteoles, 2-4 cm long, often um-
belloid in outline, the peduncle to 2.5 cm long, ca. 0.7 cm wide, often with 3
radiate branches terminating the peduncle, to 1 cm long, the flowers of cymules
few, on short branches or on a short rachis extending to 0.7 mm long; bracts and
bracteoles minute, ovate, to 1 mm long, acute, concave, ciliolate marginally.
Flowers sessile, the hypanthium turbinate, 0.8 mm long, glabrous, the calycine
cup short, the teeth 5, oblong, 0.5 mm long, obtuse, carnose, glabrous except
minutely ciliolate toward the apex; corolla white, the tube cylindrical, to 2.5 mm
long, 1.2 mm wide, petaloid, glabrous outside, densely villose within above the

^^ Psychotria hakeri Dwyer, spec. nov. Frutices ad 1.5 m alti. Folia elliptico-salicoidea vel
elliptico-rhomboidea, 1.5-6.5 cm longa, 0.6-1.5 cm lata, venis lateralibus 8-10, venis minoribus in-
conspicuis, margine leviter calloso, rigido-chartacea, glabra laevia opaco-rubra; petiolis nullis vel ad
3 mm longis; stipulis liberis, ad 4.5 mm longis, apice truncatis cornubus 2 subulatis. Inflorescentiae
terminales solitariae 2-4 cm longae glabrae. Flores sessiles dentibus calycis 0.5 mm longis, glabris
praeter minuto-ciliolatis apicem versus; corolla tubo ad 2.5 mm longo. Fructus oblongi, ca. 5 mm
longi, glabri costati in sicco rubri.

348 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

middle of the tube, the lobes 5(?), ovate oblong, to 1.7 mm long, obtuse; stamens
5, the filaments slender, to 1.5 mm long, attached near the middle of the tube;
style 2 mm long, slender at base, dilated toward the apex, sparsely ciliolate; the
stigmas 2, 0.1 mm long. Fruits sessile, oblong, ca. 5 mm long, glabrous, obviously
ribbed, drying red.

Psychotria hakeri is named in honor of Dr. Kenneth Baker, long a student of
the flora of Panama. It is known only from the type collection. It is closely related
to P, chiriqiiina from Cerro Horqueta, Chiriqui, but it differs in its shorter petioles
and leaves with fewer lateral veins. The stipules, unlike those of P. chiriqidna,
may persist at the apices of the twiglets and are 2-lobed.

CHiRiQUi: Cerro Punta, IVi km SB of town, Mori & Kallunki 5660 (MO), los santos: Loma
Prieta, Cerro Grande, 2400-2800 ft, Lenis et al. 2249 (MO. holotype). Panama: Cerro Azul, 2000 ft,
Tyson & Blum 4066 (SCZ).

6. Psychotria berteriana DC, Prodr. 4: 515. 1830. type: Puerto Rico and Santo
Domingo, Bertero, Not seen,

p. angustiflora Krause, Bot. Jahrb. Syst. 54. Beibl. 119: 43. 1916. type: Costa Rica, Tonduz 12996,

Not seen.
P. crehrinervia Standley, Publ. Field Columbian Mus., Bot. Ser. 4: 343. 1929. type: Honduras,

Standley 55286 (F, holotype).

Shrubs or trees, to 2.5 m tall, the branchlets 4-angled, minutely puberulent,
often red when dry, the nodes well spaced, the subnodal area often constricted
on drying. Leaves elliptic or obovate elliptic, 7.5-21.0 cm long, 1.5-11.5 cm wide,
deltoid or obtuse at the apex, acute or attenuate acute at the base, the costa
prominulous above, prominulous beneath, puberulent, the lateral veins 13-20,
arcuate, scarcely prominulous but conspicuous beneath, chartaceous, often stiffly
so, glabrous above, glabrous beneath except on the veins; petioles slender, 1.0-
5.5 cm long; stipules often deciduous, free or shortly connate, to 1 cm long, often
cylindrical, each part slightly retuse at the apex, the lobes of each part conspic-
uously rounded, often spreading with age, scarious marginate, subcoriaceous,
glabrous. Inflorescences terminal, pyramidal paniculate, to 14 cm long, 5-10 cm
wide; peduncle to 7 cm long, relatively stiff, to 0.3 cm wide, puberulent, often
drying dark red, the lowermost primary branches opposite, to 5 cm long, the
superior primary branches usually well spaced, opposite or alternate and bearing
several well-spread pairs of opposite or occasionally alternate secondary branches
or sometimes 3 secondary branches terminating a primary branch, the cymules
or occasionally individual flowers sessile and disposed along the branches to give
a zig-zag effect; bracts and bracteoles persistent, linear or narrowly obovate
oblong, the bracts of the lowermost primary branches 0.4-1 .0 cm long, paralleling
the branches or somewhat reflexed, ciliate. Flowers sessile, the calyx and hy-
panthium 2 mm long, the calycine lobes oblong or triangular, 0.2-1.0 mm long,
obtuse or acute, spreading, puberulent; corolla yellow or greenish white, the tube
cylindrical or subcampanulate, to 6 mm long, scattered puberulent or minutely
puberulent all over outside, puberulent within, petaloid, the lobes oblong, ca. V4
the length of the tube; anthers oblong, ca. 3 mm long, the filaments ca. 1 mm

