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Flora of Panama. Part IX. Family 172.
Bignoniaceae
Author(s): Robert E. Woodson, Jr., Robert W. Schery and Alwyn H. Gentry
Source: Annals of the Missouri Botanical Garden, Vol. 60, No. 3 (1973), pp. 781-977
Published by: Missouri Botanical Garden Press
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FLORA OF PANAMA'
BY ROBERT E. WOODSON, JR. AND ROBERT W. SCHERY
AND COLLABORATORS
Part IX
FAMILY 172. BIGNONIACEAE2

ALWYN H. GENTRY3
Woody trees, shrubs, or vines, rarely herbaceous; outer scales of axillary buds
often pseudostipular and sometimes foliaceous; glandular fields often present
between the petioles and rarely at the petiole apex. Leaves usually opposite,
palmately or pinnately compound, sometimes simple, the terminal leaflet often
replaced by a tendril. Inflorescence terminal or axillary, a panicle or raceme,
sometimes reduced to a fascicle or a single flower. Flowers with the calyx syn-
sepalous, usually more or less cupular, truncate to variously lobed or spathaceously
split; corolla sympetalous with a conspicuous tube and 5 usually imbricate lobes;
stamens inserted on corolla tube, didynamous, usually 4, rarely 2, a posterior
staminode usually present, anthers with 1 or 2 thecae, these usually divergent or
divaricate and held against the adaxial side of corolla tube, the filaments slender,
ovary superior, 1 or 2 locular, the ovules numerous, attached to 4 vertical placen-
tal ridges, these axile in 2-locular ovaries and parietal in unilocular ovaries (a
single placental ridge in each locule in some species of Schlegelia), style slender,
stigma bilamellate; a nectariferous disc usually conspicuous, sometimes lacking.
Fruit a capsule with dehiscence perpendicular or parallel to the septum or a berry
or hard-shelled pepo (calabash); seeds without endosperm, somewhat flattened,
usually winged, the cotyledons foliaceous.
A family of about 120 genera and 650 species, mostly tropical with a few
genera- in warm temperate regions. The family is divided into three major tribes
and several minor ones. The Crescentieae and Tecomeae are found in both the
Old and New Worlds. The Bignonieae sensu stricto are restricted to the New
World. In Panama 35 genera and 75 species occur naturally, and an additional
7 genera and 11 species occur in cultivation.
The family is predominantly woody and includes several genera of large trees
and many lianas. Opposite, compound leaves characterize the great majority of
species of the family. The bud scales, termed pseudostipules, are often conspicu-
1Assisted by National Science Foundation Grant GB-5674 (Thomas B. Croat, principal

investigator).
2 Assisted by a National Science Foundation graduate fellowship and pre-doctoral disserta-
tion improvement grant.
'Missouri Botanical Garden, 2315 Tower Grove Avenue, St. Louis, Missouri 63110.
ANN. MISSOURi BOT. GARD. 60: 781-977. 1973.
782 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 60
ous and provide useful taxonomic characters. Presence or absence of concen-

trations of peltate glands into glandular fields at the nodes (interpetiolar glandu-
lar fields) or, less commonly, at the apex of the petiole (petiolar glandular fields)
is another taxonomically useful character. However, glandular fields disappear
on older branchlets and in some species may be either present or absent, so the
character must be used with caution. The flowers are variously pollinated by
bees, birds, butterflies, moths, or bats. Seeds are mostly wind-dispersed, but
some are dispersed by water or by mammals.
The major economic importance of the family is in horticulture with many
species widely cultivated for their showy flowers. A number of species of
Tecomeae are important timber trees. The hard shells of the fruits of Crescentia
are widely employed as household utensils.
Useful references:
Bureau, E. & K. Schumann. Bignoniaceae. In C. F. P. von Martius, "Flora
Brasiliensis." 8(2). 1896-97.
Gentry, A. H. Bignoniaceae of OTS lowland study sites. In C. Schnell (edi-
tor), "Handbook for Tropical Biology in Costa Rica." Pp. 26-16 (6 pp). San
Jose. 1971.
. Generic realignments in Central American Bignoniaceae. Brittonia
25: 226-242. 1973.
Sandwith, N. Y. Bignoniaceae. In "Flora of Trinidad and Tobago." 2. Port of
Spain. 1954.
Seibert, R. J. Botany of the Maya Area: Miscellaneous papers XXI: The
Bignoniaceae of the Maya Area. Carnegie Inst. Washington Publ. 522: 375-434.
1940.
a. Herbaceous vines; inflorescence subspicate, the upper flowers mostly sterile; ovary
4-locular; capsule densely uncinate-spiny, bur-like, thd valves not splitting to the base
------------------------------------------------------------------------ 40. Tourrettia
aa. Woody lianas or trees; inflorescence never spicate, without sterile flowers; ovary 1-2-
locular, capsule not spiny with uncinate barbs, or if spiny not bur-like and the 2 valves
completely dehiscent.
b. Trees or shrubs (Podranea and Tecomaria, both cultivated, are often vines); leaves
various but not 2-foliolate and never tendrillate; stems without phloem arms in
cross section; fruits dehiscing perpendicular to the septum or indehiscent and more
than 7 cm long.
c. Leaves pinnately compound.
d. Leaves bipinnate, flowers purplish-blue; staminode longer than fertile sta-
mens- 15. Jacaranda
dd. Leaves once compound; flowers not bluish; staminode much shorter than
fertile stamens.
e. Shrubs, vines, or small trees; corolla small (less than 6 cm long and 2 cm
wide), tubular-infundibuliform to tubular-campanulate; native or exotic
species.
f. Usually vines; flowers pink to lavender; calyx more than 1.0 cm long;
cultivated -29. Podranea
1973] GENTRY-FLORA OF PANAMA (Family 172. Bignoniaceae) 783
ff. Usually trees or shrubs; flowers yellow to orange-red; calyx less than
0.8 cm long; cultivated or native.
g. Stamens included; flowers yellow; native shrub or small tree
--- - - - - - - - - - - - - - - - - - - - - - - - - - - - -- - - - - - -- - - - - - - - 3 8 . T eco m
gg. Stamens exserted; flowers orange-red; exotic shrub or vine
--- -- - --- - - -- - --- --- - -- - - - -- --- -- - - -- -- - - --- - -- -- - 39. T ecom a
ee. Trees; corolla large (more than 7 cm long and 4 cm wide), broadly cam-
panulate; exotic species.
h. Calyx spathaceous; corolla red-orange; fruit a glabrous capsule
--- - - - -- - -- -- - - - -- - - -- - - - - - - -- - - --- - - -- -- - -- - -- - - 34. S p ata ho d
hh. Calyx cupular, irregularly 2-5-lobed; corolla yellow or brownish ma-
roon; fruit indehiscent or a pubescent capsule.
i. Corolla yellow, strongly pubescent; fruit dehiscent 14. Haplophragma
ii. Corolla brownish-maroon, glabrous; fruit indehiscent - 16. Kigelia
cc. Leaves simple, 3-foliolate, or palmately compound.
j. Leaves alternate, opposite, or fasciculate; corolla thick, usually with a trans-
verse fold in throat, whitish or greenish; the flowers borne singly or in
fascicles (bat pollination syndrome), fruit indehiscent, fibrous-fleshy, or
pulpy with a hard shell.
k. Leaves 3-foliolate, opposite; calyx spathaceously split (see also Den-
drosicus spathicalyx); fruits without hard shell- 25. Parmentiera
kk. Leaves mostly simple, alternate or fasciculate (if compound then fascicu-
late in clusters); calyx not spathaceous (except in D. spathicalyx); fruits
with hard shell (calabash).
1. Leaves alternate; placentation axile, at least at extreme base of ovary,
above parietal on two placentae; seeds large (greater than 13 X 16
mm) -11. Dendrosicus
11. Leaves fasciculate; placentation parietal on four placentae; seeds
small (less than 8 X 6 mm) -8. Crescentia
jj. Leaves opposite; corolla thin, without transverse fold in throat, color various;
inflorescence a raceme or panicle or reduced to a few flowers; fruit a de-
hiscent capsule.
m. Flowers less than 1.8 cm long; anthers pilose; fruits spirally twisted;
leaflets 7-9, often cuneate to the base -13. Godmania
mm. Flowers more than 2 cm long; anthers glabrous; fruits more or less
straight; leaflets (1- )3-7, conspicuously petiolulate (unless 3-foliolate)
--- - - - -- - - - - - - - - - - -- -- - -- - - - - - - - - - - - -- -- - - - - -- - - - - 36. T ab eb u ia
bb. Lianas, sometimes shrubby when young (see also Podranea and Tecomaria); leaves
various but often 2-foliolate and usually in part tendrillate; stems (except Schle-
gelia) with 4-16 phloem arms in cross section; fruits dehiscing parallel to the sep-
tum or indehiscent and less than 4 cm long.
n. Leaves simple, tendrils lacking; placentation axile with a single central placenta
in each locule, sometimes parietal with two bifid placentae toward top of ovary;
fruit spherical, indehiscent -32. Schlegelia
nn. Leaves usually compound, tendrils often present; placentation axile with 2 pla-
centae in each locule; fruit more or less elongate or flattened, dehiscent.
o. Branchlets strongly 6-sided, the angles ribbed.
p. Tendrils apically 3-forked; leaves with dendroid trichomes; calyx double;
corolla conspicuously bilabiate, the upper 2 and lower 3 lobes fused at
anthesis -2. Amphilophium
pp. Tendrils apically 3-15-forked; trichomes all simple; calyx simple; corolla
slightly bilabiate, the lobes separate, subequal, reflexed at maturity
------------------------------------------------------27. Pithecoctenium
oo. Branchlets terete to tetragonal, not 6-ribbed.
q. Leaves 2-3-ternate; branchlets acutely tetragonal and the tendril trifid

------------------------------------------------- ---- 28. P leonotom a
qq.Leaves 2-5-foliolate (sometimes simple in part); branchlets various, if
acutely tetragonal the tendril simple.
r. Corolla split about half its length, white; margins of capsule valves
sharply raised; vegetative parts with odor of cloves- 41. Tynnanthus
rr. Corolla only slightly bilabiate, color various; margins of capsule
valves not sharply raised; without odor of cloves.
s. Corolla orange, lobes valvate in bud; exotic -30. Pyrostegia
ss. Corolla not orange, lobes imbricate in bud; native or exotic.
t. Branchlets conspicuously hollow; fruit narrow (less than 7 mm
wide); leaves punctate -35. Stizophyllum
tt. Branchlets not hollow; fruit more than 8 mm wide; leaves not
punctate.
u. Tendrils uncinate-trifid with claw-like tips; flowers yellow;
stem irregularly many-armed in cross section. I
v. Calyx with a recurved apicule; capsule valves oblong,
woody, splitting in half at maturity -20. Melloa
vv. Calyx apicule wanting or incurved; capsule valves linear,
not woody, not splitting in half -18. Macfadyena
uu. Tendrils simple or trifid without claw-like tips; flowers lav-
ender, white, or yellow; stem symmetrically 4-16-armed in
cross section.
w. Leaves + dendroid-pubescent, at least in vein axils be-
neath; capsule oblong, the valves woody, compressed,
smooth, the seeds thin with broad wings.
x. Interpetiolar glandular fields present; calyx small
(less than 9 X 5 mm); corolla lavender; seed wings
hyaline - -- 42. Xylophragma
xx. Interpetiolar glandular fields absent; calyx large
(greater than 22 X 22 mm); corolla yellow; seed
wings brown -5. Callichlamys
ww.Leaf pubescence of simple trichomes or lepidote scales
or wanting; capsule various but without above combina-
tion of characters.
y. Calyx with narrow, elongate (to 5 mm), twisted
teeth; capsule oblong, conspicuously large tubercu-
late - 22. Onohualcoa
yy. Calyx various but marginal teeth lacking or less
than 1 mm long; capsule oblong or linear, not large-
tuberculate.
z. Anthers pubescent (see also Pachyptera kerere
and Cuspidaria floribunda, these with curved
anther thecae and obvious disc), the thecae
straight; disc absent; capsule pubescent, the mid-
rib and margins raised -17. Lundia
zz. Anthers mostly glabrous (pubescent in some
species of Pachyptera and Cuspidaria); thecae
straight or curved; disc present or absent; cap-
sule glabrous or lepidote, if pubescent then
evenly flattened.
A. Flowers yellow, sometimes with reddish splotches (Mussatia), or white (see also
Cydista aequinoctialis and Arrabidaea spp.); disc present; capsule valves more or less
woody, more than 2.5 cm wide, not echinate.
B. Twigs sharply tetragonal, the angles ribbed; corolla small, less than 2 cm long;
calyx shallowly campanulate, less than 2 mm long - 21. Mussatia
BB. Twigs subterete to subtetragonal, without conspicuous angles; corolla more than
2.5 cm long; calyx cupular, more than 5 mm long.
C. Capsule stipitate, orbicular or elliptic; ovary usually stipitate; corolla glabrous
(see also Tanaecium nocturnum with white flowers) or lepidote, cream or
yellow, not white - -- 3. Anemopaegma
CC. Capsule + oblong; ovary not stipitate; corolla pubescent outside (except
Tanaecium nocturnum), cream, yellow or white.
D. Leaves with petiolar glandular fields; pseudostipules vertically 3-seriate
(see Fig. 23C, page 891) -23. Pachyptera
DD. Leaves without petiolar glandular fields; pseudostipules not vertically
3-seriate.
E. Tendril trifid.
F. Capsule + flattened, wider than thick, basally rounded; inter-
petiolar glandular fields lacking -12. Distictella
FF. Capsule subtetragonal, basally gibbous; interpetiolar glandular
fields present -6. Ceratophytum
EE. Tendril simple.
G. Leaves with almond (cyanide) odor, never with cartilaginous
margin; corolla elongate-tubular, salverform, white- 37. Tanaecium
GG. Leaves without almond odor, usually with at least traces of a
cartilaginous margins; corolla tubular-campanulate or tubular-
infundibuliform, yellow -1. Adenocalymma
AA. Flowers magenta to lavender (some species have white-flowere
or absent; capsule valves not woody unless echinate, mostly less
H. Calyx spathaceous; surface of capsule valves gray, densely lepidote, appearing
rough-crystalline under a lens - 26. Phryganocydia
HH. Calyx not spathaceous; surface of capsule valves rarely
under a lens.
I. Capsule suborbicular, echinate; pseudostipules clusters of cataphylls resem-
bling miniature bromeliads, most conspicuous at base of inflorescence
--- 7. Clytostoma
II. Capsule linear (rarely linear-oblong), not echinate (sometimes verrucose);
pseudostipules not clusters of cataphylls, the inflorescence without cataphylls
at base.
J. Plant with strong onion odor (except P. parvifolia with disc-tipped
tendrils); tendril trifid or ending in a peltate disc; mature capsule drying
straw-colored or light brown, smooth, more than 1.7 cm wide, the mar-
gins not raised; petiolar and interpetiolar glandular fields often present
--- -- - -- - -- - - - _ - -23. Pachyptera
JJ. Plant without onion odor; tendrils simpl
peltate disc; mature capsule usually drying b
cose to sandpaper-surfaced or smooth, less than 2 cm wide (except in
Cydista aequinoctialis with raised margins), the margins raised or not;
petiolar glandular fields only in Paragonia with simple or minutely bifid
phylla) -19. Martinella
tendrils; interpetiolar glandular fields only in Arrabidaea and Scobinaria
both with simple tendrils.
K. Tendril conspicuously trifid; fruit more than 55 cm long, uniformly
flat, glabrous; inflorescence a raceme (see also Cydista hetero-
KK. Tendril simple or minutely bifid; fruit less than 60 cm long,, com-
pressed but with edges raised or sharply angled or the midline
thicker, glabrous or pubescent inflorescence usually a panicle (a
raceme in Cydista heterophylla).
L. Calyx large (more than 18 mm long) and bilabiate; capsule
verrucose -33. Scobinaria
LL. Calyx less than 9 mm long, usually more or less truncate; cap-
sule not verrucose.
M. Corolla glabrous outside; leaflets basally attenuate; pseu-
dostipules foliaceous on terete branchlets; cultivated
--- -- - -- - -- - - -- - -- - -- - -- - - -- - -- - -- - -- - - -- - -- - -- - - 3 1 . S arita e a
MM. Corolla puberulous or lepidote outside at least on lobes;

leaflets basally rounded to broadly cuneate; pseudostipules
only foliaceous if early deciduous or on sharply tetragonal
branchlets; native.
N. Disc wanting; interpetiolar glands absent; stems with
8-16 phloem arms in cross section; branchlets some-
times tetragonal- 10 Cydista
NN. Disc present, obvious; interpetiolar glands often pres-
ent; stems with 4 phloem arms in cross section;
branchlets not tetragonal.
0. Leaflets elliptic, drying gray, with faint sweet
odor when fresh; pseudostipules appressed-coni-
cal; tendril usually minutely bifid; capsule valves
convex, rough -24. Paragonia
00. Leaflets ovate to elliptic or round, not drying
gray, without sweet odor when fresh; pseudo-
stipules inconspicuous or wanting; tendril sim-
ple; capsule valves not convex and rough-surfaced.
P. Capsule valves with raised lines bordering a
median furrow; anther thecae strongly re-
flexed, pollen in tetrads -9. Cuspidaria
PP. Capsule valves uniformly flattened, or the
median nerve and/or margins slightly raised;
anther thecae straight or slightly bent, pollen
grains single -4. Arrabidaea
1. ADENOCALYMMA
Adenocalymma Mart. ex Meisn., Gen. 1: 300; 2: 208. 1840. LECTOTYPE: A.

comosum (Cham.) DC.
Lianas; stems with 4 phloem arms in cross section; branchlets terete, without
interpetiolar glandular fields, pseudostipules small, more or less subulate, not
foliaceous. Leaves 3-foliolate, the terminal leaflet often replaced by a simple
tendril. Inflorescence a narrow axillary or terminal bracteate raceme, each bud
subtended by a caducous glandular bract. Flowers with the calyx cupular, bila-
biate to more or less truncate, usually with conspicuous plate-shaped glands in
upper part; corolla yellow, tubular-infundibuliform to tubular-campanulate, usu-
ally pubescent outside; anthers glabrous, the thecae straight, divaricate or some-
what divergent; pollen ecolpate, the exine alveolate; ovary narrowly cylindrical,
lepidote or puberulent, the ovules 2-seriate in each locule; disc pulvinate. Fruit
an oblong capsule, the valves parallel to the septum, only slightly compressed,
woody, usually with more or less raised lenticels, the median nerve not conspicu-
ous; seeds bialate or almost wingless, the body thick and corky, the wings when
present poorly demarcated, brown at the base and hyaline toward the ends.
a. Leaves ovate, drying shiny gray with a distinct cartilaginous margin; twigs usually dry-
ing dark with pale flecks; corolla tubular-infundibuliform; anther thecae slightly di-
vergent, the connective apiculate -1. A. apurense
aa. Leaves narrowly ovate, drying gray-green without a distinct cartilaginous margin; twigs
drying tannish; corolla tubular-campanulate; anther thecae widely divaricate, the con-
nective not extended -2. A. arthropetiolatum
1. Adenocalymma apurense (H.B.K.) Sandw., Lilloa 3: 461. 1938.-FIG. 1A-D.

Bignonia apurensis H.B.K., Nov. Gen. Sp. P1. 3: 138. 1819. TYPE: Venezuela, Rio Apure,
Humboldt & Bonpland 812 (P).
Adenocalymma inundatum Mart. ex DC., Prodr. 9: 201. 1845. TYPE: Brazil, Para, Martius s.n.
(1819) (M).
A. grenadense Urb., Fedde Repert. 14: 306. 1916. TYPE: Grenada, prope Belmont, Broadway
1798 (C, GH, M, NY).
Tabebuia calderonii Standl., Jour. Washington Acad. Sci. 14: 244. 1924. TYPE: El Salvador,
Sonsonate, Calderon 1666 (US).
Adenocalymma hintonii Sandw., Kew Bull. 1936: 10. 1936. SYNTYPES: Mexico, Mexico, Hinton
3784 (K, MO); Guerrero, Hinton 7259 (K, MO, US).
A. calderonii (Standl.) Seib., Carnegie Inst. Washington Publ. 522: 428. 1940.
Liana to 10 cm in diameter, the bark smooth, gray, often 4-furrowed; twigs

subterete, glabrous to slightly lepidote, usually drying black with whitish lenticels
when young; pseudostipules small, pointed, usually ? ovate. Leaves 3-foliolate
or 2-foliolate with a tendril or tendril scar; leaflets ovate to elliptic ovate, acute
to acuminate, basally rounded, truncate or almost subcordate, 4.5-17 cm long and
2.2-8.8 cm wide, membranaceous to chartaceous, secondary veins 5-7 on a side,
the nerve axils beneath with inconspicuous glandular fields, somewhat lepidote,
at least beneath, usually puberulous at base of midvein above, drying shiny dark
gray or dark olive with a conspicuous narrow cartilaginous margin; tendril sim-
ple, 4-13 cm long; petiolules 0.9-4 cm long, petioles 3.2-6.5 cm long, petiole and
petiolules flattened above, inconspicuously lepidote, slightly puberulous above at
least at base and apex. Inflorescence an axillary raceme, an enclosing deciduous
bract ca. 1 cm long and 1 cm wide with conspicuous sunken glands subtending
each bud, the rachis and pedicels puberulous. Flowers with the calyx cupular,
minutely 5-toothed but bilabiately split for 1-2 mm, 5-8 mm long and 4-6 mm
wide, puberulous, usually with plate-shaped glands; corolla bright yellow, 2.5-6.9
cm long and 0.9-1.9 cm wide at the mouth, the base cylindric, 6-7 mm long and
4-5 mm wide above, infundibuliform-campanulate, the tube often bent somewhat
anteriorly, 1.8-5.0 cm long, the lobes all ? reflexed, 1.0-2.0 cm long; puberulous
outside with simple and thick-stellate trichomes, glandular-lepidote inside on the
lobes and sparsely so in the tube, with longer 1-2-celled trichomes at the level of
stamen insertion; stamens didynamous, included, the anther thecae slightly di-
vergent, slender, 2-3 mm long, the connective with a 1 mm apicule, the longer
filaments 1.4-1.7 cm long, the shorter filaments 0.9-1.3 cm long, staminode 4-5
mm long, inserted 4-7 mm from the base of the corolla tube; pistil 2.4-2.5 cm
long, ovary cylindric, 3-3.5 mm long and 1.5-2 mm wide, lepidote, the ovules
2-seriate, disc pulvinate, 2 mm long and 3 mm wide. Capsule oblong, rounded at
both ends, not compressed, the valves woody and somewhat thickened, the midrib
FIGURE 1. Adenocalymma.-A-D. A. apurense (H.B.K.) Sandw.-A. Habit (X %).