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 349

long, affixed below the middle of the tube; ovarian disc compressed rotund, to
0.2 mm long, wider than long, the style short, the stigmatic lobes ca. 2 mm long.
Fruits sessile, ovate oblong or oblong rotund, 3-4 mm long, rounded at the apex,
delicately costate, glabrous, the raphides abundant.

Mexico

West

variable species which seems closely related to P. luxurians Rusby.

BOCAS DEL TORo: Cloud foFcst above Quebrada Huron on Cerro Bonyic, 500-1200 ft, Kirkhride
& Duke 614 (MO), darien: Ascent Cerro Pirre, 100-1600 m, Duke 5295 (MO), veraguas: 5 mi W
of Santa Fe on road past Escuela Agricola Alto Piedra on Pacific side of divide, 800-1200 m, Liesner
896 (MO). Lower slopes of Cerro Tute. 750-1000 m, Mori 6244 (MO, US). N of Santa Fe, Escuela
Agricola Alto Piedra, Mori & Kallunki 2534 (MO).

7. Psychotria boquetensis Dwyer.^" type: Panama, Davidson 709 (MO, holotype).

Shrubs to 1.5 m tall, the branchlets smooth, red when dry, the hairs few, the
nodes well spaced, the lenticels dense but inconspicuous. Leaves narrowly ellip-
tic or narrowly obovate elliptic, 9.5-13.5 cm long, 2.0-3.5 cm wide, narrowly
acute toward the apex, acuminate, the acumen ca. 1 cm long, acute at the base,
the costa prominulous or plane above, prominent beneath, the lateral veins ca.
17, arcuate, strongly ascending, papyraceous, drying concolorous, red, the hairs
few, elongate or short, disposed on the costa and veins, often aggregated in the
axils; petioles 0.3-0.5 cm long, slightly alate on the adaxial side; stipules oblong,
to 1.5 cm long, to 0.5 cm wide, obtuse or subacute at the apex, papyraceous,
drying red, the hairs few, red. Inflorescences terminal, erupting from the envel-
oping terminal stipule, to 2 cm long, to 1 cm wide, densely ferrugineous puber-
ulent. Flowers to 3 mm long; hypanthium short, the calycine cup spathaceous,
deeply fissured, the fissure solitary, the teeth 5, in groups of 2 and 3, the cup
petaloid, carnose, the hairs outside few, spreading, eglandular within, the teeth
unequal, subulate, 1-2 mm long; corolla with the tube short, 3.5 mm long, ca. 1
mm wide, villose within near the filament attachment position, the lobes oblong,
ca. 1.5 mm long; stamens 5, the anthers oblong, 0.7 mm long, the filaments 0.7
mm long, attached near the mouth, villose at the base; style ca. 1 mm long, thick,
the stigmas ca. 0.35 mm long; ovarian disc pulviniform, ca. 0.5 mm long. Fruits
not seen.

Psychotria boquetensis is known only from Panama. It is the only Psychotria
I have encountered with the calyx spathaceous.

CHiRiyui': Boquete, 3800 ft, Davidson 709 (MO).

^'^ Psychotria boquetensis Dwyer, spec. nov. Frutices ad 1.5 m aiti, ramulis laevibus in sicco
rubris ciliis paucis ornatis. Folia angusto-elliptica vel anguste obovato-elliptica, 9.5-13.5 cm longa,
2.0-3.5 cm lata, ad apicem attenuato-acuta, venis lateralibus ca. 17. papyracea. ciliis paucis solum
in venis costaque; petiolis 0.3-0.5 cm longis; stipulis oblongis, ad 1.5 cm longis, apice obtusis vel
subacutis ciliis paucis. Inflorescentiae terminales ex stipulis terminalibus eruptae, ad 2 cm longae.
Flores cupula calycis spathacea fissuraalta solitaria notata dentibus 5 sic binis ternique dispositis, I-
2 mm longis; corolla tubo ca. 3.5 mm longo; antheris ca. 0.7 mm longis. Fructus non visi.

350 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

8. Psychotria brachybotrya Muell.-Arg. in Mart., Fl. Bras, 6(5): 327, 1881. type:
Brazil, Spruce 2190. Not seen.

P. iquitosensis Standi., Field Mus. Nat. Hist., Bot. Ser, 8: 195. 1930. type: Peru, Llewelyn Williams
1391 (F, holotype).

Shrubs to 2.5 m tall, the stems smooth, glabrous, often drying yellow, the
nodes well spaced, the pith sometimes porate. Leaves elliptic to ovate elliptic,
7_15(_22) cm long, 2.5-9.0(-13.0) cm wide, usually deltoid or attenuate acute,
the acumen often vague, to 1.5 cm long, often falcate, basally acute, attenuate
acute to cuneate, the costa prominulous above and beneath, the lateral veins ca.
9(-12?), strongly ascending, arcuate, uniting near the margin to form an undulate
vein, thinly papyraceous, usually drying green or tan, concolorous, glabrous
above and beneath or with well spaced, small appressed white hairs; stipules
persistent, connate, the sheath short, turgid, 0.1-0.3 mm long, each part with 2
awns to twice the length of the body, glabrous. Inflorescences terminal, solitary,
usually 1.0-1.5 cm long at early anthesis, in fruit to 3(4?) cm long, the peduncle
absent or to 1.2 cm long, glabrous to puberulent, the branches either reduced and
scarcely measurable or to 1 cm long, disposed in 2-3 opposite pairs perpendicular
or deflexed to the rachis; bracts arising from the proximal portion of the branches,
foliose, oblong lanceolate or narrowly oblong, 0.6-0.8 cm long, to 0.2 cm wide,
glabrate, narrowly deltoid at the apex, ultimately obtuse; bracteoles surrounding
the few-flowered clusters scarcely smaller than the bracts, ascending, involucrate,
hiding the small flowers. Flowers sessile, the hypanthium short, glandular shiny,
glabrate to pubescent, the margin erosulose or with 5 triangular teeth, to 0.3 mm
long, ciliolate, usually with a minute gland at the sinuses; corolla white, the tube
ca. 2 mm long, petaloid, puberulent outside, villose within, the lobes 5, oblong,
to 1.5 mm long, obtuse; anthers narrowly oblong, ca, 1 mm long, obtuse, the
filaments ca. 0.5 mm long, affixed above the middle; style ca. 4 mm long, the
stigmas 0.2 mm long; ovarian disc compressed rotund, 0.5 mm long. Fruits ses-
sile, rotund, to 3 mm in diam., glabrous with abundant raphides, conspicuously
costate, the remnants of the calyx scarcely visible.

Psychotria brachybotrya is known from Panama, Trinidad, northern South

America to Bolivia.

Martius collection fr

Missouri (MO)

BOCAS DEL TORo: W of Almirantc, Blum 1358 (MO). Water Valley, Chiriqui Lagoon, Wedel 1638
(MO). CANAi zone: Juan Mina, Bartlett & Lasser 16559 (MO). Rio Piedras, road to Portobelo, Blum
et al. 2521 (MO). Pipeline Road. 14 mi NW of Gamboa, Croat 12365 (MO). 2 mi NW of Summit
Garden, Croat 16588 (MO). Toro Point, Fort Sherman, Tyson 4420 (MO). Coco Solo, U.S. Army
Tropic Test Center, Dwyer & Duke 7873 (GH, MO, UC, US). Navy Pipeline Reserve Area, NW of
Gamboa, Stimson 5342 (MO). 6 mi N of Gamboa, S of Rio Frijol, Tyson 1506 (MO), barro Colorado
island: Croat 11132 (MO); Foster 1783 (MO), cocle: El Valle, cloud forest, 800-1000 m, Duke
13210 (MO), colon: Maria Chiquita, E of Rio Piedras, Dnyer 7805 (GH, K, MO, NY, UC, US).
Achiote, Tyson et al, 4513 (MO), darien: Cativo Swamp, near Tusijuanda, Duke 13523 (MO). Trail
between Cana and Boca de Cupe, Stern et al. 612 (MO). Cana, 1750 ft, Stern et al. 659 (MO).
herrera: Without other locality, Ehinger 174 (MO). Panama: Cerro Azul, Dwyer 2752, 3109, 4051 A
(all MO). 1.5 mi S of Goofy Lake, Dnyer & Hayden 8022 (GH, MO, US). Fort San Lorenzo, Hayden
110 (MO). Cerro Campana. 2800 ft, MeDaniel 6910 (MO). Isla Taboga, Woodson et al. 1457 (MO).

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhioceae) 351

9. Psychotria brachiata Swartz, Prodr. 45. 1788. type: not seen.

P, caerulea Ruiz & Pavon, PI. Peruv. Chil. 2: 62. 1799. type: Peru, Rihz & Pavon (MO, photo of

isotype? in Delessert Herb.).
Palicourea caerulea (Ruiz & Pavon) R. & S., Syst. Veg. 5: 194. 1819.
Cephaelis polycephala Schiecht., Linnaea 28: 532. 1856. type: Peru, Lechler 2364, Not seen.