[After Woodson et al. 1578 (MO).]-B. Corolla split open (X '/2).-C. Anther (X 5).
[After Gentry & Tyson 5023 (MO).]-D. Fruit- (X 3/io). [After Hinton 7259, Mexico
(MO).]-E-J. A. arthropetiolatum A. Gentry.-E. Ovary and disc (X 1/2). -F. Calyx and
pistil (x 1/2).-G. Corolla split open (x 1/2).-H. Corolla (x 1/2).-I. Leaf (X 1/2).-J.
Ovary cross section ( X 9). [After Gentry 2460 (MO).]
obscure, 9.5-27 cm long and 2.5-3.1 cm wide, 1.4-2.2 cm thick, glabrous but
rough-surfaced, gray with numerous raised tan lenticels; seeds 1.7-2.1 cm long
and 5.1-7.6 cm wide, the body thick and corky, the wings thin, brown and mem-
branaceous, subhyaline to hyaline at tips and not sharply demarcated.
A vine of the tropical dry forest and edaphically dry regions of the tropical
moist forest, this species is known only from riverine gallery forests over much
of its range; from Mexico to the Guianas. Flowering mostly during the late dry
season and early wet season from mid-April through August, it has also been
collected in flower in February. Although the heavy-bodied seeds have wings,
they are probably water dispersed at least in part. This may account for mature
plants occurring almost invariably adjacent to streams.
The ovate leaflets of this species are useful characters for differentiating it
from A. arthropetiolatum, which has narrowly ovate leaflets. The included anthers
with slightly divergent thecae separate the flowers from those of A. arthro-
petiolatum.
This species has in the past been well known from Mexico to El Salvador as
A. calderonii (Standl.) Seib. or A. hintonii Sandw. and in northern South America
as A. inundatum Mart. ex DC. In large part because of the seeming disjunction
across most of Central America, Sandwith (Kew Bull. 1953: 481. 1954) main-
tained the two as separate species. However collections from Costa Rica and
Panama close the range gap and re-examination of the minor characters purport-
ing to separate the two species disclosed that they are the same. The name
A. apurense (H.B.K.) Sandw. has priority over either of the names in current use.
CANAL ZONE: Salamanca Hydrographic Station, Woodson et al. 1577, 1578 (both A, MO,
NY). CHIRIQUI: 7.5 mi. W of Puerto Armuelles Croat 22471 (MO). Near Puerto Armuelles,
Woodson & Schery 809 (GH, MO, US). DARIEN: Rio Chucunaque between Rios Membrillo
and Subcuti, Duke 8607 (MO). Punta Patifio NNE of Garachine, Gentry 4044 (MO). Rio
Chucunaque above Rio Tuquesa, Stern et al. 946 (G, GH, K, MO, US). LOS SANTOS: Gua'nico
near Tonosi, Stern et al. 1850 (MO, US). PANAMA: Near Madden Lake, Gentry 5023 (MO).
2. Adenocalymma arthropetiolatum A. Gentry.4 TYPE: Panama, Canal Zone,

Pipeline Road, Gentry 2460 (MO).-FIG. 1E-J.
Liana to 3 cm in diameter, the bark smooth, greenish, with inconspicuous

lenticels; branchlets subterete to subtetragonal, glabrous to subscabrous, drying
tan to greenish, pseudo-stipules small, 4-scaled, usually narrow. Leaves 3-folioate
or 2-foliolate with a tendril or tendril scar; leaflets drying olive, narrowly ovate
to almost linear, acute, basally truncate, 5.2-17 cm long and 1.9-7.5 cm wide, ?
chartaceous, secondary veins 5-7 on each side, above scabrous along the midvein,
beneath almost glabrous or puberulous along main veins and sometimes sparsely
4Adenocalymma arthropetiolatum A. Gentry, sp. nov. Frutex scandens, cortice laevi, viridi;
ramuli subteretes, sine consociebus glandularum in nodis inter petioles; pseudostipulae parvae;
folia bifoliolata vel trifoliolata, saepe cirrata simplici cirrho, foliolis anguste ovatis; inflorescentia
floribus in racemis axillaribus dispositis, unoquisque flore per bracteam glandulosam caducam
subtento; calyx cupulatus, 5-dentatus sed leviter bilabiatus, puberulus; corolla aurata, campanu-
lata super basem tubularem; stamina subexserta, thecis glabris, divaricatis; ovarium anguste
cylindricum, puberulum, ovulis in quoque loculo biseriatis; discus cupulatus-pulvinatus; capsula
ignota.
on the surface, with q. few scattered plate-shaped glands; tendril simple, to 14 cm

long; petiolules 0.5-4.1 cm long, the petiole 1.1-6.4 cm long, puberulous, woody
proximally and conspicuously jointed with the petiolules. Inflorescence an axillary
or terminal raceme, a subtending glandular, deciduous bract enclosing each bud,
the rachis and pedicels puberulous. Flowers with a faint musky odor, the calyx
cupular, 5-toothed but more or less bilabiate, 5-15 cm long and 4-10 mm wide,
puberulous with simple and thick-stellate trichomes, usually with conspicuous
submarginal sunken cupular glands; corolla uniformly bright yellow, often a
slightly deeper shade within, campanulate, 4.2-8.2 cm long and 1.1-2.1 cm wide
at the mouth, the base cylindrical, 2.3-2.6 cm long and 0.6-0.7 cm wide, the tube
2.5-5.6 cm long, the lobes often folded or rolled, the upper shorter two lobes
1.2-1.7 cm long, extending horizontally in front of the tube, the longer three lobes
1.8-2.3 cm long, variously reflexed, outside and inside the lobes puberulent with
simple and thick-stellate trichomes, the tube glabrous inside except for slender
2-3-celled gland-tipped trichomes at and just below the level of stamen insertion,
the base of the lobes with plate-shaped glands outside; stamens slightly exserted,
the anthers held underneath the hood formed by 2 projecting upper corolla lobes,
the anther thecae more or less divergent, 5-6 mm long, the filaments more or less
equal, the anterior pair 3.5-3.9 cm long, the posterior pair 3.3-4.0 cm long, the
staminode 12-21 mm long, inserted 2.1-2.6 mm from base of corolla tube, pistil
5.0-5.9 cm long, the ovary narrowly cylindrical, 4-5 mm long and 1.5-2.0 mm
wide, puberulent, the ovules 2-seriate; disc pulvinar-cupulate, 1.5-2.5 mm long,
4-6 mm wide. Capsule unknown.
Apparently restricted to stream banks and lake shores of the tropical moist
forest and premontane wet forest of central Panama, this plant flowers during the
late wet season in October and November and is thus temporally isolated from
its congener A. apurense.
The smooth greenish bark and wiry, strongly bent petiole and petiolule joints
are distinctive. Narrowly ovate leaflets which are slightly 3-nerved at the base
also aid in distinguishing it from A. apurense.
This species is most closely related to A. magdalenense Dugand of the Mag-
dalena Valley of Colombia, which is vegetatively distinguishable mainly by its
much larger, glandular, subfoliaceous pseudostipules. The corolla of A. mag-
dalenense, however, is evenly and narrowly tubular-infundibuliform with
strongly exserted anthers as opposed to A. arthropetiolatum which has its corolla
abruptly tubular-campanulate above a narrowly tubular base and subexserted
anthers held under a hood formed by the forward-projecting upper corolla lobes.
Adenocalymma arthropetiolatum is unusual in the family and especially in
the Bignonieae in its restricted geographical range. Since all collections of this
species have come from along rivers or lakes, the seeds, when discovered, may
prove to be water dispersed. Probably because of this it is restricted to the
drainages of the Rios Bayano and Chagres plus the adjacent Rios Pasiga and
Maestro.
CANAL ZONE: Pipeline Road, Gentry 1435, 2460 (both MO); Barro Colorado Island,
Standley 40912 (US). PANAMA: Rio Aguacate near El Llano, Duke 5828 (MO, SCZ). Near
mouth of Rio Pasiga, Gentry 2205A (MO). Rio Maestro, Gentry 2240 (MO). Rio Piragua
1973] GENTRY--FLORA OF PANAMA (Family 172. Bignoniaceae) 791
(Tabardi), Gentry 2519 (MO). Between Rios Espave and Agua Clara, Gentry 3772 (MO).
Near Madden Lake, Gentry 5011 (MO).
2. AMPHILOPHIUM
Amphilophium Kunth, Jour. Phys. Chim. Hist. Nat. Arts 87: 451. 1818. LECTO-
TYPE: A. paniculatum (L.) H.B.K.
Endoloma Raf., Sylva Tell. 79. 1838. TYPE: E. purpurea Raf. = Amphilophium paniculatum
(L.) H.B.K.
Lianas, stems with 4(-5) phloem arms in cross section; branchlets sharply
hexagonal with conspicuous ribs, without interpetiolar glandular fields; pseudo-
stipules foliaceous, often early deciduous. Leaves 2-3-foliolate, the terminal leaf-
let often replaced by a trifid tendril. Inflorescence a terminal racemose panicle
usually on a short lateral branch. Flowers with the calyx campanulate, double,
with a thick inner tube and a spreading, frilly outer limb; corolla purple at ma-
turity, thick, tubular, bilabiate, split about half its length and the upper 2 lobes
and lower 3 lobes fused, glabrous or somewhat pubescent outside; anthers gla-
brous, the thecae thick, divaricate; pollen grains single, 6-8-colpate, the exine
alveolate; ovary ovate-cylindric, the ovules multiseriate in each locule. Fruit an
oblong-elliptic capsule, the valves parallel to the septum, somewhat compressed,
woody, smooth to tuberculate; seeds thin, puberulous winged, the wings mem-
branaceous, brownish to hyaline at tip and not sharply demarcated from the seed
body.
About 5 species ranging from Mexico and the West Indies throughout tropical
America to northern Argentina.
a. Pseudostipules persistent and conspicuous; capsule compressed, rugose-tuberculate; vege-

tative parts with pronounced pubescence, at least partially of long (1-2 mm) trichomes;
tropical wet forest -2. A. pannosum
aa. Pseudostipules early caducous; capsule only slightly flattened, smooth or rugose-wrin-
kled; vegetative parts lepidote to variously short pubescent with mostly dendroid tri-
chomes; throughout Panama -1. A. paniculatum
1. Amphilophium paniculatum (L.) H.B.K., Nov. Gen. Sp. P1. 3: 116. 1819.
-FIG. 2.
Bignonia paniculata L., Sp. P1. 623. 1753. TYPE: (not seen).
Amphilophium macrophyllum H.B.K., Nov. Gen. Sp. P1. 3: 150. 1819. TYPE: Colombia, near
Turbaco, Bonpland s.n. (F, fragment).
A. mutisii H.B.K., Nov. Gen. Sp. P1. 3: 149, tab. 219. 1819. TYPE: Colombia, Mutis 913 (US).
A. molle Schlecht. & Cham., Linnaea 5: 120. 1830. TYPE: Mexico, Veracruz, Schiede & Deppe
s.n. (not seen).
Endoloma purpurea Raf., Sylva Tell. 80. 1838. (nom. nov. for Bignonia paniculata L.).
Amphilophium vauthieri DC., Prodr. 9: 193. 1845. TYPE: Brazil, Rio de Janeiro, Vauthier
249 (G-DC, P; F, fragment).
A. xerophilum Pittier, Jour. Washington Acad. Sci. 18: 121. 1928. TYPE: Venezuela, Lara,
Sauer 253 (K).
A. paniculatum var. molle (Schlecht. & Cham.) Standl., Publ. Field Mus. Nat. Hist., Bot. Ser.
18: 1114. 1938.
Liana to at least 10 cm in diameter, the bark loose, fibrous, vertically striate;

stem with 4(-5) arms in cross section; branchlets sharply hexagonal with ribbed
FIGURE 2. Amphilophium paniculatum (L.) H.B.K.-A. Habit ( X%//). [After Gentry

3120 (MO).]-B. Ovary cross section ( X 9).-C. Fruit (x X%7). [After Allen 205 (MO).]
-D. Pistil and disc ( X 35) .-E. Flower in dorsal and ventral view ( Xe).. -F. Corolla slit
open (x X%/). [After miscellaneous fresh material.]
angles, lepidote to strongly dendroid pubescent, especially on the ribs, interpetio-

lar glandular fields lacking, a conspicuous ridge connecting opposite petioles;
pseudostipules foliaceous, 3-10 mm long and 3-6 mm wide, early deciduous.
Leaves 2-foliolate except on young plants, sometimes with a tendril; leaflets ovate
to subrotund, acuminate to obtuse, truncate to asymmetrically cordate, 2.5-16 cm
long and 2.1-10.7 cm wide, membranaceous, palmately veined at base, the sec-
ondary veins 4-6 on a side, strongly lepidote above and beneath, variably den-
droid pubescent, sometimes also with simple trichomes especially above, pubes-
cence beneath restricted to midveins and nerve axils or scattered throughout, gray
to brown when dry, lighter beneath; tendril trifid, 7-16 cm long to branching, the
3 arms 0.4-2.5 cm long, often unequal; petiolules 0.6-4.3 cm long; petiole 2.3-7.1
cm long; petiole, petiolules, and tendril all ? tetragonal to hexagonal, lepidote
and dendroid pubescent. Inflorescence a caducously bracteate ? racemose termi-
nal panicle on a short erect lateral branch, the branches lepidote and variably
dendroid pubescent. Flowers with some odor, calyx double, the inner calyx 2-3-
labiate, the outer calyx more or less 5-lipped, densely lepidote, sometimes den-
droid pubescent, especially at the base, 0.7-1.2 cm long and 0.8-1.5 cm wide;
corolla yellowish white when young, turning deep purple, tubular, bilabiate,
2.4-3.5 cm long and 0.7-1.6 cm wide, the upper 2 lobes fused, thick, 1 mm long,
the lower 3 fused, 1.3-1.8 cm long, the tube split 1/3_2/3 its length, all 5 lobes re-
maining loosely fused until forced apart by pollinator, viscid, glabrous to sparsely
short simple pubescent outside, inside mostly glabrous, glandular-lepidote on the
ridge at the level of stamen insertion, below this the epidermis highly convoluted
and glandular; stamens didynamous, the anther thecae divaricate, 2 mm long, the
longer filaments 1.7-1.8 cm long, the shorter filaments 1.4-1.5 cm long, the stami-
node 3-5 mm long, inserted 4-5 mm from base of corolla tube; pistil 2.4-2.7 cm
long, bent 0.7-0.8 cm from the base, the ovary ovate-cylindric, slightly contracted
at base, 2-3 mm long, ca. 2 mm wide, short pubescent with simple and dendroid
trichomes, the ovules more or less 10-seriate in each locule below to only 4-seriate
distally; disc annular-pulvinate, 1 mm long, 3-4 mm wide. Capsule rounded
oblong, somewhat flattened, 3.9-9.5 cm long and 2.5-3.8 cm wide, 1.4-2 cm thick,
the surface dark, somewhat warty-lenticellate, densely lepidote; seeds 1-1.9 cm
long and 2.7-3.0 cm wide, puberulous, the wings membranaceous, brownish, not
sharply demarcated, irregularly hyaline only at extreme tips.
Ecologically this is the most wide-ranging species of Central American Big-

noniaceae, occurring in the lowland tropical dry, moist, and wet forests and also
to 6000 feet in montane cloud forest. The higher elevation forms tend to be more
pubescent and have been designated as A. molle or A. paniculatum var. molle in
Central America. Amphilophium macrophyllum of northern South America is
this same form. There is complete intergradation, and the variation in extent of
pubescence is more or less clinal with elevation. The species ranges from Mexico
and the West Indies to Argentina.
Amphilophium paniculatum flowers in the wet season and early dry season
from May through January. The corolla is pseudocleistogamous, never opening
completely. Its lobes are reduced to two lips which cover the tube and remain
fused until forced apart; even after having been opened they are held strongly
together. The peculiar double calyx serves a dual purpose as well; the thick inner
sleeve surrounds and protects the nectar-filled corolla base, while the outer sleeve
even in bud secretes a sticky substance which attracts wasps, trigonids, and ants,
the latter presumably forming an "ant guard" of the type typical for xylocopid-
flowers. It possibly also acts as a scent attractant. Pollination is of necessity by
robust bees. The flowers reach maximum size at least a day before anthesis but
are yellowish white in color, turning purple with anthesis. Purple flowers are
preferentially visited by pollinators, which enter by landing at the tip of the
corolla, inserting their proboscis into the fusion line between the upper and lower
lips, and pushing the lips apart with their head.
The most useful field character for recognition of this species is the hexagonal
branchlet. The relatively short, ovate pseudostipules, tendrils never with more
than three arms, and the presence of dendroid trichomes distinguish A. panicu-
latum from Pithecoctenium echinatum, which also has 6-sided twigs buit elongate
pseudostipules, up to 15-branched tendrils, and only simple trichomes.
BOCAS DEL TORO: Water Valley, von Wedel 1749 (MO). CANAL ZONE: Near Cerro Viejo,
Blum 1258 (MO, SCZ). Venado Beach, Correa 509 (MO, PMA, SCZ). Boy Scout Camp
Road, Gentry 1376 (MO). Pipeline Road, Gentry 1434, 1579 (both MO); Gentry &
Dressler 1980 (MO). Curundu, Gentry 1594 (MO). Highway K10, Gentry 1959 (MO).
Balboa, Standley 26091 (MO). Sosa Hill, Standley 26487 (GH, K). Curundu, Tyson &
Blum 2530 (MO, SCZ). Barro Colorado Island, Aviles 22 (F); Croat 4684, 9789 (both MO),
12499 (NY, SCZ), 12870 (MO); Foster 1918 (GH, PMA); Frost 213 (F). CHIRIQUI: W
of Rio Chorchita, Gentry 5850 (MO). David airport, Lewis et al. 760 (MO). Near Boquete,
Stern et al. 1144 (GH, K, MO). Cerro Punta, Tyson 5821 (SCZ). COCLE: Rio Grande,
Burch et al. 1164 (GH, MO). El Valle, Gentry 3571 (MO); Lewis et al. 2578 (MO, SCZ);
Pittier 5002 (NY). Rio Grande, Tyson 5243 (SCZ). COLON: Giral near Bat Cave, Blum &
Tyson 520 (MO, SCZ). Santa Rita Ridge, Gentry 1432, 1881 (both MO). DARIEN: Santa
Fe, Duke 14243 (MO). Isla Encanto, Gentry 4021 (MO). 3-4 mi. S of Jaque, Gentry 4130
(MO). Rio Jaque, Gentry 4201 (MO). El Real, Gentry 4511 (MO). Cerro Pirre, Gentry
4645 (MO). HERRERA: Rio Santa Maria, Burch et al. 1186 (GH, MO). Along main road
between Ocui and Chitre, D'Arcy & Croat 4146 (MO). S of Ocu', Gentry 3120( (MO). E of
Las Minas, Gentry 3139 (MO). Santa Maria, Seemann 153 (K). S of Pese, Tyson et al. 2885
(MO, SCZ). N of Macaracas, Tyson et al. 3140 (MO, SCZ). LOS SANTOS: S of Chitre,
Burch et al. 1219 (F, GH, K, MO). PANAMA: Near Nueva Gorgona, Duke 4533 (MO).
Between Chepo and Rio Mamoni, Duke 6056 (MO). San;Jose Island, Erlanson 64 (GH, NY);
Johnston 422 (GH, MO), 644 (GH). S of Pacora, Gentry 1412 (MO). Rio Piragua (Tabardii),
Gentry 2531 (MO). Icanti (Rio Agua Clara), Gentry 2621 (MO, SCZ). Near Rio Espave,
Gentry 3725 (MO). Between Bayano crossing and Rio Partli, Gentry 3810 (MO). Cerro
Campana, Gentry 4925 (MO). Taboga Island, Standley 27112 (GH). Juan Diaz, Zetek 905
(F). SAN BLAS: Puerto Obaldlia, Gentry 1551 (MO). VERAGUAS: Trail between Cafiazas
and foot of Cordillera Central, headwaters of Rio Cafiazas, Allen 205 (MO). Sonai, Allen 1064
(GH, MO). S of Santa Fe, Gentry 2929, 2940 (both MO). Mountains above Santa Fe,
Gentry 2984, 3098 (both MO). Rio Santa Maria S of San Francisco, Gentry 3116 (MO).
W of Santiago, Tyson 5164 (SCZ).
2. Amphilophium pannosum (DC.) Bur. & K. Schum. in Mart., Fl. Bras. 8(2):
209. 1896.
Bignonia pannosa DC., Prodr. 9: 148. 1845. TYPE: Peru, Pavon s.n. (F, fragment and photo).
B. pubescens Pav. ex DC., Prodr. 9: 148. 1845, pro syn, non L.
A. oxylophium Donn. Sm., Bot. Gaz. (Crawfordsville) 31: 119. 1901. TYPE: Costa Rica,
Tucurrique, Tonduz 13045 (MO, US).
Liana to at least 2.5 cm in diameter, branchlets sharply hexagonal with the