Trees to 5 m tall, the branchlets smooth, glabrous, the nodes well spaced.
Leaves oblong or lanceolate, occasionally obovate elliptic, 7-16 cm long, 3-7 cm
wide, obtuse or widely cuneate at the apex, the acumen to 1 cm, basally acute
or rarely obtuse, often slightly inequilateral, the costa prominulous above, prom-
inent beneath, usually laterally villose, the lateral veins 9-12, arcuate or occa-
sionally subperpendicular, the intervenal areas spreading reticulate, papyraceous,
concolorous; petioles to 2.5(-3.5) cm long, glabrous, slender, 0.15 cm wide; sti-
pules free, oblong or oblong rotund, 2-4 mm long, often bilobed, the lobes ca.
1 mm long, obtuse or rounded, 3-4 mm wide basally, appressed, scarious, gla-
brous or occasionally pubescent, appressed setose submarginally within. Inflo-
rescences terminal, solitary, 5-22 cm long, 3.5-7.0 cm wide, thrysoid or racemi-
form paniculate, usually rigidly upright, glabrous to puberulent, rarely
epedunculate and then with branches in pairs or in 3's arising from the stem tip,
usually with peduncles to 5.5 cm long, 0.8-1.0 mm wide, angular, usually with
2-4 pairs of well spaced, opposite branches l-3(-4) cm apart, to 9 cm long, the
uppermost branches occasionally about as long as the lowermost pair, usually
perpendicular, occasionally deflexed, the flowers aggregated into compressed ro-
tund heads terminating the branches, these wider than long, to 1 cm wide; bracts
at the base of the branches involute, oblong, to 8 mm long; floral bracts usually
carinate, the outer bracts larger, to 6 mm long, to 5 mm wide, the inner bracts
ca. 4 mm long, narrower. Flowers with the hypanthium short, the calycine cup
a membranous, inflated cuplike structure, 1.5-3.0 mm long, the margin irregular
or with irregular, triangular teeth, to 0.8 mm long, ciliolate outside; corolla yellow
or greenish white, the tube 4-5 mm long, carnose, pubescent outside, the lobes
5, ovate oblong or triangular, 1 mm long; stamens 5, the anthers oblong, 1.3-2.0
mm long, obtuse, often minutely apiculate, the filaments 3 mm long, attached (in
bud) below the middle of the tube; ovarian disc compressed rotund, ca. 0.35 mm
long, 0.5 mm wide, the style slender, 1.5-2.0 mm long, the stigmas 2, digitiform,
1.0-1.8 mm long. Fruits succulent, china blue or blue black, ellipsoid to subro-
tund, to 1 cm long, the pyrenes costate, glabrous to occasionally puberulent, the
persistent calyx cupuliform, to 1.5 mm long.

Psychotria brachiata is known from Central America, South America, the
West Indies, and Trinidad. Steyermark (1972) regards P. caerulea as distinct
from P, brachiata, stressing characteristics of the calyx, corolla, and inflores-

F-

cence. I regard them as conspecific, agreeing with Standley's treatment in the
Flora of Peru (1936) in which P, caerulea is reduced to the synonomy of P.
brachiata, Duke 13615 lists the common name ''Pugabaca pacuru" of the Choco
Indians, the leaves being used as a purgative.

BOCAS DEL toro: Ri'o Terlbe between Quebrada Huron and Puerto Palenque, 400-800 ft, Kirk-
hride & Duke 503, 506 (both MO). Chiriquicito, Lenis et al, 1973 (COL, MO, UC, VEN). Without

352 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol, 67

specific locality, Wedel 177 (MO). Chiriqui Lagoon, Wedel 1329 (MO), 1638, 2196 A (both US). Rio
Cricamola between Finca St. Louis and Konkinloe, Woodson et al. 1928 (MO), canal zone: France
Field, Blum & Dnyer 2123 (MO). Road S-11 NW of Escobal, Croat 12457 (MO). Fort Sherman,
Dwyer 6686 (MO); Dwyer & Robyns 173 (MO). Chagres, Fendler 63 (MO). Coco Solo, Gentry 6489
(MO). Pipeline Road, Limbo Hunt Club, Kennedy & Tyson 1868 (MO). Agua Clara, F////Vri99i (MO).
Fort Sherman, Standlev 31072 (US); Tyson 1649 (MO); Tyson & Blum 1657 (MO), barro Colorado
island: Croat 11231, 11699, 12184, 13251 A, 14555, 15149, 15252 A (all MO); Shattuck 1069 (MO);
Standley 40997 (US), cocle: Boca del Toabre, confluence of Rio Toabre and Rio Code del Norte,
Lewis et al. 4435 (COL, MO, SCZ, UC, VEN). colon: Las Cruces Trail at Portobelo, Ebinger 109,
257 (both MO). Achiote, Tyson et aL 4555 (MO), darien: Without other locality, Bristan 165 (MO).
Cana, Croat 27316 (MO). Loma Cuasi behind Manene, Duke 13615 (MO). Between Paya and Palo de
Los Letras, Duke & Kirkbride 14036 (MO). E of Tres Bocas, shortest headwaters of Rio Cuasi, Kirk-
bride & Duke 1158 (MO). Cana, Stern et aL 467, 478, 668 (all MO, US). LOS santos: Loma Prieta,
Cerro Grande, 2400-2800 ft, Lewis et al. 2198 (MO, UC). Panama: Altos del Rio Pacora, 2500 ft,
Lewis et al. 2322 (COL, DUKE, MO), san blas: 15-20 km WSW of Puerto Obaldia, Mori et al. 6852
(MO). VERAGUAS: Valley of Rio Dos Bocas, 11-13 km beyond Agriculture School, Santa Fe, 350-500
m, Croat 25717 (MO). 5 mi W of Santa Fe, past Escuela Agricola, Liesner 846 (MO).

10. Psychotria calophylla Standley, Contr. U.S. Natl. Herb. 18: 129. 1916.
type: Panama, Fendler 60 (US, holotype).

P. gatunensis Dwyer & Hayden, Ann. Missouri Bot. Card. 54: 142. 1967. type: Panama, 7a/7/7.v/oAi

1554 (MO).
/*. grandicarpa Dwyer & Hayden, Ann. Missouri Bot. Card. 55: 42. 1968. type: Panama, Dwyer

7244 (MO).

Trees to 8 m tall or shrubs, the branchlets at first terete, ultimately angular,
often with porous pith, minutely farinose puberulent or ferrugineous villose.
Leaves oblong, oblong rotund or obovate, 13-28 cm long, 3.7-15.5 cm wide,
tapering obtusely, widely cuneate or rounded at the apex, the acumen none or
to 0.5 cm long, basally acute, often abruptly contracted with a spatulate appear-
ance, often markedly inequilateral at the base, the costa subplane above, often
grooved, prominent beneath, to 0,4 cm wide proximally, the lateral veins (1 1-)13-

19(-30), the lowermost subperpendicular, the upper veins often arcuate, subcor-
iaceous, drying deep red brown or gray green, glabrate or mealy punctate above,
beneath ferrugineous villosulose, especially on the costa and the veins; petioles
stout, 1.5-4.0 cm long, to 0.35 cm wide, glabrate to pubescent, the lamina often
decurrent; stipules deciduous, free, compressed rectangular, ca, 4 mm long,
densely ferrugineous pubescent, coriaceous, with 2 proximate, subulate awns, to
6 mm long, ca. 2 mm wide at the base. Inflorescences terminal, solitary, thrysoid
paniculate, to 15 cm long, to 15 cm wide, densely ferrugineous pubescent, sessile
or with peduncles 2.5-5.5 cm long, stout, 0.3-0.4 cm wide, the branches of 2-3
well-spaced pairs, 2.5-5.5 cm apart, at least the lowermost 2 pairs opposite,
arcuate or sharply ascending, the lowermost to 7 cm long, the flowers in globose
fascicles, to 1.5 cm wide; bracts triangular ovate, to 2 mm long, acuminate.
Flowers subsessile or sessile; hypanthium subrotund, ca. 1.8 mm long, densely
ferrugineous puberulent, the calycine cup scarcely measurable, glabrous and
eglandular inside, the teeth 5, triangular or widely oblong, 0.2-2.0 mm long;
corolla white or greenish white, the tube cylindrical, inflated above, 3-6 mm long,
minutely puberulent outside and densely villose near the mouth, the hairs to 2.2
mm long, the lobes 5, rectangular oblong, to 3 mm long, galeate and crassate;
stamens 5, the anthers oblong, 1.2-1.7 mm long, the filaments ca. 1 mm long;
styles short and long, 1.5-6.0 mm long, the stigmas 2, 0.65-0.8 mm long. Fruits

1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 353

elliptic, often falcate (by abortion of one seed?), 1.3-1.5 cm long, 0.6-0.7 cm

wide, each pyrene 5-ribbed, sparsely pubescent, drying black; pedicels to 0.4 cm
long.

Psychotria calophylla is limited to three provinces of Panama and to Costa
Rica. Dr. Croat's two collections in Costa Rica have an unusual number of lateral
veins, numbering between 20 and 30.