angles ribbed, lepidote and strongly villous with long (to 2 mm) reddish yellow
trichomes, especially on ribs, these simple, forked and variously dendroid; inter-
petiolar glands lacking; pseudostipules foliaceous, 0.6-1.6 cm long and 0.6-1.4 cm
wide, persistent. Leaves 3- or 2-foliolate with a tendril or tendril scar; leaflets
ovate, acuminate, cordate to subcordate, 7.6-12.6 cm long and 5.4-8.8 cm wide,
membranaceous, palmately veined at the base, the secondary veins 4-5 on each
side, strongly lepidote above and beneath, villous above with simple or weakly
dendroid trichomes and beneath with much branched dendroid trichomes, golden
brown when dry; tendril trifid, 8-18 cm long to branching, the 3 arms 0.8-1.7 cm
long, often recurved at the tip; petiolules 1.3-3.7 cm long, petiole 3.0-5.1 cm long,
lepidote and villous. Inflorescence a terminal raceme on a short erect lateral
branch, the branches lepidote and dendroid-villous, each flower subtended by a
pair of narrow bracts below the calyx and a single bract below the pedicel. Flower
with the calyx double, the inner calyx 2-3-labiate, the outer calyx more or less
5-lipped, 1.2-1.4 cm long and 1.2-1.9 cm wide, densely lepidote, densely den-
droid-villous below the outer calyx lobes, sparsely so on them; corolla yellowish
white when young, turning deep purple, tubular, 2.2-2.6 cm long and 0.6-0.9 cm
wide, the upper 2 lobes fused and thick, 1 mm long, the lower 3 fused, 6-8 mm
long, the tube bilabiate, split about half its length, the 2 lips remaining loosely
fused until forced apart by pollinator, viscid, mostly glabrous, glandular-lepidote
on the ridge at the level of stamen insertion; stamens didynamous, the anther the-
cae divaricate, 2 mm long, the longer filaments 1.4-1.6 cm long, the shorter fila-
ments 1.2-1.3 cm long, the staminode 3-4 mm long, inserted 5 mm from the base
of corolla tube; pistil 2.1-2.3 cm long, bent near the base, ovary truncate-conical,
2.5 mm long, 2 mm wide, rough-surfaced and lepidote, the ovules multi-seriate
in each locule; disc annular-pulvinate, 1.5 mm long, 3.5 mm wide. Capsule
(Lehmann 7691, Cali, Colombia, K) elliptic, compressed, 7.5 cm long and 4 cm
wide, the valves woody, densely rugose-tuberculate with the closely packed tuber-
cles ca. 2 mm long, pubescent with dendroid and simple trichomes, appearing
yellowish from the trichomes; seed 1.2-1.3 cm long and 4.7-4.9 cm wide, bialate,
the wings thin, brown to almost hyaline at the tip, distinct from the darker brown
puberulous seed body.
This species is a rarely collected but locally common vine restricted to the
tropical wet forest; it ranges from Costa Rica to Argentina. The flowers have
been collected in January, July, and August which is no indication of seasonality.
This species is closely allied to A. paniculatum. The species can be distinguished
vegetatively by the conspicuous, persistent pseudostipules, more conspicuous
pubescence, and longer trichomes in A. pannosum. On Santa Rita Ridge, the two
species occur within a kilometer of each other with no indication of introgression.
I have seen only a fragment of the type of Bignonia pannosa, but a photograph
of the type specimen agrees with the Central American material. Acceptance of
the name A. pannosum is also based on Sandwith's notation on a herbarium sheet
of it at Kew (Haught 3399) that he had matched that specimen with the Pavon
type collection at Madrid (MA). Some recent workers have argued that A. pan-
nosum is synonymous with A. paniculatum (including A. macrophyllum), but
until the type is re-examined Sandwith's interpretation should be accepted.
COLON: Santa Rita Ridge, Dwyer & Gentry 9380 (MO); Dwyer et al. 9037 (MO).
3. ANEMOPAEGMA
Anemopaegma Mart. ex Meisn., Gen. 1: 300; 2: 208. 1840, nom. cons. TYPE: A.
mirandum (Cham.) DC.
Cupulissa Raf., Fl. Tell. 2: 57. 1837, nom. rej. TYPE: C. grandifolia (Jacq.) Raf. = Anemo-
paegma grandifolia (Jacq.) Merill & Sandw.
Platolaria Raf., Sylva Tell. 78. 1838, nom. rej. TYPE: P. flavescens Raf. = Anemopaegma
orbiculatum (Jacq.) DC.
Lianas (in Central America), stems with (4-)8 phloem arms in cross section;
branchlets terete without interpetiolar glandular fields; pseudostipules foliaceous
or lacking. Leaves 2-5-foliolate, often with a simple or trifid terminal tendril.
Inflorescence an axillary or terminal raceme. Flowers with the calyx cupular,
truncate, usually with glands near the margin; corolla bright to pale yellow, tubu-
lar-campanulate, glabrous or glandular-lepidote outside; anthers glabrous, the
thecae straight, divaricate; pollen grains simple, ecolpate or 5-colpate, the exine
reticulate; ovary ellipsoid, usually stipitate, lepidote or puberulous, the ovules
2-6-seriate in each locule; disc pulvinate. Fruit an elliptic to orbicular capsule,
the valves parallel to the septum, usually strongly compressed, smooth; seeds
flat, completely surrounded by a broad hyaline-membranaceous wing, or wingless
and the seed body larger, brown and corky.
About 30 species; continental tropical America from Brazil and Argentina to

Mexico. Anemopaegma is taxonomically one of the more difficult genera of
Bignoniaceae with many species showing great phenotypic plasticity and many
of the local phenotypes designated as species. Whether all the morphologically
and geographically distinguishable species currently recognized are valid is ques-
tionable, but where to draw specific lines is unclear.
a. Leaves palmately 5-foliolate or 4-foliolate with a tendril or tendril scar; fruit suborbicu-
lar -3. A. orbiculatum
aa. Leaves 2-foliolate, often with a tendril or tendril scar; fruit elliptic or oblong-elliptic
(unknown in A. santa-ritense).
b. Pseudostipules foliaceous; tendril simple; leaflets narrowly elliptic; fruit ellipsoidal,
not compressed, the seeds corky, wingless -2. A. chrysoleucum
bb. Pseudostipules inconspicuous or lacking; tendril trifid; leaflets ovate to ovate-
elliptic; fruit (where known) oblong, compressed, the seeds with broad hyaline mem-
branaceous wings.
c. Leaves conspicuously pubescent beneath -4. A. puberulum
cc. Leaves glabrous beneath.
d. Corolla tube glabrous outside; surface of leaflets shiny, varnished--looking, the
margin not involute -1. A. chrysanthum
dd. Corolla tube glandular-lepidote outside; surface of leaflets dull, the margin
more or less involute -5. A. santa-ritense
1. Anemopaegma chrysanthum Dugand, Caldasia 4: 307. 1947. TYPE: Colom-

bia, Cauca, Pacific coast, Cuatrecasas 14269 (COL, holotype; F, isotype).
Liana to at least 2 cm in diameter, the bark smooth, vertically striate; stem
with 4 phloem arms in cross section; branchlets subtetragonal, glabrous, the nodes
without interpetiolar glandular fields; pseudostipules minute, acute, paired in
each axil. Leaves 2-foliolate with a tendril or tendril scar; leaflets ovate to ovate-
elliptic, acute or shortly acuminate, basally cuneate to rounded with a brief at-
D F
FIGURE 3. Anemopaegma.-A-B. A. orbiculatum (Jacq.) DC.-A. Habit (x '/2).

[After Dressler 4051 (MO).]-B. Fruit (X %). [After Croat 12742 (MO).]-C-F. A.
chrysoleucum (H.B.K.) Sandw.-C. Pistil and disc (X 1/2).-D. Calyx (X '/2) .-E. Corolla
(X '/2).-F. Ovary cross section (x 10). [After Gentry 3848 (MO).]
tenuation at the petiole apex, 9.4-19.5 cm long and 4.7-12.5 cm wide, chartaceous,
the secondary veins 4-7 on each side, strongly minute-lepidote beneath, especially
in the lower vein axils, usually minutely and inconspicuously scabrous at the base
of the main vein above, sometimes beneath, olive when dry, slightly paler be-
neath, above shiny and varnished-looking under a lens; tendril usually trifid, 6-13
cm long to branching, the arms 0.1-1.2 cm long; petiolules -+- terete, often twisted,
0.8-3.7 cm long, petiole 2.9-3.9 cm long, somewhat lepidote or minutely subpu-
berulous. Inflorescence a 4-8-flowered, ebracteate axillary raceme; rachis and
pedicels somewhat glandular-lepidote, glabrous to minutely subpuberulous at
the nodes. Flowers with inconspicuous musky odor, calyx cupular, coriaceous,
truncate to shallowly 5-lobed, -+- lepidote with plate-shaped glands in the uppe
part, 5-9 mm long and 5-9 mm wide; corolla uniformly bright yellow, tubular-
campanulate, 5.2-7.1 cm long and 1.2-2.2 cm wide at mouth of the tube 4.5-5.8
cm long, the lobes 1.1-1.3 cm long, the lobes glandular-lepidote outside and in,
the tube glabrous outside, inside sparsely glandular-lepidote near the mouth and
pubescent with simple, several-celled, mostly gland-tipped trichomes at the level
of stamen insertion; stamens didynamous, the anther thecae divaricate, 3-4.5 mm
long, the longer filaments 2.5-3.2 cm long, shorter filaments 1.9-2.3 cmi long, the
staminode 6-15 mm long, inserted 11-13 mm from base of corolla tube; pistil
4.2-4.6 cm long, the ovary flattened-cylindric, 6-7 mm long and 2.5 mm wide,
1.5-2.0 mm thick, lepidote, the ovules 2-seriate in each locule; disc annular-pulvi-
nate, 2-2.5 mm long and 6-7 mm wide. Capsule oblong-elliptic, flattened, 14-20
cm long and 7.2-8.3 cm wide, the basal stipe 1-2 cm long, drying tan, glabrous;
seeds thin, suborbicular, 4-5.5 cm long and 4-5.5 cm wide, the wing not sharply
demarcated, membranaceous, marginally hyaline, basally brown, surrounding the
seed body.
A plant of the tropical wet forest ranging from southern Mexico to the Pacific
coast of Colombia, this species apparently flowers mostly in the dry season. It is
rarely collected and has not previously been reported outside Colombia. It can
be recognized vegetatively by the large, varnished-looking leaflets, lack of foli-
aceous pseudostipules, and trifid tendrils.
CANAL ZONE: Pipeline Road, TTC Transect 3, Gentry 7395 (MO). COCLE: N of El Valle
de Anton, Allen 2831 (US). El Valle near La Mesa, Gentry 7417 (MO). COLON: Rio
Guanche, Gentry 6315 (MO). Santa Rita Ridge, Dwyer & Gentry 9374 (MO, SCZ). DARIEN:
Cerro Pirre, Gentry & Clewell 6964 (MO, PMA). PANAMA: Cerro Campana, Gentry 4907
(MO).
2. Anemopaegma chrysoleucum (H.B.K.) Sandw., Lilloa 3: 459. 1938.-FIG.

3C-F.
Bignonia chrysoleuca H.B.K., Nov. Gen. Sp. P1. 3: 134. 1819. TYPE: Colombia, Rio Magda
Anemopaegma belizeanum Blake, Contr. Gray Herb. 52: 91. 1917. TYPE: British Honduras,
Toledo, Peck 957 (GH).
A. punctulatum Pittier & Standl., Jour. Washington Acad. Sci. 15: 461. 1925. TYPE: Panama,
Colon, Pittier 4009 (GH, NY, US).
A. macrocarpa Standl., Publ. Field Columbian Mus., Bot. Ser. 4: 262. 1929. TYPE: Panama,
Bocas del Toro, Cooper 202 (F, NY, US).
Liana to at least 2.5 cm in diameter, usually less than 1 cm in diameter, the

bark smooth, finely longitudinally ridged; stem with 8 phloem arms in cross sec-
tion; branchlets subterete, longitudinally striate when dry, mostly glabrous, the
nodes puberulous, without interpetiolar glandular fields; pseudostipules foli-
aceous, 4 at each node, 3-13 mm long, 1.5-10.0 mm wide. Leaves 2-foliolate
(rarely 3-foliolate), sometimes cirrhose; leaflets narrowly elliptic, acute, cuneate

at base, 5.5-15 cm long and 2.0-7.1 cm wide, membranaceous, secondary veins 5-8
on a side, densely lepidote-papillose with scattered, often deciduous, lepidote
scales, puberulous on the midvein above, otherwise glabrous, drying olive, con-
colorous; tendril simple, 8-17 cm long; petiolules 0.3-2 cm long, petiole 0.7-3.4
cm long, puberulous on the upper surface. Inflorescence an axillary raceme, often
reduced to 3 flowers or a single flower in the leaf axil, the peduncle and pedicels
glabrous or inconspicuously lepidote. Flowers with a sweet aroma, calyx cupular,
truncate, 7-12 mm long and 7-10 mm wide, inconspicuously lepidote to essentially
glabrous, inconspicuously ciliate on the margin, usually with submarginal plate-
shaped glands, corolla lobes cream-colored, the tube yellow outside, inside yellow
with a longitudinal orange stripe below each of the lower 4 sinuses, tubular-
campanulate above a narrowed base, 6.2-10.3 cm long and 1.1-2.2 cm wide at
the mouth of tube, the tube 4.3-7.5 cm long, the lobes 1.1-2.0 cm long, glabrous
outside, inside glandular-lepidote on the lobes, mostly glabrous in the tube, pu-
bescent with simple trichomes at the level of stamen insertion; stamens didyn-
amous, the anther thecae divergent, 3.5-4.5 cm long, the longer filaments 2.3-3.1
cm long, the shorter filaments 1.6-2.1 cm long, the staminode 3-6 mm long; in-
serted (2.4-)2.7-3.1 mm from base of corolla tube; pistil 5.1-5.6 cm long, the
ovary rounded-cylindric, 2.0-2.5 mm long and 2.0-2.5 mm wide, 1.5-2.0 mm
thick, densely lepidote, the ovules 4-seriate in each locule; disc conical-pulvinate,
3-4 mm long, 3-4 mm wide at base, 1.5 mm wide at top. Capsule elliptic, com-
pressed, acute, 6.2-12.2 cm long and 2.9-6 cm wide, on a stipe 1-1.2 cm long,
drying yellow-brown, the surface slightly and inconspicuously lepidote, appearing
rough and varnished under a lens, the calyx persistent; seeds 1.4-1.8 cm long and
1.9-2.4 cm wide, woody, somewhat flattened, essentially wingless, the hilum large,
elliptic, to 3 mm by 7 mm.
This species is restricted to lowland swampy areas and river banks in the
tropical wet forest and wetter parts of the tropical moist forest, ranging from
British Honduras to Venezuela. It flowers irregularly throughout the year, most
frequently in the wet season, seldom with more than one or two flowers opening
at a time. The vine tends to remain slender and can flower when only a few
millimeters in diameter and almost herbaceous. The flowers are sweetly aromatic
when fresh with an odor resembling that of Cydista aequinoctialis but more deli-
cate. The seeds are water-dispersed.
Foliaceous pseudostipules provide the most useful field character. The slender,
terete twigs distinguish A. chrysoleucum from the angular twigs of other similarly
pseudostipulate species of Bignoniaceae.
Anemopaegma chrysoleucum was more broadly interpreted by Sandwith
(Kew Bull. 1953: 470. 1954) to include A. paraense Bur. & K. Schum. (including
A. surinamense Sprague ex Pulle and A. grandiflorum Sprague) of the Guianas
and Amazonia. A. chrysoleucum has small, membranaceous leaflets, small pseu-
dostipules, and few flowers in an inflorescence. Anemopaegma paraense has
larger, coriaceous leaflets, larger pseudostipules, and a many-flowered inflores-
cence. Field study shows that the distinguishing characters of A. chrysoleucum
are consistent and do not vary from top to bottom of a vine as supposed by Sand-
with. On the other hand, A. flavum Moro

livia and west central Brazil to Argentina is similar to A. chrysoleucum and may
be conspecific.
BOCAS DEL TORO: Almirante, Cooper 202 (F, NY, US); Gentry 2688 (MO). Nances Cay
Island, von Wedel 538 (MO, US). Water Valley, von Wedel 968 (GGH, MO, US). Fish Creek,
von Wedel 2218A, 2356 (both MO), 2445 (GH, MO, US). Shepherd Island, von Wedel 2674
(GGH, MO, US). Western River, von Wedel 2702 (GH, MO, US). Near Almirante, Woodson
& Schery 1031 (MO). CANAL ZONE: Barro Colorado Island, Bangham 544 (A); Croat 12683
(MO); Shattuck 1131 (MO); Starry 309 (MO). Near Gatuin, Goldman, 1869 (US). Cano
Quebrado, Pittier 6669 (US). S of Colon, Tyson et al. 4490 (MO, SCZ). COLON: E
of Rio Piedras, Gentry 2188 (MO). Along Rio Sirri, Trinidad Basin, Pittier 4009 (GH, NY,
US). DARIEN: Near El Real, Duke 4826, 4915 (both MO), 4936 (MO, SCZ). Between
Cerro Pirre and El Real, Gentry 4784 (MO). Near Marraganti, Williams 649 (NY, US).
PANAMA: Rio Sulugandi, Gentry 3745 (MO). Junction of Rios Espave and Bayano, Gentry
3848 (MO). SAN BLAS: Mainland opposite Playon Chico, Gentry 6382 (MO, PMA).
3. Anemopaegma orbiculatum (Jacq.) DC., Prodr. 9: 190. 1845.-FIG. 3A-B.
Bignonia orbiculata Jacq., Sel. Stirp. Amer. Hist. 184, tab. 180, fig. 79. 1763. TYPE: Colombia,
Cartagena, Jacquin s.n. ( BM, single leaf which matches illustration) .
Jacaranda orbiculata (Jacq.) Spreng., Syst. Veg. 2: 834. 1825.
Platolaria flavescens Raf., Sylva Tell. 78: 1838. TYPE: (not seen).
Pithecoctenium panamense Benth., Bot. Voy. Sulphur 129. 1844. TYPE: Panama, Sinclair
s.n. (K).
Peltospermum orbiculatum (Jacq.) DC., Prodr. 9: 191. 1845, pro. syn.
Liana to at least 2.5 cm in diameter, the bark light gray, smooth; stem with 8
phloem arms in cross section; branchlets subtetragonal, hollow, longitudinally
ridged, puberulous, interpetiolar glandular fields lacking, an interpetiolar ridge
sometimes connecting opposite petioles; pseudostipules usually lacking, rarely
minutely foliaceous in young branches. Leaves digitately 5-foliolate or 4-foliolate
with a tendril or tendril scar; leaflets ovate, acuminate, rounded to subcordate,
membranaceous to subcoriaceous, secondary veins 5-7(-8) on each side, minutely
papillate and impressed-lepidote above and beneath, scabrous along most veins
beneath and on the midvein- above, drying olive, the terminal leaflet 4.3-22 cm
long and 2.2-10 cm wide, intermediate leaflets 4.7-18.2 cm long and 2.3-9.5 cm
wide, basal leaflets 3.0-14.5 cm long and 2.1-8.2 cm wide, the terminal petiolule
0.8-5.5 cm long, intermediate petiolules 0.8-3.8 cm long, basal petiolules 0.4-2.6
cm long; petiole 3.2-13.5 cm long, puberulous; tendril simple or trifid, 8-17 cm
long, the ends when split, 1-5 mm long. Inflorescence a 1-10-flowered axillary
or terminal raceme, its branches puberulous. Flowers faintly fragrant, calyx cupu-
lar, truncate, with 5 irregular, minute, submarginal teeth, 5-8 mm long and 5-9
mm wide, puberulous, sometimes with glands; corolla tube yellow, the lobes
cream, without nectar guides, tubular-campanulate, 3.2-5.0 cm long and 1.4-1.9
cm wide at the mouth of tube, the tube 2.6-4.1 cm long, the lobes 0.7-1.0 cm long,
strongly glandular-lepidote outside, the trichomes sometimes drying orange, in-
side scattered glandular-lepidote, pubescent with gland-tipped trichomes at and
below the level of stamen insertion; stamens didynamous, the anther thecae more
or less divaricate, 3-4 mm long, the longer filaments 2.0-2.6 cm long, the shorter
filaments 1.3-1.9 cm long, the staminode 2-7 mm long, inserted 6-11 mm from
base of corolla tube; pistil 2.4-3.0 cm long, the ovary stipitate, rounded-cylindric,
2 mm long and 1.5 mm wide, 1 mm thick, lepidote, the ovules 4-seriate; disc coni-
cal-pulvinate, 3 mm wide for basal 1 mm, upper 1.5 mm tapering to a 1-mm-wide
stipe below ovary. Capsule circular with a 1.0-1.5 cm long basal stipe, 7.5-9.5 cm
in diameter, flattened, the valves slightly convex, ca. 2 cm thick at the center, the
surface smooth, scattered lepidote or lepidote-punctate; seeds circular, 2.8-4.5 cm
in diameter with a well demarcated membranaceous wing surrounding the seed
body, brownish near the center and hyaline near the margin.
Occurring most commonly in drier parts of the tropical moist forest, this
species is also found to some extent in tropical wet forest, mostly in disturbed
areas. It ranges from Nicaragua to Colombia, though previously unreported
northwest of the Canal Zone. It flowers irregularly throughout the year. The
seeds are effectively wind dispersed. This is the only liana of the family in Cen-
tral America with palmately compound 5-foliolate leaves. The 3-foliolate first
seedling leaves are unusual in the Bignonieae.
CANAL ZONE: Pipeline Road, Correa & Dressler 408 (MO); Croat 12742 (MO, SCZ);
Gentry 1775 (MO), 1799 (MO, SCZ), 4856 (MO). Ancon Hill, Duke 4603 (K, MO);
Killip 12110 (US); Mason s.n. (US); Standley 26399 (US); Williams 37 (NY, US). Near
Summit Hills Golf Course, Gentry 1587, 2820 (both MO). Between Corozal and Ancon,
Pittier 6507 (BM, F, GH, MO, NY, US); Pittier 6803 (GH, US). Balboa, Standley 29272
(US). Obispo, Standley 31774 (US). Near Miraflores, White 114 (F, GH, MO). DARIEN:
Isla Maria in Golfo de San Miguel, Gentry 3999 (MO). Near La Palma, Gentry 4296 (MO);
Pittier 5493 (NY, US). LOS SANTOS: Loma Prieta, Duke 11877 (MO); Lewis et al. 2247
(MO). PANAMA: Near Bejuco, Allen 974 (MO); Duke 4558 (K, MO). Nueva Gorgona,
Allen 3765 (GH, K, MO, NY). Perlas Islands, without locality, Anderson 1852 (GH). Be-
tween Rio Pacora and Chepo, Dwyer et al. 5092 (MO, NY, SCZ). Perlas Islands, San Jose
Island, Erlanson 14 (GH, US), 34 (GH, NY, US), 391 (GH, US); Harlow 40 (GH, US);
Johnston 198 (GH, MO, US), 469 (GH), 668 (GH, US), 714 (GH, MO). Near Rio Bayano
beyond Caniitas, Gentry 1664 (MO). Rio Piragua (or Tabardi), Gentry 2550 (MO). Icanti,
Gentry 2619 (MO). Near Rio Espave, Gentry 3701 (MO). Near Rio Parti, Gentry 3798
(MO). Near Panama Cruces, Hayes 132 (BM, K). Near Juan Diaz, Maxon & Harvey 6736
(BM, US); Standley 30612 (US). Near Bella Vista, Piper 5325 (US). Along Corozal road
near Panamai, Standley 26767 (US). WITHOUT LOCALITY: Cuming 1110 (BM, K); Hayes 594
(NY); Seemann s.n. (K); Sinclair s.n. (K).
4. Anemopaegma puberulum (Seib.) Miranda, Anal. Inst. Biol. Mexico 24:

93. 1953.
Chodanthus puberulus Seib., Carnegie Inst. Washington Publ. 527: 425. 1940. TYPE: Mexico,
Chiapas, Chiapa, Goldman 999 (US).
Liana, branchlets subterete, puberulous or glabrate, the nodes without int

petiolar glandular fields; pseudostipules small, inconspicuous, paired in eac
Leaves 2-foliolate with a tendril or tendril scar; leaflets ovate-elliptic, acute,
base rounded, 8-13 cm long and 5.6-7.5 cm wide, chartaceous, the secondary
veins 3-5 on each side; strongly puberulous beneath, glabrous and shiny above;
tendril trifid; petiolules terete, 2.5-3.0 cm long, petiole 2.2-3.1 cm long. Inflores-
cence a several-flowered ebracteate puberulous raceme. Flowers with the calyx
cupular, subcoriaceous, truncate, puberulous with glands in the upper part, 6-8
mm long and 8-9 mm wide; corolla yellow, tubular-campanulate, 5-5.4 cm long
and 1.5-1.6 cm wide at the mouth of tube, the tube 4-4.3 cm long, the lobes
0.8-1.0 cm long, glandular-lepidote inside and out, the tube glabrous except inside
at the level of stamen insertion; stamens didynamous, the anther thecae divaricate,
ca. 3 cm long, the longer filaments ca. 2 cm long, the shorter filaments ca. 1.3 cm
long, the staminode 3-4 mm long; inserted 8-9 mm from the base of the corolla
tube; pistil 3.2-3.3 cm long, ovary narrowly oblong, 3 mm long, 1-1.5 mm wide,
lepidote; disc annular-pulvinate, 1 mm long and 2-2.5 mm wide. Capsule (Mi-
randa) elliptic-oblong, flattened, puberulent and glandular-lepidote, 12-21 cm
long and 6.5-8, cm wide, on a stipe 1.5-2 cm long.
This species was supposedly restricted to Veracruz to Chiapas in southern

Mexico and to adjacent Guatemala, although a plant cultivated at Fairchild
Tropical Garden, Florida, was reported as acquired from Costa Rica. The single
Panamanian collection cited here represents a significant range extension and
elimination of most of the disjunction between A. puberulum and its South Ameri-
can relative, A. lehmanii Sandw. The latter species, known only from the type
collection from the Department of Cauca, Colombia, is closely related to and
perhaps not distinct from A. puberulum, reportedly differing in shorter trichomes,
shorter petioles, and larger calyx.
Anemopaegma puberulum apparently differs from A. chrysanthum Dugand
only in its pubescence, especially on the leaves, and the two may prove to be
conspecific.
PANAMA: El Llano-Carti highway, 6-10 km N of El Llano, Dressler 4329 (MO, PMA)
5. Anemopaegma santa-ritense A. Gentry, Ann. Missouri Bot. Gard. 58: 93.

1971. TYPE: Panama, Colon, Gentry 454 (MO, holotype; WIS, isotype).
Liana, branchlets more or less subtetragonal with the wrinkled surface form-
ing longitudinal ridges when dry, the epidermis finely and regularly papillate
with scattered lepidote scales and sometimes a minute puberulence at the nodes,
the nodes without interpetiolar glandular fields or pseudostipules. Leaves 2-foli-
olate with a tendril or tendril scar; leaflets ovate, obtuse, short acuminate, basally
truncate 5.9-11 cm long and 2.8-5.3 cm wide, coriaceous to chartaceous, the
margins revolute, the epidermis regularly and densely papillose with scattered
lepidote punctations, the smaller veinlets obscure, glabrous above and beneath to
rarely minutely puberulent at the base of the midvein beneath, green to olive
above, light green to yellowish beneath, the petiolules 0.6-1.3 cm long, the petiole
1.2-2.5 cm long; tendril usually trifid, rarely simple, 3-12 cm long to branching,
the 3 arms 0.4-1.0 cm long. Inflorescence an axillary one to few-flowered raceme
with opposite flowers. Flowers with the calyx cupular, glabrous, truncate, eglan-
dular, 5-8 cm long and 7-9 cm wide; corolla yellow, tubular-campanulate above
the narrowed tubular base, 5-7.5 cm long and 1.7-1.9 cm wide at the mouth of
tube, the tube 3.8-5.3 cm long, the lobes 0.8-1.1 cm long, lepidote outside, inside
lepidote on the lobes, glandular pubescent below the point of stamen insertion
and sparsely so on the lobes; stamens didynamous, the anther thecae divaricate,
3-4 mm long, the longer filaments 1.9-2.1 cm long, the shorter filaments 1.7-1.9
cm long, the staminode 0.7-0.9 cm long, inserted ca. 10 mm from base of corolla
tube; pistil 4.0-4.2 cm long, the ovary stipitate, flattened-ovate, 2 mm long, 1-1.5
1973] GENTRY FLORA OF PANAMA (Family 172. Bignoniaceae) 803
mm wide, 0.5 mm thick; disc conspicuous, conical-pulvinate, 1.5 mm long, 2.5 mm

wide. Capsule unknown.
This species is known only from the tropical wet forest in central and eastern
Panama, mostly above 800 m elevation. Recent sterile collections from Cerro Jefe,
Cerro Pirre, and the mountains above Puerto Obaldia are referable to this species,
formerly known only from Santa Rita Ridge. Its fruits have never been collected.
Anemopaegma santa-ritense is a puzzling form with close relatives from which
it seems to differ consistently, but by only one or two significant characters. Its
closest relatives are A. carrerense Armitage (possibly = A. karstenii Bur. & K.
Schum.) and A. chamberlaynii (Sims) Bur. & K. Schum., which have leafy pseu-
dostipules and minor leaf differences but are similar in other respects.
COLON: Santa Rita Ridge, Allen 5105 (MO); Correa & Dressler 772 (MO); Croat 13832
(MO); Dressler & Lewis 3721 (MO); Gentry 454 (MO, WIS), 734, 735, 736 (411 MO);
Gomez-Pompa et al. 2990, 3305 (both MO); Lewis et al. 5288 (MO, SCZ). DARIEN: Summit
of Cerro Pirre, Gentry 4599 (MO). PANAMA: Cerro Jefe, Gentry 2879 (MO). SAN BLAS:
Mountaintop above Puerto Obaldia, Gentry 1506 (MO).
4. ARRABIDAEA
Arrabidaea DC., Rev. Bign. (Bibl. Univ. Geneve) 10. 1838. TYPE: A. rego
(Vell.) DC.
Chasmia Schott ex Spreng., Syst. Cur. Post. 409. 1827, nomen nudum.
Pentelesia Raf., Sylv. Tell. 78. 1838. TYPE: P. discolor Raf. = Arrabidaea carichanensis
(H.B.K.) Bur. & K. Schum.
Vasconcellia Mart., Flora 24(2): Beibl. 12. 1841, nomen nudum.
Cremastus Miers, Proc. Roy. Hort. Soc. 3: 187. 1863. LECTOTYPE: C. glutinosa (DC.) Miers
= Arrabidaea sceptrum (Cham.) Sandw.
Panterpa Miers, Proc. Roy. Hort. Soc. 3: 196. 1863. TYPE: P. leucopogon (Cham.) Miers
Arrabidaea leucopogon (Cham.) Sandw.
Petastoma Miers, Proc. Roy. Hort. Soc. 3: 194. 1863. TYPE: P. samydoides (Cham.) Miers -
Arrabidaea samydoides (Cham.) Sandw.
Paramansoa Baill., Hist. P1. 10: 27. 1888. TYPE: P. grosourdyana Baill. = Arrabidaea
grosourdyana (Baill.) Sandw.
Lianas, rarely shrubs or small trees, stems with 4 phloem arms in cross section;
branchlets terete, usually with interpetiolar glandular fields; pseudostipules small
and inconspicuous. Leaves usually 3-foliolate, the terminal leaflet often replaced
by a simple tendril. Inflorescence usually a large, many-flowered axillary or
terminal panicle. Flowers with the calyx cupular, truncate, usually minutely
5-denticulate, pubescent; corolla pale pink to purple (white-flowered forms in
some species), relatively small, more or less campanulate, densely pubescent
outside at least on the lobes; anthers glabrous, the thecae straight or slightly
curved, usually divaricate; pollen grains single, tricolpate, the exine finely granu-
lar or punctate; ovary narrowly cylindrical, usually minutely lepidote, the ovules
2-seriate; disc cupular-pulvinate. Fruit a compressed, linear capsule, the valves
parallel to the septum, essentially smooth, usually with a slightly raised median
nerve; seeds bialate, the wings usually more or less hyaline and distinct from the
seed body.
About 70 species ranging in tropical America from Mexico and the West
Indies (rare) to Argentina.
a. Capsule valves without noticeably raised median nerve, softly puberulous or with
sharply angled margins; corolla usually over 3.5 cm long, the tube pubescent outside;
calyx usually drying dark with lighter or thinner rim.
b. Capsule valves pubescent, the margins not angled; calyx evenly 5-denticulate with
a white-fringed margin ------- ---------- 6. A. mollissima
bb. Capsule valves glandular-pitted; the margins sharply angled; calyx unevenly truncate
to sub-bilabiate, not evenly denticulate, thinner at the margin but not evenly whitish-
fringed - 4. A. corallina
aa. Capsule valves with somewhat raised median nerve, glabrous or lepidote, the margins
rounded; corolla usually less than 4 cm long (if longer, the tube glabrous); calyx rim
not noticeably lighter or thinner than rest of calyx when dried.
c. Corolla tube glabrous outside, dark in bud with a conspicuous white tip from the
pubescence of the lobes; calyx reflexed away from base of corolla; leaves simple or
2-foliolate -------------------------------------------- 8. A. patellifera
cc. Corolla tube pubescent outside, the buds not bi-colored; calyx cupular, enclosing
base of corolla; leaves always 3-foliolate in part, never simple.
d. Leaflet underside conspicuously whitish from the dense short puberutence, the
main veins less puberulous and drying brownish or tan and conspicuously darker
than the lamina -1. A. candicans
dd. Leaflets variously pubescent or glabrate but undersides not conspicuously whitish
in contrast with the darker main veins.
e. Leaflets (including veins) evenly and densely tannish-puberulous beneath;
corolla less than 1.8 cm long; capsule usually less than 9 cm long-
------------------ --------------- 9. A. pubescens
ee. Leaflets glabrate or pubescent mostly along the main nerves beneath (if in
part tannish-puberulous, then the corolla more than 2.2 cm long); mature
capsules usually more than 9 cm long.
f. Leaves drying conspicuously brick red at least in patches, usually glabrous
except for simple trichomes in axils of lateral nerves beneath; interpetiolar
glandular fields present -2. A. chica
ff. Leaves not drying reddish, usually slightly puberulous at least along main
veins below; interpetiolar glandular fields present or absent.
g. Mature branchlets with loose, conspicuously wrinkled epidermis and
inconspicuous lenticels, young branchlets drying black with contrast-
ing light lenticels; inflorescence contracted, few-flowered (3-7(-25)-
flowered including buds) -7. A. oxycarpa
gg. Mature branchlets with tight, finely, and evenly striate epidermis with
noticeable small lenticels; young branchlets drying grayish or brown-
ish; inflorescence open, many-flowered.
h. Corolla less than 2 cm long; calyx drying grayish, eglandular,
small (3-4 mm long and 2-3 mm wide); inflorescence branches
long-pedunculate (1-5 cm), the peduncles terminated by dense
ultimately trichotomous corymbs; capsule drying blackish, 0.8-1.0
cm wide; seed wings hyaline-membranaceous; usually without in-
terpetiolar glandular fields- - . A. florida
hh. Corolla more than 2.2 cm long; calyx drying whitish-gray with
conspicuous submarginal darker glands, larger (4-7 mm long and
4-6 mm wide); lateral branches of inflorescence narrow, sub-
racemose, without elongate peduncles, the flowers borne along
their length; capsule drying brownish, more than 1.0 cm wide at
maturity; seed wings basally brownish with hyaline tips; usually
with interpetiolar glandular fields -3. A. conjugata
1. Arrabidaea candicans (L. C. Rich.) DC., Prodr. 9: 185. 1845.-FIG. 4D-H.

Bignonia candicans L. C. Rich., Act. Soc. Hist. Nat. Paris 1: 110. 1792. TYPE: French Guiana,
LeBlond s.n. (P).
B. colorata Poir. in Lam., Encycl. Bot. Suppl. 1: 634. 1800. TYPE: French Guiana, ex herb.
Poiret (P).
Arrabidaea candicans var. latifolia DC., Prodr. 9: 186. 1845. TYPE: French Guiana, Perrottet
s.n. (BM, P).
FIGURE, 4. Arrabidaea.-A-C. A. florida DC.-A. Habit (X 3/5).-B

( X 3/5).-C. Ovary cross section ( X 12). [After Gentry 3120 (M
(L. Rich.) DC.-D. Fruit ( X 1/2). [After Croat 5228 (MO).]-E. O
-F. Flower ( X 3/5).-G. Ovary cross section ( X 12).-H. Corolla split
Gentry 2865 ( MO ).]
A. orbignyana DC., Prodr. 9: 184. 1845. TYPE: Bolivia, Santa Cruz, d'Orbigny 1135 (P).
A. subincana DC. sec. Seem., Bot. Voy. Herald 180. 1854, non DC.
A. pruinosa K1. ex Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 57. 1896. TYPE: British Guiana,
Schomburgk 433 (not seen).
A. schomburgkii KI. ex Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 58. 1896. TYPE: British
Guiana, Schomburgk 1332 (K).
A. pachycalyx Sprague, Bull. Herb. Boiss., ser. 2. 6: 373. 1906. LECTOTYPE: Colombia, Mag-
dalena, H. H. Smith 744 (BM, BR, GH, K, MO, NY, P, US).
A. rhodothyrsus Kranzl., Fedde Repert. 17: 20. 1921. TYPE: Costa Rica, Guanacaste, Tonduz
13643 (BM, F, GH, K, NY, US).
A. calodictyos Sprague, Kew Bull. 1927: 358. 1927. TYPE: Venezuela, Distrito Federal,
Pittier 11883 (K).
A. zulianensis Pittier, Jour. Washington Acad. Sci. 18: 341. 1928. TYPE: Venezuela, Zulia,
Pittier 10992 (US, VEN).
Liana to 7 cm in diameter, the bark yellowish to pale gray, smooth, 4-furrowed

in age; stem with 4 phloem arms in cross section; branchlets subterete with raised
warty lenticels, minutely puberulous when young, soon glabrate, interpetiolar
glandular fields conspicuous; pseudostipules lacking. Leaves 3-foliolate or 2-
foliolate with a tendril or tendril scar; leaflets ? ovate, membranaceous to char
taceous, secondary veins (4-)5-6(-9) on a side, above glabrate, minutely pu-
berulous at the base of the midvein, drying grayish-brown to reddish-brown,
beneath densely white puberulous, less so on the main veins, the short simple
trichomes completely covering the surface, drying whitish from the pubescence
with contrasting darker main veins, the terminal leaflet acuminate to subretuse,
broadly cuneate to truncate, 6.2-15.5 cm long and 4.2-9.1 cm wide; the lateral
leaflets acute to retuse, broadly cuneate to asymmetrically subcordate, 5.8-14.8
cm long and 3.6-10.7 cm wide; the terminal petiolule 1.0-3.3 cm long, the lateral
petiolules 0.8-3.2 cm long, the petiole 1.6-6.1 cm long, minutely puberulous as
the petiolules; tendril simple, 11-21 cm long. Inflorescence a terminal or axillary
panicle, its branches puberulous. Flowers with a sweet aroma, calyx cupular,
simple puberulous and lepidote, mostly with some glands, truncate to minutely
subdenticulate, 3-6 mm long and 2-4 mm wide; corolla magenta with white throat,
tubular-campanulate, 2.2-4.0 cm long and 0.8-1.4 cm wide at the mouth of tube,
the tube 1.4-2.7 cm long, the lobes 0.4-1.0 cm long, moniliform pubescent outside
and on lobes inside, the tube mostly glabrous, sparsely pubescent in the floor of
throat with several-celled simple or rarely forked trichomes, and at the level of
stamen insertion with long simple gland-tipped trichomes; stamens didynamous,
inserted 5-6 mm from base of corolla tube, the anther thecae divaricate, 2-2.5 mm
long, the longer filaments 1.5-1.6 cm long, the shorter filaments 1.1-1.4 cm long,
the staminode 4-6 mm long; pistil 1.7-1.8 cm long, the ovary linear oblong, 2-2.5
mm long and 0.5-1.0 mm wide, finely lepidote, the ovules 2-seriate; disc cupular-
pulvinate, ca. 1 mm long and 1 mm wide. Capsule linear, acuminate, flattened,
12-35 cm long and 0.8-1.3 cm wide, the surface slightly warty and with lepidote
glands, the midrib and margins slightly raised; seeds 0.6-1.0 cm long and 2.0-3.4
cm wide, the wings hyaline-membranaceous, sharply demarcated from the seed
body.
A common liana typical of the tropical moist forest, A. candicans occurs also
sporadically in the tropical wet forest and pre-montane wet and moist forest. It
ranges from southern Mexico to Brazil. This species flowers during the early dry
season from November through February. The fruits mature and seeds are re-
leased in the late dry season.
The whitish pubescence of the leaf undersurface of this species is its best
recognition mark. When dry the main veins are darker than the more pubescent
ground surface in contrast to A. pubescens, which has veins and undersurface
equally pubescent, with a more tannish color. On young leaves of A. candicans
whitish pubescence may not be obvious except under a lens.
Arrabidaea calodictyos is apparently a partially glabrescent form of this
species. It is intermediate between A. candicans and A. chica and could be a
result of hybridization between the two species. Sandwith identified many of
these intermediates with A. chica rather than A. candicans.
CANAL ZONE: Barro Colorado Island, Brown 123, 128 (both F); Croat 4833 (SCZ), 5228
(MO, SCZ), 7201 (MO, NY, SCZ), 7329 (MO), 7347, 7698A (both MO, SCZ), 7884 (MO),
7902, 8124 (both MO, SCZ), 8312 (MO, NY, SCZ), 8375 (MO), 11099 (MO, SCZ), 12592,
12631 (both MO), 12705 (MO, NY); Dwyer et al. 8441, 8457 (both MO); Foster 681, 1416
(both PMA); Gentry 435 (MO, WIS); Graham 200 (GH); Oppenheimer & Hladik 97 (MO);
Shattuck 499 (F, MO, US); Standley 41071 (US); Wetmore & Abbe 2 (F), 13 (A, F, MO),
27 (F), 58 (A, MO); Wetmore & Woodworth 27, 848 (both A). Along road to Venado
Beach, Gentry 2865 (MO). Road to Gamboa airfield, Gentry 3267 (MO). Rio Grande Station,
Panamat Railroad, Hayes s.n. (BM). Near the Mindi, Hayes 589 (NY). Howard Air Force
Base, Nowicke et al. 3608A (MO, SCZ). Along Las Cruces Trail between Fort Clayton and
Corozal, Standley 29226 (US). CHIRIQUI: Near San Felix, Pittier 5129 (NY, US). COCLE:
2 miles E of Rio Hondo, Gentry 2907 (MO). Between Aguadulce and Chico River, Pittier
5003 (NY, US). 1-5 miles S of Anton, Tyson & Blum 2575, 2776 (both MO, SCZ). COLON:
1.5 miles E of Rio Piedras on road to Portobelo, Gentry 2191 (MO). Rio Agua Sucio, Gentry
3195 (MO). Between Rio Piedras and Puerto Pilon, Lewis et al. 3220 (MO, SCZ). DARIEN:
Isla Encanto in Golfo de San Miguel, Gentry 4013 (MO). S of Jaque, Gentry 4120, 4127
(both MO). Rio Jaque 1-2 hours upriver from Jaque, Gentry 4186 (MO). Upriver from
Boca de Cupe on Rio Tuira, Gentry 4321 (MO). El Real, Gentry 4565 (MO). HERRERA.
Vicinity of Oci, Allen 4070 (K, MO). Road from La Arena to outskirts of Pese, Burch et al.
1284 (GH, US). S of Ocui, Gentry 3125 (MO). E of Las Minas, Gentry 3127 (MO).
PANAMA: Vicinity of Pacora, Allen 1113 (GH, MO, NY, US). Rio Las Lajas, Allen 2038
(F, GH, MO, NY, US). Along road between Panama and Chepo, Dodge et al. 16625 (BR,
K, MO, U). Trail S from Pacora along Rio Pacora, Gentry 1410, 1411 (both MO). Rio
Pasiga, Gentry 2199, 2247, 2260 (all MO). Rio Maestro, Gentry 2235 (MO). Rio Piragua
(Tabardi), Gentry 2576 (MO). 10-15 km from Rio Bayano crossing on trail to Santa Fe,
Gentry 3821 (MO). W of Chepo, Gentry & Tyson 1640 (MO). Dam site on Rio Bayano,
Gentry & Tyson 1663B (MO). San Jose Island, Johnston 969 (BM, GH, MO, US), 1324,
1407 (both GH, MO, US). San Francisco de la Caleta, Paul 237 (US). Near Panamat,
Sargent s.n. (US). Near Punta Paitilla, Standley 26310 (MO, US). E of Rio Tocumen,
Standley 26691 (BM, GH, US). Tumba Muerto Road near Panama, Standley 29729 (GH,
US). VERAGUAS: Hills W of Sona, Allen 1029 (GH, MO, NY, US). 25 mi. W of Aguadulce,
Gentry 2922 (MO). Near Santiago, Seemann 152 (K). WITHOUT LOCALITY: Duchassaing
s.n. (K, P), Seemann 152 (K).
2. Arrabidaea chica (H. & B.) Verl., Rev. Hortic. 40: 154. 1868.
Bignonia chica H. & B., PI. Aequin. 1: 107, pl. 31. 1808. TYPE: Venezuela, Orinoco, Humboldt
& Bonpland s.n. (P).
B. cuprea Cham., Linnaea 7: 655. 1832. TYPE: Brazil, Sellow s.n. (K).
Arrabidaea acutifolia DC., Prodr. 9: 185. 1845. TYPE: Brazil, Sdo Paulo, Martius s.n. (M).
A. rosea DC., Prodr. 9: 185. 1845. SYNTYPES: Brazil, Sdo Paulo, Guillemin s.n. (not seen);
Amazonas, Martius s.n. (M).
Bignonia triphylla Willd. sec. DC., Prodr. 9: 151. 1845, pro syn, non L. nec Vell.
Temnocydia carajura Mart. ex DC., Prodr. 9: 185. 1845, pro syn.
Vasconcellia acutifolia Mart. ex DC., Prodr. 9: 185. 1845, pro syn.
Lundia chica (H. & B.) Seem., Bot. Voy. Herald 180. 1854.
Adenocalymma portoricensis Stahl, Est. 6: 186. 1888. TYPE: Puerto Rico (not seen, fide
Britton, Sci. Surv. Puerto Rico & Virgin Islands 6. 1925).
Bignonia erubescens S. Moore, Trans. Linn. Soc., ser. 2. 4: 412. 1895, non DC.
Arrabidaea larensis Pittier, Jour. Washington Acad. Sci. 18: 336. 1928. TYPE: Venezuela,
Lara, Jahn 1186 (GH, US, VEN).
A. cuprea (Cham.) Born., Revista Sudamer. Bot. 2: 10. 1935, non Pittier.
Liana to 6 cm in diameter, the bark pale gray, smooth, 4-furrowed in age;