CANAL zone: Chagres, Fendler 60 (MO, US). Pipeline Road, 5-10 mi N of Gamboa, Gentry 2659
(MO). 6-8 mi N of Gamboa, Gentry & Dwyer 3361 (MO). W of Limon Bay, Gatiin Locks and Gatun
Lake, Johnston 1554 (MO). Road S-8 and road to towers at Loma la Toba, Nee 8946 (MO). Pipeline
Road, 10 km NW of Gamboa, Nee 9600 (MO), colon: Miguel de la Borda, Croat 10036 (MO). Santa
Rita Ridge, Dwyer 8572 (MO). Rio Guanche, Gentry 8790 (MO). Santa Rita Ridge Road, 4 mi from
Transisthmian Highway, Gentry^ et al. 8867 (MO, US). Salud, Lao & HolJridge 207 (MO). Santa Rita
Road, ca. 8.5 km from Boyd-Roosevelt Highway, Mori & Kallunki 2076 (MO). Rio Guanche, ca. 2.5
km upriver from bridge to Porlobelo, Mori <& Kallunki 3719 (MO). Santa Rita Ridge Road, 10.2 km
from Boyd-Roosevelt Highway, Mori & Kallunki 5065 (MO). Ri'o Guanche, Mori & Witherspoon
7951 (MO). Rio Guanche, 6 km S of Portobelo, Nee & Gentry 8688 (MO). Achiote, Tyson et al. 4532
(MO). Rio Guanche, 2 km upstream from bridge, Witherspoon 7973 (MO). Panama: Cerro Azul,
Duke 9363 (MO). Cerro Jefe, Dwyer et al, 7244 (MO); Gentry et al 3513 (MO). El Llano-Carti Road,
9,6-11.0 km from Interamerican Highway, 1100-1200 ft, Mori & Kallunki 3528 (MO), costa rica:
limon: Near Rio Sixaola ca. Vi mi SW of Bambu, 15 m, Croat 43250 (MO). Between Limon and
Shiroles, along Rio Sixaola, 0.9 mi SW of Bambii, Croat 43303 (MO).

11. Psychotria capacifolia Dwyer. ^^ type: Panama, Kirkhride & Duke 601 (MO,

holotype).

SubshrubSy succulent, erect, unbranching, to 1 m tall, the stem smooth.
Leaves large, oblong, to 40 cm long, to 20 cm wide, widely deltoid at the apex,
often ultimately obtuse, basally widely cuneate to obtuse, the costa subplane and
canaliculate above, somewhat prominent beneath, to 0.4 cm wide, the lateral
veins ca. 30, spreading arcuate, to 1.8 cm apart, conspicuous beneath, the inter-
venal areas with the veinlets slender, spreading, thinly coriaceous when dry,
brown, discolorous; petioles to 10 cm long, slightly alate, diffusely ciliate; stipules
not seen. Inflorescences axillary, narrowly paniculate, to 50 cm long, to 24 cm
wide, the peduncle elongate, the few branches opposite, disposed in about 6
pairs. Flowers with the hypanthium subrotund, the calycine cup to 1 mm long,
the lobes 5, triangular, to 0.4 mm long, obtuse or acute, sparsely ciliate; corolla
white, the tube subcampanulate, to 3 mm long, expanded at the apex, to 2.5 mm
wide, ca. 0.4 mm wide toward the base, thinly carnose, glabrous outside, diffusely
pilose within near the filament attachment, the lobes 5, scarcely shorter than the
tube, ovate oblong, 1.5-1.9 mm long, cuneate toward the apex with a round mass
of hairs on the adaxial surface at the apex; stamens 5, exserted, the anthers
oblong, ca. 1.1 mm long, the filaments ca. 1 mm long, arising near the mouth;

^"^ Psychotria capacifolia Dwyer, spec. nov. Suffrutices succulenti erecti simplices, ad 1 m alti.
Folia magna oblonga, ad 40 cm longa, ad 20 cm lata, apice lato-deltoidea venis lateralibus ca. 30,
tenui-coriacea in sicco; petiolis ad 10 cm longis. Inftorescentiae axillares angusto-paniculatae, ad 0.5
m longae, ad 24 cm latae, pedunculo longo ramis oppositis in jugis 6 dispositis. Flores cupula calycis
ad 1 mm longa, lobis 5, ad 0.4 mm longis, sparso-ciliatis; corolla tubo ad 3 mm longo, extus glabro
lobis 1.5-1.8 mm longis apice caespite rotundo adaxiali ciliorum praeditis; staminibus 5, antheris
oblongis, ca. 1.1 mm longis. Fructus viridi-lutei.

354 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

Style slender, ca. 4 mm long, the stigmas 2, linear, ca. 2 mm long. Fruits yellow
green.

Psychotria capacifolia is known only from Panama. It is distinguished by
being a small succulent subshrub with elongate leaf blades supported by elongate
petioles. As is characteristic of succulent species of Psychotria, the inflorescence
is axillary. The inflorescence is as long or longer than any Psychotria in Panama.
A diagnostic floral character is the ball of hairs borne on the inside tip of the
corolla lobe.

BOCAS DEL TORo: Cloud forest above Quebrada Huron on Cerro Bonyik, 500-1200 ft. Kirkhride
& Duke 601 (MO), darien: Serrani'a de Pirre, along ascent of Serrania de Pirre above Cana gold mine
between Ri'o Cana and Rio Escucha Ruido, 600-1000 m, Croat 37731 (MO).

12. Psychotria capitata Ruiz & Pavon, Fl. Peruv. Chil. 2: 59. 1799. type: Chin-
chao, Huanuco, Peru, Ruiz & Pavon. Not seen.

P. inunJata Benth.. J. Bot. (Hooker) 3: 229. I84L type: Essequibo, British Guiana, Schomhurgk 27.
Not seen.

P. alhonervia Standley, Contr. U.S. Natl. Herb. 18: 131. 1916. Not Jacq. type: Panama, FenJIcr 62
(US, holotype; isotype, MO).

Palkoiirea stcvensonil Standley, Trop. Woods 16, 32. 1928. type: Belize, Stevenson 10683 (NY,
holotype).

12a. Psychotria capitata Ruiz & Pavon ssp. capitata.

Shrubs or trees to 4 m tall, the branchlets ramose, terete, smooth, glabrous,
the pith abundant. Leaves elliptic oblong, 4-17 cm long, 1.5-7.0 cm wide, apically
deltoid to subacute, occasionally rounded with an acumen to 1.5 cm long, often
falcate, basally cuneate or occasionally obtuse, the costa prominulous above,
prominent beneath, the lateral veins 12-18(-20), conspicuous, widely arcuate,
usually drying white in contrast to the lamina, prominulous above and beneath
with 1 (or 2-3) irregular subparallel veins between adjacent lateral veins, these
soon disappearing or branching, the intervenal areas spreading reticulate, stiffly
papyraceous, concolorous, drying green, glabrous; petioles to 2 cm long, often
reflexed, glabrous; stipules free, often persistent, especially at the tips of the
twigs, oblong, ovate oblong, 10-20 mm long, 4.5-5.5 mm wide at the base, vari-
able at the apex, usually 2-lobed, the lobes 1-2 mm long or more than 2 times the
length of the body, rarely undivided and simply acute, scarious and puberulent.
Inflorescences terminal, solitary, ovate oblong, oblong or pyramidal, 4-14 cm
long, 2.5-8.0 cm wide, puberulent to glabrate, the peduncles 3-6 cm long, usually
strict, the lowermost branches 3-4-verticillate or disposed as a pair, the upper
pairs usually 2 or 3 and opposite; bracts linear or lanceolate, to 1.5 cm long,
about equaling the flowers. Flowers sessile; hypanthium rotund, ca. 1.5 mm in
diam., the calycine cup ca. 0.5 mm long, glabrous, petaloid, the lobes 4-5, dis-
posed as irregular points or to 1 mm long; corolla white, white with yellow throat
or green, the tube 5-9 mm long, glabrous outside, pilose within, the lobes 5,
oblong, 2.5-4.5 mm long, glabrous; stamens 5, the anthers oblong, ca. 1.5 mm
long, sessile or the filaments to 4 mm long; ovarian disc bilobed, to 1 mm long,
the style 3-7 mm long, somewhat expanded above the middle. Fruits ovate.

1980] DWYER— FLORA OF PANAMA (Family 179. Ruhiaceae) 355

oblong or subrotund, 4-6 mm long, ca. 3.5 mm wide, each pyrene 4-5 costate,
glabrous, black or purple.

Psychotria capitata is known from Central America, tropical South America,
and Trinidad and Tobago. This is one of the easiest of the Panamanian species
of Psychotria to recognize because of the stiff leaves which in the field rattle in
the breeze, as well as the usually ghostly white appearance of the inflorescence.
Croat on two collections from Barro Colorado Island notes that the flowers are

butterflies

Heliconius

terfly, a famihar polhnator of the Cucurbitaceae, Steyermark (1972) distinguished
P. capitata subsp. amplifolia (Raeuschel) Steyermark as occurring in Central
America including Panama, as well as in French Guiana, Surinam, and British
Guiana,