stem with 4 phloem arms in cross section, the pith reddish, even when fresh;
twigs subterete with raised warty lenticels, glabrous to subpuberulous, the
younger branchlets drying reddish black, interpetiolar glands usually present but
often inconspicuous except on very young twigs; pseudostipules not noticeable.
Leaves 3-foliolate or 2-foliolate with a tendril or tendril scar; leaflets ovate to
narrowly ovate, acuminate to acute, cuneate to truncate, membranaceous, sec-
ondary veins 5-7 on a side, above glabrous, sometimes minutely scabrous at the
base of midvein, glabrous beneath except for simple trichomes in the axils of
lateral nerves or sometimes slightly scabrous along main veins, rarely puberulous
on the surface, dark glossy green when fresh, green with red mottling to uni-
formly brick red when dry; the terminal leaflet 4.4-12 cm long and 1.3-7.0 cm
wide; lateral leaflets 3.3-11.5 cm long and 1.3-5.5 cm wide, petiole and petiolules
glabrous to subpuberulous, the terminal petiolule 0.6-4.6 cm long, the laterals
0.3-2.1 cm long, the petiole 1.5-7.1 cm long; tendril simple, 6-12 cm long. In-
florescence a terminal panicle, its branches puberulous. Flowers aromatic, calyx
simple-puberulous, mostly eglandular, truncate to minutely subdenticulate, 3-5
mm long and 2-3 mm wide; corolla 1.6-3 cm long and 0.5-1.4 cm wide at the
mouth of tube, the tube 1.1-2.1 cm long, the lobes 0.6-1.2 cm long, pubescent, the
tube pubescent outside, mostly glabrous inside, pubescent at the level of stamen
insertion; stamens didynamous, the anther thecae divaricate, 1.5-2 mm long, the
longer filaments 1.1-1.3 cm long, shorter filaments 0.9-1.0 cm long, the staminode
3-4 mm long, inserted 3-4 mm from base of corolla tube; pistil 1.1-1.3 cm long,
the ovary narrowly cylindric, finely lepidote, 2-2.5 mm long and 0.5-1.0 mm wide,
the ovules 2-seriate; disc cupular, 1 mm long, 1 mm wide. Capsule linear, acute,
flattened, 12-23 cm long and 0.9-1.2 cm wide, the surface smooth, slightly irregu-
lar-ridged longitudinally, midrib and margins slightly raised, drying brown to
reddish brown, lepidote; seeds thin, bialate, 0.7-0.9 cm long and 2.3-2.6 cm wide,
the wings hyaline-membranaceous, distinct from the often reddish-drying body.
This vine is most abundant in tropical wet forest and premontane wet forest
but also occurs sporadically in wetter parts of the tropical moist forest. It ranges
from Mexico to Brazil and Argentina. It flowers at the end of the dry season and
through most of the wet season from mid-April to October.
The leaves of the plant are widely used among tropical Indians as a cosmetic
to color the skin red. A note on Duke 14406 reports additional Cuna uses to
include protection against sunburn and a medication used between the toes
against "mazamora."
The bright red color (sometimes only in patches) of the dry leaves is an ob-
vious herbarium distinguishing character. When fresh, the plant is relatively
difficult to recognize but the combination of mostly 3-foliolate leaves, narrow
dark green leaflets, and interpetiolar glandular fields makes recognition possible.
This is another of the species of Bignoniaceae with taxonomically confusing

pubescence forms. The leaves range from essentially glabrous to bearded in the
axils beneath (commonest) to puberulous over the lower leaf surface. The latter
form intergrades with A. candicans.
BOCAS DEL TORO: Water Valley, von Wedel 160 (GH, MO), 899 (GH, MO, US).
Shepherd Island, von Wedel 2676A, 2697 (both GH, MO, US). CANAL ZONE: Barro Colorado
Island, Croat 5038, 6258 (both MO, NY, SCZ), 9266 (MO), 11791 (MO, SCZ); Foster 931
(PMA). CHIRIQUI: Remedios, Seemann 1610 (GH). COCLE': Distrito de la Pintada, Escobar
s.n. (MO, PMA). DARIEN: Outskirts of La Palma, Gentry 3932 (MO). S of Jaque, Gentry
4138 (MO). Summit of Cerro Pirre, Gentry 4675 (MO). Manene to the mouth of Rio Cuasi,
Kirkbride & Bristan 1471 (MO). PANAMA: La Victoria hills E of Panama City, Allen 4454
(GH, MO, P). Piria, Duke 14406 (SCZ). Near Rio Espave, Gentry 3716 (MO). Between
Rios Espave and Sulugandi, Gentry 4987 (MO). San Jose Island, Harlow 28 (GH, US);
Johnston 14 (BM, GH, MO, US). SAN BLAS: Near stream above Puerto Obaldlia, Gentry
1535 (MO). VERAGUAS: Mountains N of Santa Fe, Gentry 2995, 3086A (both MO).
3. Arrabidaea conjugata (Vell.) Mart., Flora 24(2): Beibl. 46. 1841.
Bianonia conjugata Vell., Fl. Flum. 245. 1825; 6: Tab. 18. 1827. TYPE: Brazil (not seen).
B. farinosa Salzm. ex DC., Prodr. 9: 185. 1845, pro syn.
Arrabidaea virescens DC., Prodr. 9: 1184. 1845. TYPE: Brazil, Bahia, Blanchet s.n. (G-DC).
A. virescens var. farinosa DC., Prodr. 9: 185. 1845. TYPE: Brazil, Bahia, Salzmann 395
(G-DC).
Adenocalymma paniculatum Benth. ex Miers, Ann. Mag. Nat. Hist., ser. 3. 7: 393. 1861.
TYPE: Brazil, Para, Spruce s.n. (NY, W).
Bignonia ciliata Wawra, Oester. Bot. Zeitschr. 12: 237. 1862. TYPE: Brazil (not seen, fide
Bureau & K. Schumann, 1896-97).
Arrabidaea amazonica K. Schum. in Engler & Prantl, Nat. Pflanzenf. 4(3b): 213. 1894,
nomen subnudum. TYPE: Brazil (not seen, fide Bureau & K. Schumann, 1896-97).
A. chironeura Bur., Bull. Soc. Bot. France 58, Mem. 3f: 517. 1911. TYPE: Brazil, Rio de
Janeiro, Glaziou 4707 (P).
Liana; stem with 4 phloem arms in cross section; branchlets terete, puberulous
or glabrate, usually with conspicuous interpetiolar glandular fields, these some-
times reduced or almost lacking; pseudostipules short, obtuse, not conspicuous.
Leaves 3-foliolate or 2-foliolate with a tendril or tendril scar; leaflets elliptic or
ovate-elliptic, acute to acuminate, basally rounded,. 9-21 cm long and 4.5-12.2 cm
wide, membranaceous to chartaceous, secondary veins 4-6 on a side, underside ?
glabrous to densely puberulous, the main veins puberulous, above glabrous or
with a few short trichomes above the main veins, scattered lepidote on both sides
with some plate-shaped glands scattered along midvein beneath, drying dark olive
above and green or greenish olive beneath; petiolules 0.7-4.2 cm long, petiole
3-13 cm long, puberulous to glabrate. Inflorescence a terminal panicle of numer-
ous narrow subsidiary panicles, the branches puberulous. Flowers fragrant, calyx
cupular, truncate, minutely 5-denticulate, 4-7 mm long and 4-6 mm wide, drying
whitish gray, puberulous outside, usually with a subapical row of dark drying
glands; corolla magenta, white inside the tube, tubular-campanulate, 2.2-4.0 cm
long and 0.6-1.1 cm wide at the mouth, the tube 1.6-2.9 cm long, pubescent out-
side mostly glabrous inside, somewhat pubescent at the point of stamen insertion,
the lobes 3-8 mm long, pubescent; stamens didynamous, the anther thecae
straight, slender, divaricate, 2-2.5 mm long, the longer filaments 1.5-1.7 cm long,
the shorter filaments 1.0-1.2 cm long, the staminode 6-7 mm long, inserted 5-6
mm from the base of the corolla tube; pistil 1.7-1.8 cm long, the ovary cylindrical,
lepidote, 2 mm long and 1 mm wide, the ovules 2-seriate in each locule; disc
cupular-pulvinate, 0.5-1 mm long and 1.5-2 mm wide. Capsule linear, com-
pressed, acuminate, the median nerve and margins slightly raised, 12-31 cm long
and 1.1-1.4 cm wide; seeds thin, bialate, 0.7-1.1 cm long and 1.9-2.7 cm wide,
the wings brown except for a terminal hyaline-membranaceous fringe.
A liana of sporadic occurrence in the tropical moist forest and gallery forests
of the premontane moist and tropical dry forest, it ranges from Costa Rica to
Brazil. This species flowers in the wet season from July to September. The seeds
are released in the dry season.
Like most species of Arrabidaea, A. conjugata is difficult to distinguish vege-
tatively. The dark green, ovate, glabrate leaves provide the best character, but
it can be differentiated in the field only with experience. The inflorescence with
its narrow lateral branches and whitish calyces, usually with dark submarginal
glands, is quite distinctive.
I have seen no type material and have accepted the interpretation of Martius
(ex herb.) as to the application of this Vellozo name. The application of this
name to this species has also been accepted by many other authors. Should the
name Bignonia conjugata prove referable elsewhere, Arrabidaea virescens DC.
would be the correct name for this species.
CANAL zoNE: Near Juan Mina, Flat Rock, Chagres River, Bartlett & Lasser 16837 (MO).
Mohinga Swamp, Bartlett & Lasser 16825 (MO). West arm of Quebrada Salamanca, Dodge
et al. 17004 (BM, BR, MO, U). DARIEN: Rio Sabana above Santa Fe, Duke 14093 (MO).
Santa Fe, Gentry 4077 (MO). PANAMA: Rio Bayana, altos de Maje, Isla Corgas, Galindo
B-2 (MO).
4. Arrabidaea corallina (Jacq.) Sandw., Kew Bull. 1953: 460. 1954.
Bignonia dichotoma Jacq., Enum. PI. Carib. 25. 1760; Select. Stirp. Amer. Hist. 183. 1763,
non Arrabidaea dichotoma (Vell.) Bur. TYPE: (not seen, possibly represented by one
of the 2 sheets of B. corallina at W).
B. corallina Jacq., Fragm. Bot. 37, tab. 42, fig. 1. 1800-1809. TYPE: Venezuela, Caracas,
Jacquins.n. (W).
B. glabrata H.B.K., Nov. Cen. Sp. PI. 3: 137. 1819. TYPE: Venezuela, Cumana, Humboldt
& Bonpland 53 (P).
B. obliqua H.B.K., Nov. Cen. Sp. PI. 3: 135. 1819. TYPE: Venezuela, Laguna de Valencia,
B. villosa Vahl sec. Spreng., Syst. Veg. 2: 830. 1825, non Vahl.
B. hibiscifolia Cham., Linnaea 7: 705. 1832. TYPE: Brazil, Sellow s.n. (not seen, fide Sandw.).
Arrabidaea obovata DC., Prodr. 9: 185. 1845. TYPE: Brazil, Bahia, Martius s.n. (M).
Bignonia balbisiana DC., Prodr. 9: 153. 1845. TYPE: Colombia, Magdalena, Bertero s.n.
(C-DC, MO).
B. rotundata DC., Prodr. 9: 163. 1845. TYPE: Brazil, Bahia, Blanchet 2757 (K, MO, NY, P).
Cremastus balbisianus (DC.) Miers, Proc. Roy. Hort. Soc. 3: 187. 1863.
Arrabidaea obliqua (H.B.K.) Bur., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenh
1893: 99'. 1893.
Bignonia colombiana Morong in Morong & Britton, Ann. New York Acad. Sci. 7: 186. 1893.
TYPE: Paraguay, Pilcomayo River, Morong 1535 (MO, NY).
Arrabidaea rotundata (DC.) Bur. ex Schum. in Engler & Prantl, Nat. Pflanzenf. 4(3b): 213.
1894.
Cuspidaria hibiscifolia (Cham.) Bur. ex K. Schum. in Engler & Prantl, Nat. Pflanzenf. 4(3b):
216. 1894.
Tabebuia chapadensis S. Moore, Trans. Linn. Soc., ser. 2. 4: 422. 1895. TYPE: Brazil, Moore
79 (BM).
Arrabidaea rhodantha Bur. & K. Schum. in Mart., Fl. Bras. 8 (2): 44. 1896. SYNTYPES:
Paraguay, along Rio Paraguay, Balansa 499 (K); near Concepcion, Kuntze s.n. (not
seen); Brazil, Matto Grosso, Kuntze s.n. (not seen).
A. praecox Hassl., Bull. Herb. Boiss., ser. 2. 6 (append. 1): 25. 1898. TYPE: Paraguay,
Hassler 840 (K).
Bignonia boliviana Rusby, Bull. Torrey Bot. Club 27: 70. 1900. TYPE: Bolivia, junction of
Beni and Madre de Dios Rivers, Rusby 1142 (CIH, K, NY, US).
B. acuminata Johnston, Proc. Amer. Acad. Arts 40: 696. 1905. TYPE: Venezuela, Margarita
Island, Johnston 345 (F, GiH, K, M, NY, US).
Arrabidaea acuminata (Johnston) Urb., Fedde Repert. 14: 306. 1916.
Adenocalymma friesiana Kranzl., Fedde Rep. 17: 115. 1921. SYNTYPES: Argentina, Fries 446B
(S, fide Fabris, Rev. Mus. La Plata 9: 383. 1965); Lorentz & Hieronimus 435 (CORD,
fide Fabris Rev. Mus. LaPlata 9: 383. 1965.).
Arrabidaea barquisimetensis Pittier, Jour. Washington Acad. Sci. 18: 339. 1928. TYPE:
Venezuela, Lara, Saer d'Heguert 214 (K, VEN).
A. guaricensis Pittier, Jour. Washington Acad. Sci. 18: 337. 1928, pro parte. TYPE: Venezuela,
Guarico, Pittier 11308 (leaves only) (GH, US, VEN).
A. lenticellosa Pittier, Jour. Washington Acad. Sci. 18: 340. 1928, non Bur. & K. Schum.
SYNTYPES: Venezuela, Guarico, Pittier 12481 (GH, K, US, VEN), 12370 (CH, VEN).
A. ovalifolia Pittier, Jour. Washington Acad. Sci. 18: 337. 1928. TYPE: Venezuela, Aragua,
Pittier 12311 (C, CIH, K, US, VEN).
A. spraguei Pittier, Jour. Washington Acad. Sci. 18: 339. 1928. TYPE: Venezuela, Aragua,
Pittier 11476 (K, US, VEN).
Piriadacus hibiscifolius (Cham.) Pichon, Bull. Soc. Bot. France 92: 226. 1945.
Arrabidaea obliqua var. hirsuta (DC.) Dugand, Mutisia 9: 3. 1952.
A. rhodantha var. oxyphylla Sprague ex Sandw., Kew Bull. 1953: 460. 1954, pro syn.
Liana to 7 cm in diameter, the bark gray, rather smooth; stem with 4 phloem
arms in cross section; branchlets terete with raised lenticels, glabrous or glabrate,
nodes with or without interpetiolar glandular fields; pseudostipules short and
inconspicuous. Leaves 3-foliolate or 2-foliolate with a tendril or tendril scar, some-
times simple on young plants; leaflets ovate, acute to obtuse, truncate to subcor-
date or broadly cuneate at base, 3.1-18.2 cm long and 2.3-11.6 cm wide, mem-
branaceous to subcoriaceous, secondary nerves 4-7 on a side, variably pubescent,
from glabrate to strongly tannish-pubescent, especially in nerve axils beneath,
drying olive to gray, the pubescent form olive to tannish; tendril simple, 8-15 cm
long; petiolules 0.6-4.0 cm long, petiole 2.3-3.6 cm long, puberulous. Inflorescence
a series of narrow axillary panicles in the axils of fallen leaves, the branches
puberulous; buds minutely 5-parted with an acuminate tip. Flowers with the
calyx cupular, bilabiately subtruncate or with 5 inconspicuous teeth, 5-8 mm long
and 5-8 mm wide, minutely lepidote and somewhat scabrous, the lobes often with
tufts of pubescence; corolla lavender to magenta, the tube white within, tubular
campanulate, 2.7-4.6 cm long and 0.8-1.2 cm wide at the mouth of tube, the tube
2.1-2.8 cm long, the lobes 0.5-1.0 cm long, puberulous outside, the lobes and
abaxial surface of tube glandular-lepidote and puberulous inside, glandular-pu-
bescent at the level of stamen insertion; stamens didynamous, the anther thecae
divaricate, 2.5-3 mm long, the longer filaments 1.4-1.7 cm long, shorter filaments
0.9-1.1 cm long, the staminode 4-5 mm long, inserted 6 mm from the base of the
corolla tube; pistil 2.0-2.2 cm long, the ovary cylindric, 2-2.5 mm long and 1 mm
wide, lepidote, the ovules 2-seriate; disc annular-pulvinate, 1.5 mm long and
2.5 mm wide. Capsule linear, 12-47 cm long, 1.6-2 cm wide, conspicuously
glandular-pitted, the surface drying tannish-brown with the pits darker, midrib
not visible, the margins slightly raised, more or less square-cornered; seeds
1.3-1.8 cm long and 3.6-5.2 cm wide, the wings hyaline-membranaceous at the

tips, brown at the base, not sharply distinct from the seed body.
A plant of tropical dry forest and premontane moist forest, Arrabidaea coral-
lina also occurs locally in tropical moist forest. In moist forests, this species is
commonest along more or less exposed sandy or rocky shores where dry edaphic
conditions seem to favor it. It ranges from Mexico to Argentina. This species
flowers during the dry season from February to April.
Arrabidaea corallina is an extremely variable species. However, Central Ameri-
can specimens fall into two separable natural groups: Panamanian specimens
have mostly glabrous leaves with conspicuous tufts in the vein axils beneath;
Costa Rican specimens have strongly pubescent leaves and little development of
tufts. Specimens from Mexico agree with the Costa Rican material. These two
forms may prove to be at least varietally distinct despite the similarity of their
flowers and fruits.
The large corolla and calyx of this species ally it with Arrabidaea mollissima,
but it has a glabrous, pitted fruit as opposed to the evenly pubescent fruit of the
latter. The pubescent form of A. corallina is distinguished from A. mollissima by
its acuminate, more or less cuneate leaflets and the yellowish color of its pubes-
cence and from the superficially similar Xylophragma seemannianum (0. Kuntze)
Sandw. by its exclusively simple trichomes. The less pubescent form of A. coral-
lina is similar to A. florida but differs in its tendency to round-tipped leaflets, and
conspicuously barbate nerve axils.
CANAL ZONE: Near beach at Fort Kobbe, Duke 4250 (MO). Farfan Beach, Gentry 5500
(MO). LOS SANTOS: Las Tablas, Dwyer 1179 (MO). PANAMA': Coronado Beach, Croat
14258 (MO); Gentry 752 (MO). Near archeological site at edge of Madden Lake, Gentry
5007 (MO). Near Bejuco, Miller 1793 (US). Corgona beach, Woodson et al. 1688 (A,
MO, NY).
5. Arrabidaea florida DC., Prodr. 9: 184. 1845. TYPE: Brazil, Amazonas,

Martius s.n. (G-DC, K, M ) .-FIG. 4A-C.
VascQncellia florida Mart. ex DC., Prodr. 9: 184. 1845, pro syn.

Arrabidaea divaricata Bur. & K. Schum., Fl. Bras. 8(2): 33. 1896. TYPE: Peru, Tarapoto,
Spruce 4410 (K, NY).
Bignonia nitida Britton ex Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 34. 1896, pro syn.
TYPE: Bolivia, Mapiri, Rusby 1131 (MO, NY).
Arrabidaea panamensis Sprague, Bull. Herb. Boiss., ser. 2. 6: 371. 1906. TYPE: Panama,
Canal Zone, Hayes 735 (BM, K).
A. cardenasii Rusby, Mem. New York Bot. Card. 7: 351. 1927. TYPE: Bolivia, Rurrenabaque,
Cardenas 2037 (NY).
Liana to 5 cm in diameter, the bark gray, inconspicuously longitudinally

striate; stem with 4 phloem arms in cross section; branchlets terete, with raised
warty lenticels, minutely scabrous when young, soon glabrate, without inter-
petiolar glandular fields; pseudostipules inconspicuous or lacking. Leaves 3-
foliolate or 2-foliolate with a tendril or tendril scar; leaflets ovate to broadly
ovate, acute to acuminate, broadly cuneate to truncate, more or less chartaceous,
secondary nerves 4-7 on each side, beneath with scattered lepidote scales, above
puberulous on the midvein, beneath puberulous on the main veins, drying
olive green to olive gray, the terminal leaflet 8.5-14 cm long and 4.8-8.4 cm
wide; lateral leaflets 5-10.6 cm long and 4-6.3 cm wide; petiole and petiolules
puberulous, terminal petiolule 0.9-2.6 cm long, lateral petiolules 0.5-2.7 cm
long, petiole 0.9-6.2 cm long; tendril simple, 2.5-8 cm long. Inflorescence a
terminal or axillary panicle of several to many dense trichotomous corymbs with
long (1-5 cm) peduncles, its branches puberulous. Flowers sweetly aromatic,
calyx cupular, 5-denticulate, 3-4 mm long and 2-3 mm wide, eglandular, simple-
puberulous; corolla lavender with the tube white inside or white with purple
filaments, abruptly campanulate, 1.2-1.7 cm long and 0.5-0.7 cm wide at the
mouth of tube, the tube 0.7-1.2 cm long, the lobes 0.4-0.6 cm long, pubescent
outside, on the lobes inside and on the bases of the stamens and staminode;
stamens didynamous, subexserted, the anther thecae divaricate, 1.5 mm long,
the longer filaments 0.8-0.9 cm long, the shorter filaments 0.5-0.7 cm long,
the staminode 0.3-0.4 cm long, inserted 2-4 mm from the base of the corolla
tube; pistil 0.8-1.0 cm long, the ovary linear-cylindric, 1-1.5 mm long, 0.5 mm
wide, lepidote, the ovules 2-seriate; disc pulvinate, 1.0-1.5 mm long and 0.5 mm
wide. Capsule linear, acuminate, compressed, 11-22 cm long and 0.9-1.0 cm
wide, scattered lepidote, obscurely and minutely pitted, drying dark gray,
the midrib and margins inconspicuously raised; seeds thin, bialate, 0.5-1.0 cm
long and 2.3-3.6 cm wide, the wings hyaline-membranaceous, sharply demarcated
from the seed body.
Mostly occuring in tropical moist forest, also in premontane wet forest,

this species is never common. It ranges from British Honduras to Bolivia. The
plant flowers during the wet season from July to October (November).
Recognition of sterile plants is difficult, because of lack of noteworthy char-
acters-no interpetiolar glands, no pseudostipules, tendrils not forked, leaves
not noticeably pubescent. This species is vegetatively similar to Adenocalymma
apurense (H.B.K.) Sandw. but lacks cartilaginous margins on the leaflets. It
could also be confused with Tynnanthus croatianus A. Gentry which has trifid
tendrils.
The typically short flowers of the Panamanian.-plants has led to their segre-
gation as Arrabidaea panamensis. Since in all other characters, A. panamensis
is identical with A. florida and size alone is an elusive character in this family,
it is best to consider the Panamanian plants a form of A. florida, especially
inasmuch as there is some overlap in the range of corolla size.
CANAL ZONE: Barro Colorado Island, Aviles s.n. (MO); Bailey & Bailey 560 (CH);
Bangham 449 (A), 465, pro parte (A, F), 465A (A); Chickering 59 (F); Croat 11122 (MO,
NY, SCZ), 11798 (MO, SCZ), 12647 (MO); Foster 1081 (PMA), 1338 (MO); Shattuck
1036 (F, MO); Starry 149 (F, MO). Vicinity of Camboa, Allen 1968 (F, MO, NY, US).
Caillard Highway 2 mi. NW of Summit Garden, Croat 11893 (MO, SCZ). Along old Camboa
road between Summit Hills Golf Course and Summit Radio Station, Croat 16664 (MO). Pipe-
line Road, D'Arcy 6019 (C, MO, PMA, UCWI); Gentry 1430, 4787, 5065 (all MO). Across
from Summit Hills Golf Course, Gentry 1588, 1605, 2818 (all MO), 1774 (MO, SCZ). Rio
Crande Station, Panama Railroad, Hayes 735 (BM, K). Farfan Beach road, Kirkbride & Elias
60 (MO, NY, SCZ). DARIEN: Vicinity of Boca de Cupe, Allen 909 (F, GH, MO, NY, US).
Rio Balsa between Quebrada Chusomocatre and Rio Areti, Duke 8707 (MO). Rio Pifias,
Duke 10577 (MO). Rio Tuira between Rios Punusa and Mangle, Duke 14577 (MO). Shore
near Majagualito across from La Palma, Gentry 3990 (MO). Above jaque, Gentry 4176 (MO).
S of Jaque, Gentry 4279 (MO). Rio Paya from Rio Tuira to village of Paya, Gentry 4388
(MO). PANAMA: Entrado a Nuevo Emperador cerca de Arraijan, Bakes 11 (MO, PMA);
Castillo 14 (MO); Guerra 11 (PMA). Cerro Campana above Su-Lin Motel, Gentry 1844
(MO). Rio Tatare, Woodson & Schery 1017 (CH, MO) . WITHOUT LOCALITY: Bristan 746
(MO).
6. Arrabidaea mollissima (H.B.K.) Bur. & K. Schum. in Mart., Fl. Bras. 8(2):
46. 1896.
Bignonia villosa Vahl, Eclog. Amer. 2: 44. 1798, pro parte. TYPE: Colombia, Magdalena,
von Rohr 21 (C, leaves only).
B. mollissima H.B.K., Nov. Cen. Sp. P1. 3: 133. 1819. TYPE: Venezuela, Valles de Aragua,
B. littoralis H.B.K., Nov. Gen. Sp. P1. 3: 139. 1819. TYPE: Mexico, Cuerrero, Humboldt &
Bonpland 3860 (P).
Panterpa mollissima (H.B.K.) Miers, Proc. Roy. Hort. Soc. 3: 196. 1863.
Bignonia acapulcensis H.B.K. ex Baill., Hist. P1. 10: 28. 1888, pro syn.
Arrabidaea mollicoma Blake, Contr. Cray Herb. 52: 92. 1917. TYPE: Nicaragua, La Paz,
Baker 2268 (CIH, MO, US).
A. seleriana Loes., Verhandl. Bot. Verein Prov. Brandenburg 65: 99. 1923. SYNTYPES: Guate-
mala, Chiquimula, Seler 3348 (GH); Mexico, Oaxaca, Seler 1621 (not seen).
A. isthmica Standl., Jour. Washington Acad. Sci. 15: 461. 1925, pro parte. TYPE: Panama,
Canal Zone, Pittier 2576 (flowers only) (GH, MO, NY, US).
A. littoralis (H.B.K.) Standl., Contr. U. S. Natl. Herb. 23: 1318. 1926.
A. guaricensis Pittier, Jour. Washington Acad. Sci. 18: 337. 1928, pro parte. TYPE: Venezuela,
Cuarico, Pittier 11308 (flowers only) (GH, US, VEN).
A. sanctae-marthae Sprague ex Sandw., Recueil Trav. Bot. Neerl. 34: 229. 1937, nomen nudum.
TYPE: Colombia, Magdalena, Smith 743 (BR, K, MO, NY).
Liana to 3 cm in diameter, the bark smooth, gray; stem with 4 phloem arms
in cross section; branchlets terete, glabrate, usually with interpetiolar glandular
fields, at least at some nodes; pseudostipules small, inconspicuous, narrow and
pointed. Leaves 3-foliolate or 2-foliolate with a tendril or tendril scar, deciduous
during the dry season; leaflets broadly ovate to subrotund, more or less obtuse,
the base rounded or truncate, 3.9-9.3 cm long and 2.4-7.0 cm wide, membran-
aceous, secondary veins 4-5 on each side, scattered lepidote scales on each
side, strongly pubescent with long simple trichomes, especially along main
veins, ciliate, rarely much less pubescent or glabrate, drying olive to brownish
olive, tendril simple, 8-9 cm long; petiolules 0.5-2.5 cm long, the petiole 2-3.7
cm long, pubescent as the petiolules. Inflorescence a large terminal panicle
of a series of axillary panicles, its branches pubescent. Flowers fragrant, calyx
cupular, 5-denticulate, 4-7 mm long and 3-6 mm wide, puberulous with simple
trichomes, the margin ciliate with whitish simple trichomes, usually with a
few scattered glands; corolla lavender outside and on the lobes inside, the
tube white inside with a yellow spot on the roof of corolla over anthers, occa
ionally wholly white; tubular-campanulate, 2.5-5.3 cm long and 0.8-1.6 cm
wide at the mouth, the tube 1.8-3.6 cm long, the lobes 0.6-1.2 cm long; monili-
form-pubescent outside and on margins of lobes, simple pubescent inside with
multicellular trichomes on lobes and with 1-3-celled, gland-tipped trichomes
at and just below the level of stamen insertion and on the floor of the tube;
stamens didynamous, the anther thecae divaricate, 4-5 mm long, the longer
filaments 1.5-1.7 cm long, the shorter filaments 1.1-1.2 cm long, the staminode
1-2 mm long, inserted 5-7 mm from base of corolla tube; pistil 2-2.1 cm long,
the ovary tapered-cylindric, 3 mm long, 1 mm wide, glandular-puberulous or
lepidote, the ovules 2-seriate in each locule; disc pulvinate, 1 mm long 2 mm

wide. Capsule linear, 14.5-17 cm long, 1.2-1.5 cm wide, softly pubescent, dry-
ing brownish, the midrib inconspicuous but sometimes slightly raised; seeds
thin, 1.1-1.4 cm long and 3.6-4.3 cm wide, bialate, the wings hyaline-membra-
naceous.
Restricted to tropical dry forest and drier parts of the premontane moist
forest and tropical moist forest. A. mollissima is common in Guanacaste Prov-
ince, Costa Rica, and sporadic along the Pacific slope from Mexico to Venezuela.
The species flowers during the dry season from late January to early March.
Deciduous during the blooming period, its lavender flowers often cover a
leafless dry forest tree creating a spectacular appearance similar to a flowering
tree of Tabebuia impetiginosa.
The round pubescent leaves with conspicuously ciliate margins are a good
field character. Its relatively large corolla and calyx separate A. mollissima
from other Central American species of Arrabidaea except A. corallina from
which it is most easily distinguished by its soft-pubescent fruit. A white-flowered
form is clearly conspecific and does not merit formal recognition.
Confusion surrounds the name A. littoralis, which Standley apparently used
as a catch-all for anything close to Arrabidaea which he could not otherwise
name. Arrabidaea littoralis as determined by Standley in various herbaria in-
cludes such diverse elements as Clytostoma binatum (Haenke 1022), Pachyptera
alliacea (Haenke 1518), Arrabidaea costaricensis (Molina 2725), A. corallina
(Standley 40042, Williams & Molina 12581), A. pubescens (Kenoyer A348),
A. patellifera (Palmer 362, pro parte), and A. mollissima (Edwards P-577).
- CANAL zoNE: Road K-10, Duke 15193 (MO). Near junction Gamboa Road, Harvey
5242 (F). Chiva Chiva Trail, Red Tank to Pueblo Nuevo, Piper 5751 (US). Rio Agua Salud
near Corozal, Piper 5855 (US). Vicinity of El Paraiso, Pittier 2576, pro parte (CGH, MO, US).
Near Corozal Hospital, Wetmore & Abbe 276 (A). Ancon Hill, Williams 14 (US). COCLE:
Mountains beyond La Pintada, Hunter & Allen 510 (K, MO, US), 511 (MO, NY, P).
Penonomre and vicinity, Williams 83 (NY, US), 427 (NY), 428 (US). DARIEN: Punto Sabanas
opposite La Palma, Gentry 3951, 3956 (both MO). Punta Patifia NNE of Carachine, Gentry
4052 '(MO). PANAMA: Along road to Pacora, Allen 841 (K, MO, P, US). Rio Las Lajas,
Allen 1602 (MO). Vicinity of Arraijan, Allen 1623 (F, CH, MO, NY, US). Vicinity of Bejuco,
Allen 4202 (F, MO). Rio Pasiga, Gentry 2268 (MO). Icanti, Gentry 2605 (MO). Trail
from Rio Espave toward Rio Agua Clara, Gentry 3777 (MO). 0-4 km from Bayano crossing
on road to Santa Fe, Gentry 3860 (MO). Along road to Cerro Azul, Gentry 4896 (MO). E of
Cafiitas, Gentry 4974 (MO). Road to Cerro Azul, Gentry et al. 3393 (MO). Punta Paitilla,
Piper 5395 (US). Between Matias Hernandes and Juan Diaz, Standley 32055 (K, US).
VERAGUAS: 6 mi. S of Santa Fe, Gentry 2932 (MO).
7. Arrabidaea oxycarpa Urb., Feddes Repert. 14: 301. 1916. TYPE: Trinidad,
Broadway 3188 (not seen).
Vine, the branchlets terete, lepidote to slightly puberulous, when young dry-
ing black with contrasting whitish lenticels, without interpetiolar glandular
fields; pseudostipules small, blunt, inconspicuous. Leaves 3-foliolate or 2-foliolate
with a simple tendril, the blade elliptic, somewhat acuminate, rounded to broadly
cuneate at base, 3-12 cm long and 1-6 cm wide (smaller at base of branchlet),
chartaceous (usually immature and membranaceous when in flower), slightly
lepidote, otherwise glabrous except for simple trichomes in axils of lateral nerves
beneath, drying lead gray to blackish, the main veins darker beneath, the ultimate
venation intricately reticulated; petiolules 0.2-3 cm long, the terminal longer,
laterals shorter, petiole 1.5-5 cm long, lepidote, drying black. Inflorescence a
reduced, few-flowered panicle terminal on young lateral branchlets, its branches
lepidote and puberulous, drying blackish, bracts minute. Flowers with the calyx
cupular, 3-4 mm long and 3-4 mm wide, evenly but minutely 5-denticulate,
lepidote and puberulous, usually with conspicuous plate-shaped glands near
margin, drying dark; corolla magenta, tubular campanulate, 3-3.5 cm long and
1.3-1.4 cm wide, the tube 2.2-2.4 cm long, the lobes ca. 1 cm long, densely
puberulous without and on lobes within, the trichomes somewhat glandular,
the tube inside glabrous except for trichomes at base of stamens; stamens
didynamous, the anther thecae divaricate, 3 mm long, the connective slightly
extended, the longer filaments 1.6-1.7 cm long, the shorter filaments 1.2-1.3 cm
long, the staminode 6-7 mm long; inserted 5 mm from base of corolla tube;
pistil 1.6-1.7 cm long, the ovary linear oblong, 2-3 mm long and 1 mm wide,
minutely lepidote, the ovules 2-seriate in each locule; disk pulvinate, 1 mm long
and 1.5-2 mm wide. Capsule linear, compressed, tapered at both ends, 19-23
cm long and 1 cm wide, drying brown or blackish, glabrous or slightly lepidote,
the midrib evident but slightly raised and inconspicuous, the margins somewhat
raised; seeds thin, flat, bialate, 0.7-0.8 cm long and 1.7-2.8 cm wide, the wings
sharply divided into a brown basal part and a hyaline membranaceous tip.
An uncommon vine of the premontane moist forest and edaphically dry

areas of the tropical moist forest, A. oxycarpa has been previously known only
from Trinidad.
- I have seen only a single specimen of this species from Trinidad. However,
I have recently examined a series of specimens from Venezuela which I would
identify with A. oxycarpa and am now convinced that two fruiting collections
from the Canal Zone which I had tentatively identified with A. florida are
actually A. oxycarpa. Arrabidaea oxycarpa differs from A. florida in mature
branchlets with a loose, inconspicuously lenticellate, somewhat wrinkled epi-
dermis in contrast to the tight, unwrinkled, uniformly small-lenticellate epidermis
of the latter. The leaves of A. oxycarpa also tend to dry a darker gray with a
more intricate pattern of ultimate reticulations below. In flower A. oxycarpa is
characterized by young branchlets which dry blackish with conspicuous light
lenticels and by its reduced, few-flowered inflorescences.
CANAL ZONE: Boy Scout Camp Road near Madden Lake, Gentry 5504 (MO). Farfan
Beach Road, Kirkbride & Elias 60 (MO, NY, SCZ).
8. Arrabidaea patellifera (Schlecht.) Sandw., Kew Bull. 22: 413. 1968.
Bignonia patellifera Schlecht., Linnaea 8: 516. 1833. TYPE: Mexico, Hacienda de la Laguna,
Schiede & Deppe 153 (BM, MO).
B. cupulata Splitg., Tijdschr. Natuurl. Gesch. Physiol. 9: 6. 1842. TYPE: Brazil, Para (not seen).
Tabebuia neurophylla Miq., Linnaea 26: 219. 1853. TYPE: Surinam, Kappler 1957 (C, U).
Petastoma patelliferum (Schlecht.) Miers, Proc. Roy. Hort. Soc. 3: 195. 1863.
Arrabidaea pentstemonoides Kranzl., Notizbl. Bot. Gart. Berlin 6: 370. 1915. TYPE: Brazil,
Rio Acre, Ule 9780 (K).
Cuspidaria mollis Kranzl., Notizbl. Bot. Gart. Berlin 6: 376. 1915. TYPE: (not seen).
Petastoma broadwayi Sprague & Riley, Kew Bull. 1921: 311. 1921. TYPE: Trinidad, Broadway
4164 (K, NY).
P. breviflorum Standl., Jour. Arnold Arbor. 11: 128. 1930. TYPE: Panama, Barro Colorado
Island, Bangham 465, pro parte (A, F).
P. domatiatum Sampaio, Bol. Mus. Nac. Rio de Janeiro 12: 87. 1931. TYPE: Brazil, Amazonas,
Ducke 24089 (K).
P. pentstemonoides (Kranzl.) Sandw., Kew Bull. 43: 7. 1959.
Arrabidaea broadwayi (Sprague & Riley) Sandw., Kew Bull. 22: 414. 1968.
Liana to at least 5 cm in diameter, the bark light gray; stem with 4 phloem
arms in cross section; branchlets light gray or brown usually with raised whitish
lenticels, glabrous to lepidote or scabrous, the nodes without interpetiolar glan-
dular fields, often with a ridge connecting opposite petioles; pseudostipules
usually lacking, when present foliaceous, small, soon deciduous. Leaves simple
or 2-foliolate, sometimes with a tendril; leaflets (or leaf blades) ovate to rhombic-
ovate, sometimes serrate, acuminate to obtuse, broadly cuneate to rounded 4.7-
14.5 cm long and 2.9-10.5 cm wide, membranaceous, the secondary veins 4-6
on a side, the basal nerves often making more acute angle with midvein than
upper nerves, somewhat lepidote above and beneath, conspicuously simple-
puberulous beneath mostly on the nerves, less so above (rarely glabrate, e.g.
Gentry 1955), drying more or less olive, the pubescent veins lighter beneath;
tendril simple, 4-16 cm long; petiole and petiolules conspicuously pubescent
with long simple trichomes, petiolules 0.5-2.2 cm long and petiole 0.7-1.9 cm
long in compound leaves, petiole 1.1-3.5 cm long in simple leaves. Inflorescence
an axillary or terminal, usually many-flowered panicle, the branches puberulent
and lepidote, the buds appearing conspicuously white-tipped from the pubes-
cent corolla lobes. Flowers sweetly aromatic, calyx open, patelliform, more or
less truncate, 1-4 mm long and 4-8 mm wide, scattered lepidote and simple-
pubescent, glands absent; corolla magenta, the throat usually white with magenta
ridges, sometimes wholly magenta, campanulate above the narrowed base, 2.2-
4.3 cm long and 0.5-1.7 cm wide at the mouth, the tube 1.4-2.6 cm long with
basal constriction 0.5-0.8 cm long, the lobes 0.4-1-2 cm long; the tube glabrate
outside with occasional widely scattered lepidote scales or simple trichomes,
the lobes densely simple pubescent, tube glabrous inside except for gland-
tipped trichomes at the level of stamen insertion; stamens didynamous, anther
thecae divaricate, bent forward slightly, 1.5-2 mm long, the longer filaments
1.1-1.3 cm long, the shorter filaments 0.8-1.0 cm long; the staminode 3-4 mm
long, inserted 5-7 mm above the base of the corolla tube; pistil 1.5-2.3 cm long,
the ovary linear, 1.5-2.5 mm long and 0.5-1 mm wide, slightly lepidote, the
ovules 2-seriate; disc annular-pulvinate, 0.5-1 mm long and 1.5-2 mm wide.
Capsule linear, acute, flattened, 11-39 cm long and 0.9-1.4 cm wide, the midrib
appearing as a slightly raised line, the surface generally minutely papillose and
smooth, rougher on the slightly raised margins with a few scattered trichomes
and lepidote scales; seeds 0.7-1.0 cm long and 2.5-3.9 cm wide, the wings
membranaceous with hyaline ends distinct from the brown bases, the wing
bases not demarcated from the seed body.
A widespread liana common in the tropical dry forest and premontane moist
forest, it becomes less common in tropical moist forest and rare in tropical wet
forest. It ranges from Mexico to Brazil. This species flowers during the wet
season from June to November, also rarely and irregularly in January or Feb-
ruary. The wind-dispersed seeds are released in mid dry season.
Useful vegetative charcters are the typically strongly pubescent main veins
beneath, the pubescent petioles and petiolules, and the frequent simple leaves.
The sharp angle of the basal lateral veins with the midvein is another diagnostic
character. The reflexed patelliform calyx and white-tipped bud are distinctive,
but the fruit is similar to that of several other species of Arrabidaea.
Petastoma breviflorum is no more than a form with atypically short corollas;
while I have seen no other specimens of A. patellifera with such short corollas,
intensive collecting on Barro Colorado Island has turned up no additional
material of this sort. Even greater variation in corolla length is well documented
within other species of Bignoniaceae, even on the same plant. '
Both the segregating characters (larger calyx and less pubescent or almost
glabrous leaves) of Arrabidaea broadwayi from Trinidad are within the range
of variation of A. patellifera. Also falling into this category are P. domatiatum
and A. pentstemonoides. Arrabidaea patellifera itself is close to A. samydoides
(Cham.) Sandw. of Brazil. Should they prove synonymous, the Brazilian name
would have to be adopted.
CANAL ZONE: Barro Colorado Island, Aviles 118, 889 (both F); Bailey & Bailey 280
(F, CH); Bangham 465 (A, F), 553 (A); Croat 5222 (NY, SCZ), 6065 (MO, SCZ), 6654,
6724, 9794, 10895 (all MO), 11126 (F, MO, NY, SCZ), 13161 (MO, SCZ), 13244A, 15144
(both MO); Dodge & Woodworth 3506 (GH, NY, US); Ebinger 276 (MO, US); Foster 1024
(MO, PMA), 1338B (MO); Kenoyer 531 (US); Shattuck 258 (A, F, MO), 300 (F, MO),
889 (MO); Starry 202 (F, MO); Wetmore & Abbe 71 (A, F, MO), 163 (A, F, MO); Wetmore
& Woodworth 78 (F); Woodson & Schery 970 (CGH, MO); Woodworth & Vestal 360 (MO).
Near water reservoir W of Cocoli, Croat 9162 (MO, SCZ). Ancon Hill, Duke 4606 (K, MO).
Pipeline road, Gentry 1585, 1955 (both MO). Road K-10, Gentry 1961 (MO, SCZ). Between
Corozal and Ancon, Pittier 6771 (NY, US). Balboa, Standley 26991 (MO, US). Vicinity of
Miraflores near mouth of Cocoli River, White 128 (GH, MO). Fort Kobbe road, Woodson
et al. 1416 (A, MO, NY). CHIRIQUi: Trail from San--Felix to Cerro Flor, Allen 1918 (MO).
E of Gualaca, Allen 5031 (MO). DARIEN: Rio Chica vicinity of Yaviza, Allen 4577 (F, K,
MO, P, US). Between Quebrada Venado and Peje swamp on headwaters of Rio Tuqueza,
Bristan 1054 (F, MO, NY). Rio Ucurganti, Bristan 1156 (MO). Tumaganti, Duke 14146
(MO, SCZ). Trail to Cerro Pirre, Gentry 4608 (MO). HIERRERA: Road from La Arena to
Pese, Burch et al. 1284 (MO). Road between Las Minas and Pese, Duke 12333 (MO). E of
Las Minas, Gentry 3136, 3141, 3147 (all MO). PANAMA: Vicinity of Arraijain, Allen 1621
(F, GH, MO, NY, US). SE part of Gatun Lake near Rio Pescado, Bartlett & Lasser 16595
(MO). Rio Charco-Espiritu on Tocumen Highway, Duke 5703 (MO, SCZ). Bald savanna-
like areas along road toward top of Cerro Campana, Duke 5977 (GH, MO). Rio Maestro,
Gentry 2234 (MO). Rio Pasiga, Gentry 2351 (MO). Near Rio Espave, Gentry 3707
(MO). Chiman, Lewis et al. 3352 (MO, SCZ). Cerro Campana, McDaniel 6811A (MO).
Rio Tapia, Standley 28131 (US). VERAGUAS: Santiago toward Atalaya, Dwyer & Kirkbride
7401 (GH, K, MO, US). 2 mi. W of Santiago, Dwyer et al. 7559 (MO). S of Santa Fe,
Gentry 2936 (MO). Vicinity of Santiago, Stern et al. 997 (GH, MO). 2 mi. S Caniazas,
Tyson 3729 (MO).
9. Arrabidaea pubescens (L.) A. Gentry. Brittonia 25: 239. 1973.
Bignonia pubescens L., Sp. P1., ed. 2. 2: 870. 1763. TYPE: Mexico, Campeche, Houstoun s.n.
(not seen).
Arrabidaea blanchetii DC., Prodr. 9: 186. 1845. TYPE: Brazil, Bahia, Blanchet 2363 (G-DC).
A. pleei DC., Prodr. 9: 184. 1845. TYPE: Venezuela, Maracaibo, Plee 89 (G-DC, K, P, US).
A. sieberi DC., Prodr. 9: 186. 1845. TYPE: Trinidad, Sieber 109 (G-DC, GH).
Petastoma pubescens (L.) Miers, Proc. Roy. Hort. Soc. 3: 195. 1863.
Arrabidaea carabobensis Pittier, Jour. Washington Acad. Sci. 18: 335. 1928. TYPE: Venezuela
Carabobo, Pittier 7915 (VEN).
A. lundellii Standl., Publ. Field Mus. Bot., 8: 48. 1930. TYPE: British Honduras, Orange W
Lundell 56 (F, holotype; US, isotype).
Liana to 2.5 cm in diameter, stem with 4 phloem arms in cross section;