CANAL zone: Madden Dam, Dwyer 6938 (MO). Fort Sherman, Hayden 86 (MO). Top of Cerro
Pelado, I km N of Gamboa, Nee 8826 (MO). Road C-29, 6 km E of Gamboa, Nee 8989 (MO). Fort
Sherman, Tyson & Dwyer 1239 (MO), barro Colorado island: Croat 4344, 4349, 6652, 10910,
10985, 12414, 14932, 14959, 14960 (all MO); Ebinger 570 (MO); Shattuck 1090 (MO), cocle: El Valle,
Allen 1896, 3536 (both MO). Las Margaritas and El Valle, Woodson et al. 1759 (A, MO), colon:
Santa Rita Ridge, Correa & Dressier 890 (MO); Dwyer & Gentry 8351 (MO); Dnyer 8419 (MO);
Dwyer et al. 8986, 8996 (both MO). SW of Pina, Lewis et al 1851 (MO, UC). Between Rio Piedras
and Puerto Pilon, Lewis et al 3211 (COL, MO, UC, VEN). Santa Rita Ridge, Lewis et al 5386 (MO,
SCZ, UC); Mori & Kallunki 1788 (MO). Rio Boqueron to Boqueron, Mine #2 N of Peluca Hydro-
graphic Station, Mori & Kallunki 2432 (MO). Rio Guanche, 6 km S of Portobelo, Nee & Gentry 8688
(MO). 4 km NW of Salamanca, 13 km NE of Buenos Aires, Nee 9076 (MO). San Lorenzo, Quistgaard
7 (MO). NE of Santa Rita Ridge, Wilbur & Weaver 10820 (MO), darien: Rio Sancanti, ca. 2 mi
upstream from Piria, Duke 14363 (MO). Panama: Cerro Jefe, Busey 799 (MO). Cerro Azul, Carrera
6 (MO); Croat 11552 (MO). Cerro Campana, Croat 12101 (MO). Goofy Lake to Cerro Jefe, Correa
& Dressier 581 (MO). Cerro Jefe, Davidse & D'Arcy 10, 112 (MO), Panama Viejo, Duke 5743, 5744
(both MO). Cerro Jefe, Duke 9357, 9399 (both MO). Cerro Azul, Dwyer 1877, 3075, 7075 (all MO).
Cerro Jefe, Dwyer 8483 (MO). 1.5 mi S of Goofy Lake, Dwyer & Hayden 8031 (GH, MO, US). Cerro
Jefe and La Eneida, Dwyer et al 8209 (MO). Cerro Azul, Ebinger 414 (MO). La Campana, Cerro
Campana, Ebinger 934 (MO). Cerro Jefe, Elias & Hayden 1803 (MO, UC, VEN); Gentry 2123, 2127,
6778 (all MO). 4.8 mi N of Highway, W of El Llano, Gentry 5095 (MO). Cerro Azul, Gomez-Pompa
3393, 3420 (both MO). Rio Tocumen, N of Chepo Road, Hunter & Allen 227 (MO). Cerro Jefe,
Kirkbride & Crebbs 35 (MO). Cerro Campana, Lewis et al 1944 (GH, MO). Altos del Rio Pacora,
Lewis et al 2317 (MO, UC), 2318 (MO, UC, VEN). Cerro Campana, Luteyn 963. 1031 (both MO).
Gorgas Memorial Labs, 5-10 km NE of Altos de Pacora, 600 m, Mori & Kallunki 3328 (MO). 4.5 km
N of Lago Cerro Azul, 675 m, Nee 7013 (MO). Goofy Lake, Oliver et al 2680 (MO, UC). Cerro
Azul, Porter et al 4102 (MO); Stimson et al 5165 (MO); Tyson 2074, 2085, 2092, 4065, 4086 (all
MO). Cerro Jefe, Tyson et al 3334, 3382, 4360, 4370 (all MO). Cermefio, Varela 24 (MO). Arraijan,
Woodson et al 775 (A, MO), san blas: E of Cangandi-Mandinga airport road, Duke 14795 (MO,
OH). Mainland opposite Playon Chico, Gentry 6394 (MO). Between the Atlantic Coast and Conti-
nental Divide, opposite Carti, Kennedy 2608 (MO), veraguas: Santa Fe, slopes of Cerro Tute,
Gentry 6234, 6235 (both MO).

12b. Psychotria capitata subsp. fissistipularis Dwyer. ^^ type: Panama, Croat

10037 (MO, holotype).

Similar to Psychotria capitata subsp. capitata except stipules to 15 mm long,
the 2 lobes triangular oblong, longer than the body of the stipule, erect or some-

88

Psychotria capitata suhsp. fissistipularis Dwyer, subsp. nov. Differt a. P. capitata subsp. cap-
itata stipulis ad 15 mm longis, lobis 2 triangulari-oblongis, corpore longioribus margine ciholato;
bracteis involucratis, ovatis vel ovato-oblongis, 5.0-6.5 mm longis, 2.5-3.0 mm latis.

356 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67

what falcate, ciliolate marginally; bracts of each floral cluster involucrate, ovate
or ovate oblong, 5.0-6.5 mm long, 2.5-3.0 mm wide, venose, marginally ciliolate.

CANAL zone: Chagres, Fendler 62 (MO, US), colon: Miguel de la Borda. edge of forest near
beach, Croat 10037 (MO).

12c. Psychotria capitata ssp. rubra Dwyer.^^ type: Panama, Folsom & Skof^ 2454

(MO).

Similar to P, capitata ssp. capitata except the bracts are red and t