branchlets terete, puberulous with interpetiolar glandular fields; pseudostipules
inconspicuous or lacking. Leaves 3-foliolate or 2-foliolate with a tendril or
tendril scar; leaflets ovate, acute to acuminate, rounded to broadly subcordate
at base, 4.5-13 cm long and 2.5-8 cm wide, membranaceous, secondary nerves
4-5 on a side, palmately 3-5-veined from base, densely short-puberulous beneath,
above puberulous with extremely short trichomes to glabrate with scattered tri-
chomes, drying olive to brown above, tan beneath, the veins pubescent, beneath
and colored about the same as the surface; tendril simple, 5-7 or more cm long;
petiolules 0.8-1.8 cm long, petiole 1.7-3.1 cm long, densely short-puberulous.
Inflorescence an axillary panicle, its branches puberulous. Flowers with the
calyx cupular, truncate, minutely 5-denticulate, 3 mm long and 2-3 mm wide,
puberulous, usually with some glands; corolla lavender, campanulate, 1.3-1.8
cm long and 0.5-0.7 cm wide at the mouth, the tube 0.8-1.1 cm long, the
lobes 0.3-0.5 cm long, the tube pubescent outside, abaxially pubescent inside
and at the level of stamen insertion, the lobes puberulous; stamens didynamous,
the anther thecae divaricate, 2 mm long, the longer filaments 0.9-1.0 cm long,
the shorter filaments 0.6-0.8 cm long, the staminode 2-4 mm long, inserted
2-4 mm from base of corolla tube; pistil 1.0-1.1 cm long, the ovary oblong,
densely lepidote, 1.5 mm long and 0.5-1 mm wide, the ovules 2-seriate; disc
pulvinate, 1 mm long and 1-1.5 mm wide. Capsule linear, acuminate, com-
pressed, 7.2-8.8 cm long and 0.6-0.8 cm wide, lepidote, drying dark brown,
the midline and margins slightly raised; seeds thin, bialate, 5 mm long, 1.4-1.5
cm wide, the wings hyaline-membranaceous, distinct from the dark brown
seed body.
A plant mainly of the tropical dry forest, A. pubescens is widely distributed

from Tamaulipas, Mexico, to Brazil, mostly near the coast. It flowers during
early and mid-wet season from June to August.
The complex synonymy of this species is discussed elsewhere (Gentry, Brit-
tonia 25: 238-239. 1973).
CANAL ZONE: Farfan Beach Road, Gentry & Dwyer 3545 (MO). COCLE': Between
Aguadulce and Ant6n, Woodson et al. 1224 (A, K, MO, NY). DARIEN: Punta Patifio NNE
of Garachine, Gentry 4042 (MO). Near Punta Alegre NE of Garachine, Gentry 4060 (MO).
PANAMA: Rio Las Lajas, Allen 3613 (F, GH, K, MO, NY, U, US). Mouth of Rio Pasiga,
Gentry 2201 (MO). Near Panama, Hayes 93 (BM, K). Gorgona Beach, Woodson et al.
1689 (A, MO, NY). Taboga Island, Gentry 5713 (MO), Woodson et al. 1489 (A).
5. CALLICHLAMYS
Callichlamys Miq., Linnaea 18: 254. 1844. TYPE: C. riparia Miq. = C. latifol
(L. C. Rich.) K. Schum.
Lianas, but shrubby when young, stems with 4 phloem arms in cross section;
branchlets terete without interpetiolar glandular fields; pseudostipules lacking.
Leaves 3-foliolate, the terminal leaflet sometimes replaced by a simple tendril.
Inflorescence a short axillary (rarely terminal), 1-12-flowered raceme. Flowers
with the calyx of a thick spongy material, inflated-cupular, irregularly lobed;
corolla large, infundibuliform-campanulate, the tube contracted above the ovary,
sparsely and inconspicuously glandular-lepidote outside; anthers glabrous, the
thecae straight, divaricate; pollen grains 3-colpate, microreticulate; ovary ovate-
cylindric, narrowed slightly at the base, more or less glabrous, the ovules 4-8-
seriate in each locule; disc annular-pulvinate. Fruit an oblong-elliptic or oblong
capsule, the valves parallel to the septum, compressed, woody, smooth; seeds
large, thin, the wings brown, not demarcated from the seed body.
One species ranging from Mexico to Brazil.
Callichlamys latifolia (L. C. Rich.) K. Schum. in Engler & Prantl, Nat.

Pflanzenf. 4 (3b): 223. 1894.-FIG. 5.
Bignonia latifolia L. C. Rich., Act. Soc. Hist. Nat. Paris 1: 110. 1792. TYPE: French Guiana,
Rio Kourou, Le Blond s.n. (P).
B. hondensis H.B.K., Nov. Gen. Sp. P1. 3: 135. 1819. TYPE: Colombia, Honda, Humboldt &
Bonpland 1706 (P).
Tabebuia latifolia (L. C. Rich.) DC., Rev. Bign. (Bibl. Univ. Geneve) 15. 1838.
Delostoma latifolia (L. C. Rich.) Splitg., Tijdschr. Natuurl. Gesch. Physiol. 9: 11. 1842.
Callichlamys riparia Miq., Linnaea 18: 254, 609. 1844. TYPE: Based on Bignonia latifolia
L. C. Rich.
Bignonia crucifera Bert. ex DC., Prodr. 9: 213. 1845, pro syn.
Tabebuia rufinervis DC., Prodr. 9: 213. 1845. SYNTYPES: Colombia, Santa Marta, Bertero s.n.
(G-DC, M); British Guiana, Schomburgk 575 (F, G-DC); Brazil, Para, Sieber s.n.
(not seen); Rio de Janeiro, Luschnath s.n. (not seen).
T.? mansoana DC., Prodr. 9: 214. 1845. TYPE: Brazil, Cujaba, Manso s.n. (G-DC).
Lundia schomburgkii Kl. in Schomburgk, Reisen 2: 1158. 1848, nomen nudum.
Callichlamys rufinervis (DC.) Miers, Proc. Roy. Hoit. Soc. 3: 201. 1863.
C. rubiginosa Miers, Proc. Roy. Hort. Soc. 3: 201. 1863, nomen nudum. TYPE: Brazil, Organ
Mountains, Miers s.n. (not seen).
Callichlamys splendida Miers, Proc. Roy. Hort. Soc.- 3: ,201, nomen nudum. TYPE: Braz
Rio de Janeiro, Cunningham s.n. (not seen).
C. peckoltii Bur. ex K. Schum. in Engler & Prantl, Nat. Pflanzenf. 4(3b): 223, fig. 88M-N.
1894. SYNTYPES: Brazil, Rio de Janeiro, Peckolt 533 (BR); Peckolt 592 (not seen).
Tabebuia speciosa Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 49. 1930. TYPE: British
Honduras, Middlesex, Schipp S-51 (F).
Callichlamys garnieri Standl. & L. 0. Wms., Ceiba 3: 130. 1952. TYPE: Nicaragua, Managua,
Garnier 130 (F).
Liana to 7.5 cm in diameter, the bark light gray, smooth with scattered
raised whitish lenticels; stem with 4 phloem arms in cross section; twigs sub-
terete, light gray with large, widely scattered raised lenticels, the nodes without
interpetiolar glandular fields, interpetiolar ridges, or noticeable pseudostipules.
Leaves 3-foliolate or 2-foliolate with a tendril or an inconspicuous tendril scar;
leaflets elliptic, acuminate, broadly cuneate to rounded, membranaceous to
chartaceous, secondary veins 5-9 on a side, minutely papillose above and
beneath with scattered lepidote scales, these sometimes deciduous above, pu-
bescent beneath in the axils of lateral nerves, usually subpuberulous on main
veins with simple and stellate trichomes, sometimes conspicuously dendroid
A D E
FIGURE 5. Callichiamys latifolia (L. Rich.) K. Schlum.-A. Habit ( X 1/2). [After

Gentry 1896 (MO).]-B. Fruit ( X 1/2). [After Croat 8058 (MO).]-C. Pistil and disc
(X% /) .-D. Stamen (X 3) .-E. Ovary cross section (X 6). [After Gentry 1777 (MO).]
pubescent on the undersurface, with plate-shaped glands along the midvein,

mostly in axils of the lateral nerves, drying green to olive with the main nerves
yellowish or contrastingly reddish, the terminal leaflet 9.5-37 cm long and 4.2-22
cm wide, the lateral leaflets 7-31 cm long; and 3.4-18 cm wide; terminal petiolule
1.8-9.5 cm long, lateral petiolules 1.1-4.8 cm long, petiole 2.5-18 cm long,
glabrous to minutely scabrous as the petiolules; tendril simple 9-20 cm long.
Inflorescence a short terminal or axillary, 1-12-flowered raceme, the pedicels

1-1.5 cm long, each with 2 narrow bracts, the rachis to 8 cm long, the rachis
and pedicels sparsely stellate-puberulent. Flowers with an inconspicuous sweet-
ish musky odor, calyx large, yellow when fresh, spongy, inflated-campanulate,
more or less bilabiate, mostly glabrous with scattered plate-shaped glands,
sparsely and inconspicuously thick-stellate and/or simple puberulent near the
base, or rarely conspicuously stellate pubescent, 2.2-6.0 cm long and 1.2-4.2 cm
wide; corolla bright yellow with red penciling in the throat (13 lines), tubular-
campanulate, 5.5-11 cm long and 1.3-2.8 cm wide at the mouth, the tube 4.5-7.5
cm long, the lobes 1.2-3.2 cm long; sparsely and inconspicuously lepidote inside
and out, glandular-pubescent inside from the level of stamen insertion to the
top of the enlarged base; stamens didynamous, the anther thecae divaricate,
2-3 mm long, the filaments abruptly narrowed for the apical 2 mm, the longer
pair 2.5-3.8 cm long, the shorter pair 1.5-2.3 cm long, the staminode 0.6-1.1 mm
long, inserted 1.2-2.0 cm from base of corolla tube; pistil 5.2-5.7 cm long, the
ovary ovate-cylindric, 3-4 mm long and 2.5-3.5 mm wide, narrowed slightly
at the base, essentially glabrous, the ovules 8-seriate in each locule; disc annular-
pulvinate, 2 mm long and 6-7 mm wide. Capsule oblong or elliptic-oblong,
obtuse at the ends, the valves woody, flattened, 24-32 cm long and 6.0-11.5 cm
wide, the surface smooth, glabrous with a wrinkled epidermis, drying blackish
with scattered lighter inconspicuous lenticellate spots; seeds 2.5-4.3 cm long
and 6.7-13 cm wide, the wings brown, thinly woody at the base to submem-
branaceous at the tips, not distinct from the seed body.
Widespread in tropical moist forest and tropical wet forest, but seldom
abundant, this plant ranges from Mexico to Brazil, flowering mostly toward
the end of the wet season in October and November but sporadically throughout
the year.
The plant is characterized by large leaflets which have conspicuous tufts of
dendroid trichomes in the axils of the lateral nerves beneath. The smooth gray
bark, with whitish lenticels is another field character. On drying, the leaves
can usually be recognized by the red-drying main veins. The large flat fruit
is almost unmistakable.
A second species of Callichlamys, C. garnieri Standl. and L. 0. Wms., has
recently been described from Nicaragua. This is known only from the type
and was differentiated from C. latifolia mainly on the basis of a larger flower
and strongly pubescent leaves and calyx. It is a pubescent variant of C. latifolia.
Another pubescent variant was described by De Candolle as Tabebuia mansoana.
BOCAS DEL TORO: Mountains above Almirante, Gentry 2734, 2812 (both MO). CANAL
ZONE: Barro Colorado Island, Aviles 16B (F, MO); Croat 6819 (MO, NY, SCZ), 6835 (MO),
6854A (MO, NY, SCZ), 8058 (MO, NY, SCZ); Foster 1383 (MO, PMA); Shattuck 361 (F,
MO). Near Rio Gatuncillo, Allen 5137 (MO). S of Pedro Miguel Locks, Croat 9173 (MO,
SCZ). Madden Lake, Gentry 702 (F, MO), 703 (MO). Madden Forest, Gentry 1383 (MO).
Pipeline Road, Gentry 1581 (MO). Road K-19, Gentry 1777 (MO, SCZ). Road K-10, Gentry
1896 (MO, SCZ). Boy Scout Road near Madden Lake, Gentry 2073 (MO). Empire Station,
Panama Railroad, Hayes 248 (BM, K, P, W). N of Paraiso, Lazor & Blum 5319 (SCZ). Near
Contractor's Hill, Lewis et al. 2880 (MO, NY, SCZ). COLON: Santa Rita Ridge, Dressler 3513
(PMA); Gentry 461 (MO). DARIEN: Rio Balsas above Rio Areti, Duke 8710 (MO). S of
Jaque', Gentry 4106 (MO). Above Rio Clarita on Rio Tuira, Gentry 4322 (MO). Rio Tuira
near Rio Paya, Gentry 3378 (MO). Rio Tuira between Rios Paya and Espave, Gentry 4421
(MO). Summit of Cerro Pirre, Gentry 4617 (MO). PANAMA: Near Vigia and San Juan on
Rio Pequeni, Dodge et al. 16519 (BM, C, K, MO, P, U). Dam site at Rio Bayano, Gentry &
Tyson 1662 (MO). Rio Pasiga, Gentry 2200, 2258, 2309, 2337 (all MO). Rio Maestro,
Gentry 2212, 2215 (both MO). Near Rio Parti, Gentry 3800 (MO). Rio Bayano above Rio
Espave, Gentry 3850 (MO). San Jose Island, Johnston 508 (GH), 1161 (BM, GH, MO, P,
U, US). Agricultural Experiment Station at Matias Hernaindez, Pittier 6861 (BM, F, GH,
MO, NY, US). SAN BLAS: Mountains above Puerto Obaldia, Gentry 1502 (MO). VERAGUAS:
Above Santa Fe, Gentry 3024 (MO).
6. CERATOPHYTUM
Ceratophytum Pittier, Jour. Washington Acad. Sci. 18: 62. 1928. TYPE: C.
capricorne Pittier = C. tetragonolobum (Jacq.) Sprague & Sandw.
Lianas; stems with 4 phloem arms in cross section; branchlets terete, with
conspicuous interpetiolar glandular fields; pseudostipules of 3 acute, more or
less subulate scales appressed against the twig. Leaves 3-foliolate, the terminal
leaflet often replaced by a trifid tendril. Inflorescence a few-flowered terminal,
subcorymbose panicle. Flowers with the calyx cupular, more or less truncate,
coriaceous, lepidote, with short, linear, glandular fields descending from the
rim; corolla cream to pale yellow, thick, tubular-campanulate, pubescent outside;
anthers glabrous, the thecae straight, divaricate; pollen grains simple, 3-colporate,
the exine reticulate; ovary narrowly cylindric, more or less lepidote, the ovules
6-8-seriate in each locule; disc pulvinate-cupular. Fruit an oblong-linear, sub-
tetragonal capsule; gibbous at the base and tapering to the apex, the valves
parallel to the septum, not flattened, smooth; seeds bialate, the wings papery,
brownish, the tip usually subhyaline, not clearly demarcated from the seed body.
One species ranging from Mexico to Trinidad and Guyana.
Ceratophytum tetragonolobum (Jacq.) Sprague & Sandw., Kew Bull. 1934:

222. 1935.-FIG. 6.
Bignonia tetragonoloba Jacq., Fragm. Bot. 36, tab. 40, fig. 2. 1800-1809. TYPE: Venezuela,
Caracas (not seen).
Anemopaegma tobagense Urb., Fedde Repert. 14: 311. 1916. TYPE: Tobago, Broadway 3493
(K, NY).
Ceratophytum capricorne Pittier, Jour. Washington Acad. Sci. 18: 63. 1928. TYPE: Venezuela,
Distrito Federal, Pittier 11527 (K, US, VEN).
C. brachycarpum Pittier, Jour. Washington Acad. Sci. 18: 65. 1928. TYPE: Venezuela,
Aragua, Pittier 11805 (K, fragments; US, VEN).
Adenocalymma heterophyllum Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 49. 1930,
non Kranzl. TYPE: Mexico, Yucatan, Gaumer & Sons 23889 (F).
Ceratophytum tobagense (Urb.) Sprague & Sandw., Kew Bull. 1933: 322. 1933.
Adenocalymma standleyanum Lundell, Carnegie Inst. Washington Publ. 478: 221. 1937.
TYPE: Based on A. heterophyllum Standl.
Liana to at least 7 cm in diameter, stem often indistinctly 4-parted and

twisted, the nodes with a swollen annulus, the bark gray, smooth, sometimes
with raised whitish lenticels; stem with 4 phloem arms in cross section; branch-
lets terete, longitudinally finely striate, lepidote, the nodes with interpetiolar
glandular fields; pseudostipules of 3 subulate scales, appressed against the twig
.. .... ....~ ~ ~ ~~~~~~~~~~~~~~~~. ~
FIGURE 6. Ceratophytum tetragonolobum (Jacq.) Sprague & Sandw.-A. Habit (x 3/8).

[After Gentry 722 (MO).]-B. Fruit (X 3/1s). [After Gentry 3838 (MO).]-C. Corolla
(x 1/2).-D. Calyx ( X 1/2).-E. Ovary cross section ( X 9).-F. Pistil and disc ( x 1/2)
[After Gentry 2195 (MO).]
when fresh, when dry the 2 outer scales divergent. Leaves 3-foliolate or 2-folio-
late with a tendril or tendril scar; leaflets oblong-ovate to elliptic-obovate, acute
to obtuse, truncate to asymmetrically subcordate, 6.2-18 cm long and 2.1-15 cm
wide, membranaceous to chartaceous, secondary veins 5-6 on a side, mostly
glabrous or inconspicuously lepidote, inconspicuously simple-puberulent in the
nerve axils beneath and at the base of the main veins above, drying olive; tendril
trifid, 5-20 cm long to branching, the 3 arms 0.4-2.5 cm long; terminal petiolule
2.1-4.6 cm long, laterial petiolules 0.3-5.7 cm long; petiole 3.0-12 cm long,
lepidote or sparsely puberulous as the petiolules. Inflorescence a terminal, few-
flowered, subcorymbose panicle, often contracted to one or 2 flowers, its
branches lepidote. Flowers with the calyx cupular, truncate, 8-13 mm long
and 9-10 mm wide, lepidote, with 4-6 linear glandular fields descending from
the rim; corolla thick, cream-colored to pale yellow outside and on the lobes
inside, the throat yellow inside; tubular-campanulate, 7.7-8.1 cm long and 1.7-2.1
cm wide at the mouth, the tube 5.6-5.8 cm long, glabrous inside but densely
pubescent with long, simple, multicellular hairs just below the level of stamen
insertion, the lobes 1.6-2.2 cm long, densely glandular-lepidote to glandular-
pubescent outside, glandular-lepidote on the lobes; stamens didynamous, the
anther thecae divaricate, 4.5-6 mm long, the longer filaments 2.2-2.4 cm long,
the shorter filaments 1.4-1.6 cm long, the staminode 5-6 mm long, inserted
9-12 mm from the base of the corolla tube; pistil 3.2-3.3 cm long, the ovary
narrowly cylindric, 5 mm long, 2 mm wide, lepidote, the ovules 6-8 seriate in
each locule; disc pulvinate-cupular, 2 mm long, 4 mm wide. Capsule cylindrical,
basally gibbous, tapering upwards, 26-39 cm long and 3-3.9 cm wide, 2.2-2.6
cm thick, the surface tan, more or less lepidote; seeds biseriate, 0.8-1.8 cm long
and 3.4-6.3 cm wide, the wings papery, brown, not demarcated from the seed
body.
A vine characteristic of the tropical moist forest, it is seldom common and

is poorly represented in herbaria, despite its wide range from Mexico to Surinam.
It has been collected in flower in Panama in April. (Barro Colorado Island and
Madden Lake) and late October (Rio Pasiga), in each instance two widely
separated individuals blooming on the same day.
The presence of Paragonia-like pseudostipules with interpetiolar glandular
fields serves to distinguish sterile specimens of C. tetragonolobum. Trifid tendrils
are another good field mark.
A fallen fruit collected from a stream bed at Almirante, Bocas del Toro
Province, may belong to Ceratophytum. It is of the characteristic shape but much
narrower and only 13 cm long. It could belong to a new species of this genus.
CANAL ZONE: Barro Colorado Island, Croat 10212, 10808, 10879 (all MO), 14453, 14458
(both MO, SCZ), 14865 (SCZ), 14925 (MO). Boy Scout road near Madden Lake, Gentry
2054 (MO). Margarita, Gentry & Tyson 4831 (MO). Fort Sherman, Gentry & Tyson 4838
(MO). DARIEN: S of La Palma, Gentry 4292 (MO). Rio Tuira between Rios Paya and
Espave, Gentry 4398 (MO). Between Rios Cube and Punusa, Gentry 4453 (MO). El Real,
Gentry 4537 (MO). Rio Parasenico near Cerro Pirre, Gentry 4739 (MO). PANAMA: Between
El Llano and Rio Mamoni, Duke 5368 (GH, MO). Between Cafnitas and Bayano dam site,
Gentry & Tyson 1705 (MO). Rio Pasiga, Gentry 2195, 2273, 2320 (all MO). Near Icanti,
Gentry 2601 (MO). Near Rio Espave, Gentry 3699 (MO). S of Rio Bayano, in Cuna reserva-
tion, Gentry 3814, 3838 (both MO). SAN BLAS: Near Puerto Obaldia, Gentry 1525, 1569
(both MO).
7. CLYTOSTOMA
Clytostoma Miers ex Bur., Adansonia 8: 353. 1868. TYPE: C. callistegioides

(Cham.) Bur. ex Griseb.
Lianas, stems with 8 phloem arms in cross section; branchlets tetragonal to

subterete without interpetiolar glandular fields; pseudostipules variable, usually
a cluster of subulate-lanceolate cataphylls more or less resembling a minute
bromeliad. Leaves simple to 2-foliolate, often with a terminal simple tendril.
Inflorescence a few-flowered panicle or a one to few-flowered axillary or terminal
fascicle. Flowers with the calyx cupular to campanulate, truncate, usually more
or less 5-denticulate, variously sublepidote to subpuberulous; corolla white to
purple, tubular-infundibuliform, somewhat puberulous or glandular lepidote
outside; anthers glabrous, the thecae straight, divaricate; pollen grains ecolpate,
the exine finely reticulate; ovary ovoid-cylindric, minutely blunt-tuberculate
with thick one-celled glandular trichomes, the ovules 2 (-4) -seriate in each
locule. Fruit an ellipsoid or suborbicular capsule, the valves parallel to the
septum, woody, echinate with retrorse spines; seeds thick, corky, essentially
wingless.
Nine species in continental tropical America ranging from Mexico to Uruguay,

Argentina and Brazil.
1. Clytostoma binatum (Thunb.) Sandw., Recueil Trav. Bot. Neerl. 34: 235.
1937.-FIG. 7.
Bignonia binata Thunb., P1. Bras. 3: 35. 1821. TYPE: Brazil, Freyreiss s.n. (UPS, fide Sandw.).
B. callistegioides forma p Cham., Linnaea 7: 714. 1832. TYPE: Brazil, Sellow s.n. (NY).
B. noterophila Mart. ex DC., Prodr. 9: 148. 1845. TYPE: Brazil, Para, Martius s.n., 1819. (M).
B. umbellata DC., Prodr. 9: 148. 1845. TYPE: Brazil, Lhotsky s.n. (G-DC.).
B. purpurea Lodd. ex Hook. f., Bot. Mag. tab. 5800. 1769. TYPE: (not seen).
Adenocalymma ocositense Donn. Sm., Bot. Gaz. (Crawfordsville) 18: 209. 1893. TYPE: Guate-
mala, Quetzaltenango, J. D. Smith 2688 (GH, K, M, NY, US).
Clytostoma floridum Miers ex Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 154. 1896, pro syn.
C. noterophilum (Mart. ex DC.) Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 153. 1896.
C. purpureum (Lodd. ex Hook. f.) Rehder in Bailey, Standard Cycl. Hort, 2: 806 1914.
C. isthmicum Pittier, Contr. U.S. NatI. Herb. 18: 257, pl. 106. TYPE: Panama, Canal Zone,
Pittier 4008 (BM, GH, NY, US).
Petastoma laurifolium Kranzl., Fedde Repert. 17: 57. 1921. TYPE: Bolivia, junction of Rios
Beni and Madre de Dios, Rusby 1144, pro parte (B, K, MICH).
P. multiglandulosum Kranzl., Fedde Repert. 17: 58. 1921. TYPE: Bolivia, junction of Rios
Beni and Madre de Dios, Rusby 1145 (K).
P. ocositense (Donn. Sm.) Kranzl., Fedde Repert. 17: 61. 1935.
FIGuRE 7. Clytostoma binatum (Thunb.) Sandw.-A. Habit (X 1/2). [After Gentry

3781 (MO).]-B. Tendril (x 3/5). [After Gentry 5127 (MO).]-C. Node with pseudostipules
(X 3/5). [After von Wedel 788 (MO).]-D. Fruit (X 3/5). [After Gentry 3756 (MO).]-
E. Corolla split open ( X 3/5). -F. Ovary cross section ( x 12).-G. Ovary ( x 7). [After
Gentry 3740 (MO).]
.... .. ..
A! B
j~~~~~4
DD
Clytostoma elegans Standl., Carnegie Inst. Washington Publ. 461: 86. 1935. TYPE: British
Honduras, Rio Grande, Schipp 1127 (F, holotype; K, MO, US, isotypes).
C. ocositense (Donn. Sm.) Seib., Carnegie Inst. Washington Publ. 522: 413. 1940.
Liana to at least 5 cm in diameter, the bark gray, smooth; stem with 8 phloem
arms in cross section; branchlets subtetragonal, lepidote and slightly puberulous,
the nodes without interpetiolar glandular fields, pseudostipules usually resembling
small bromeliads, composed of numerous small acute bracts. Leaves 2-foliolate,
sometimes with a tendril; leaflets elliptic, acuminate, basally cuneate, 6.5-19 cm
long and 2.4-7.9 cm wide, membranaceous to chartaceous, secondary veins 5-9
on each side, plate-shaped glands along midvein especially near base, lepidote
on both sides, otherwise glabrous beneath, minutely puberulous at the base of
the midvein above, drying greenish above, olive beneath; tendril simple, 5-12
cm long; petiole and petiolules minutely puberulous on upper surface, petiolules
0.5-2.1 cm long, slightly swollen and woody at base and apex, petiole 0.6-3.1
cm long. Inflorescence an axillary or terminal fascicle, the pedicels scabrous.
Flowers aromatic, calyx cupular, subtruncate or 5-denticulate with cusps to
1 mm long, 4-7 mm wide, subpuberulous to scabrous, usually with plate-shaped
glands in the upper half; corolla magenta to almost white outside and on the
lobes inside, the throat white inside with ca. 11 thin longitudinal magenta nectar
guide lines on the abaxial side, usually with a deeper magenta spot at the base
of the adaxial sinus, tubular-infundibuliform, 5-8.5 cm long and 1.2-2.4 cm wide
at the mouth, the tube 3.7-5.8 cm long, the lobes 1.2-2.2 cm long, puberulous
outside with simple or sometimes branched trichomes and slightly glandular-
lepidote on the lobes, inside more or less glandular-lepidote on the lobes, mostly
glabrous in the tube with plate-shaped glands scattered adaxially, glandular-
pubescent with simple trichomes at and below the level of stamen insertion;
stamens didynamous, the anther thecae divaricate, 2-3 mm long, the longer
filaments 1.4-1.9 cm long, the shorter filaments 1.0-1.3 cm long, the staminode
3-5 mm long, inserted 4-6 mm from base of corolla tube; pistil 2.4-2.7 cm long,
the ovary rounded short-cylindric, 2 cm long and 1.5-2 cm wide, 1-1.5 cm
thick, densely covered with thick subspherical glandular trichomes, the ovules
2'seriate in each locule; disc obsolete. Capsule elliptic to suborbicular, heavily
echinate with the tips of the enations recurved and often split, 4.5-6.5 cm long
and 3.2-4.5 cm wide; seeds thick, corky, 1.2-1.9 cm long and 1.9-2.4 cm wide,
the wings brown, woody, almost obsolete.
Ecologically restricted mainly to swampy or lakeshore habitats in the tropical

wet forest, this species occurs less commonly in moister parts of the tropical
moist forest, e.g. a single plant known on Barro Colorado Island. Its swampy
habitat precludes abundant herbarium representation despite its relatively high
local abundance. It ranges from Mexico to Brazil. This species flowers sporad-
ically throughout the year but is most commonly collected in late wet season
(September to November). The seeds are water dispersed as befits its swamp
habitat.
Clytostoma binatum resembles Cydista aequinoctialis when sterile and is
best separated by its bromeliad-like pseudostipules, which are unfortunately
often absent. The 4 darker lines on a subterete to subtetragonal twig are char-
acteristic as is the elliptic leaf shape.
BOCAS DEL TORO: Region of Almirante, Cooper 198 (F). Banks of Changuinola River
near Bar Mouth, Dunlap 418 (BM, F, GH, US). 10-15 mi. inland from mouth of Changuinola
River, Lewis et al. 866. Vicinity of Chiriqui Lagoon, von Wedel 9, 788 (both GH, MO), 1314
(US), 1345 (GH, MO). Water Valley, von Wedel 795 (GH, MO, US), 1660 (MO). Fish
Creek Mountains, von Wedel 2360 (GH, MO, US). Western River, von Wedel 2700 (MO).
CANAL ZONE: Barro Colorado Island, Croat 5727 (MO), 13963 (MO, SCZ). S of Margarita,
Gentry 5127 (MO). Bohio Station of Panama Railroad, Hayes 144 (K). Panama Station,
Panama Railroad, Hayes 412 (BM). Along Rio Trinidad, Pittier 4008 (BM, GH, NY, US).
COLON: E of Rio Piedras, Gentry 2190 (MO). Peluca Hydrographic Station, Steyermark &
Allen 17224 (BM, K, MO). DARIEN: El Real, Duke 5055 (GGH, MO, PMA, US); Gentry
4572 (MO). Near Pinogana, Pittier 6526 (US). PANAMA: Rio Espave, Gentry 3740, 3756,
3781 (all MO). Between Juan Mina and Vigia, Pittier 3452 (BM, NY, US).
8. CRESCENTIA
Crescentia L., Sp. PI. 2: 626. 1753; Gen. PI., ed. 5. 274. 1754. TYPE: C. cujete L.
Pteromischus Pich6n, Bull. Soc. Bot. France 92: 227. 1946. TYPE: P. alatus (H.B.K.) Pichon
= Crescentia alata H.B.K.
Small to medium-sized trees, branching rachitic, crown open. Leaves simple

or 3-foliolate, borne on thick twigs in alternate fascicles resulting from the
condensation of short-shoots. Inflorescence of 1-2 cauliflorous flowers arising
from nodes on the trunk and older branches. Flowers with the calyx large,
usually bilabiately split; corolla off-white, usually with maroon penciling, espe-
cially on lobes and on the tube inside, the lobes deltoid, acuminate; stamens
subexserted, the anthers glabrous, the thecae thick, somewhat divergent; pollen
grains single, 3-colpate, the exine microreticulate; ovary ovoid-elliptic, lepidote,
1-locular, the ovules multi-seriate on 4 parietal placentae. Fruit a pepo or cala-
bash, large, more or less spherical, indehiscent with a hard woody shell, pulpy
inside; seeds small, less than 8 mm long and 9 mm wide, flat, not winged,
embedded in the pulp.
Five species in tropical America ranging from Mexico and the West Indies
to Amazonian Brazil, also cultivated through most of the tropics. Probably not
native to Panama.
a. Leaves all simple, more than 3.4 cm long; fruit more than 13 cm in diameter, widely
cultivated --2. C. cujete
aa. Leaves 3-foliolate and simple in each fascicle, the simple leaves less than 3.2 cm long;
fruit less than 10 cm in diameter, rarely cultivated -1. C. alata
1. Crescentia alata H.B.K., Nov. Gen. Sp. PI. 3: 158. 1819. TYPE: Mexico,
Guerrero, Humboldt & Bonpland 3858 (P).-FiG. 8F-G.
C. trifolia Blanco, Fl. Filip., ed. 1. 489. 1837. TYPE: (not seen).
Parmentiera alata (H.B.K.) Miers, Trans. Linn. Soc. 26: 166. 1868.
Crescentia ternata Sesse & Moc., La Naturaleza, ser. 2. 1 (append.): 94. 1889. TYPE: Mexico
(not seen; photo of illustration at MO).
Pteromischus alatus (H.B.K.) Pichon, Bull. Soc. Bot. France 92: 227. 1945.
Small tree to 25 cm d.b.h. and 6 m tall, the branches crooked, the crown
open; branchlets essentially lacking, the smaller branches thick, subterete, with
~~~~~~B
G
FIGUpE 8. Crescentia-A-E. C. cujete L.-A. Habit (X 2/,5)-B. Fruit (x 2%). [After

,Gentry 352, Costa Rica (MO)J.-C. Ovary cross section (X 4) .-D. Flower in dorsal view
(X 2/,5) .-E. Corolla in ventral view (X 2/s5). [From miscellaneous fresh material.1-F-G.
C. alata H.B.K.-F. Habit (x 2/5)D [After roat 273, Costa Rica (MO).-G. Fruit
(X 2/5). [After Gentry 5043, Mexico (MO).]
alternate condensed, short-shoot projections each bearing a fascicle of lea

from its center. Leaves both compound and simple from each fascicle, th
pound leaves 3-foliolate with sessile oblanceolate leaflets 1.0-4.5 cm long and
0.3-1.2 cm wide, round-tipped or slightly emarginate, lepidote but otherwise
glabrous, rigid-chartaceous, the petiole winged, 2.7-11.5 cm long and 3-15 mm
wide, lepidote, oblanceolate, rigid chartaceous as the leaflets, the simple leaves
much smaller, 1.9-3.2 cm long and 0.5-1.0 cm wide, similar to single leaflets or
petioles of compound leaves. Inflorescence one or 2 cauliflorous flowers borne

on larger branches or trunk, pedicels lepidote, 0.5-0.8 cm long. Flowers with a
musky odor, calyx split nearly to the base into two subequal lobes, each 1.4-
1.9 cm long and 0.7-1.4 cm wide, mostly glabrous, slightly lepidote at the base,
usually with sunken plate-shaped glands; corolla off-white, tubular-campanulate
with a horizontal pleat midway across the lower side of throat, fleshy, 4.0-6.5
cm long and 1.8-2.5 cm wide at the mouth of tube, the tube 2.8-4.2 cm long,
the lobes 1-1.5 cm long, more or less triangular and fused at the bases, mostly
glabrous, sparsely glandular-lepidote outside; stamens subexserted, the anther
thecae ca. 2 mm thick, only partially divergent, 5-7 mm long, the anterior
filaments 2.1-3.4 cm long, inserted 0.8-1.7 cm from the base of the tube, the
posterior filaments 2.3-3.8 cm long, inserted 1.1-1.5 cm from the base of tube,
the staminode 3-4 mm long, inserted 0.9-1.0 cm from the base of the tube;
pistil 4.3-5.8 cm long, the ovary rounded-conical, lepidote, 5-6 mm long and
4-5 mm wide, the ovules multi-seriate on 4 parietal placentae; disc pulvinate,
2-3 mm long and 0.9-1.4 mm wide. Fruit a pepo or calabash, more or less
spherical, 7-10 cm in diameter; seeds small, thin, wingless, 6-7 mm long and
7-9 mm wide, scattered through the pulp of the fruit.
A plant of the tropical dry forest from Mexico to Guanacaste, Costa Rica,
C. alata is much confused with its better known congener C. cujete. It is
actually the commoner of the two in the wild and is in fact a characteristic
tree of Pacific slope dry forest savannas, its short twisted stature and open rachitic
branching making it physiognomically one of the most conspicuous elements of
this vegetation. It flowers irregularly throughout the year. The fruit is similar
to that of C. cujete but smaller. Presumably the fruit is water dispersed and
the seeds dispersed by large mammals. I have encountered young trees of this
species growing far from the usual savanna habitat on gravel bars where they
must have grown from water-carried fruits.
Apparently this tree is neither utilized nor cultivated by the campesinos,
who prefer its larger-fruited relative; there are reports, however, of the fruits
being used as cups. The 3 leaflets and petiole together form a kind of cross
and the tree has acquired some religious significance from this resemblance.
The 3-foliolate leaf of this species is the best mark distinguishing it from
the simple-leaved C. cuiete. Smaller fruit size is also a useful character. The
two species hybridize and two much-studied trees along the Pan-American
Highway at Hacienda la Pacifica near Cafias, Costa Rica, are apparent hybrids.
These trees have the small fruits of C. alata but the simple leaves of C. cujete,
mad occasional branches bear 3-foliolate leaves. Trees of both putative parent
species occur in the immediate vicinity.
CHIIQUI (VERAGUAS): David, cultivated Seemann 1130 (K).
2. Crescentia cujete L., Sp. P1., ed. 1. 2: 626. 1753. TYPE: Specimen without
information (LINN 779.1) .-FIG. 8A-E.
C. acuminata H.B.K., Nov. Gen. Sp. P1. 3: 157. 1819. TYPE: Cuba, Havana, Humboldt &
Bonpland 1300 (P).
C. arborea Raf., Sylva Tell. 81. 1838. TYPE: (not seen).
C. cuneifolia Gardn., Jour. Bot. (Hook.) 2: 422. 1840. TYPE:

C. angustifolia Willd. ex Seem., Trans. Linn. Soc., ser. 1. 23: 20, 1862, pro syn.
C. fasciculata Miers, Trans. Linn. Soc., ser. 1. 26: 171. 1868. TYPE: St. Thomas, Oersted s.n.
(not seen).
C. plectantha Miers, Trans. Linn. Soc., ser. 1. 26: 170. 1868. TYPE: Brazil, Rio de Janeiro
(not seen).
C. spathulata Miers, Trans. Linn. Soc., ser. 1. 26: 173. 1868. TYPE: Ecuador, Sinclair s.n.
C. cujete var. puberula Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 403. 1897. SYNTYPES:
Brazil, Goyaz, Dombey s.n. (not seen); Peru, Ruiz s.n. (not seen).
Tree to 10 m tall and 30 cm d.b.h., the branches usually crooked, the crown
open; branchlets mostly lacking, smaller branches thick, subterete, with alter-
nate short-shoot projections each bearing a fascicle of leaves from its center.
Leaves of various sizes within each fascicle, simple, obovate, the tip obtuse to
acute, the base attenuate, petiole lacking, 3.4-26 cm long and 1.0-7.6 cm wide,
chartaceous to rigid-chartaceous, secondary veins 5-14 on a side, the midvein
raised above; lepidote above and beneath, otherwise glabrous above, beneath
pubescent along midvein with simple and forked trichomes, plate-shaped glands
at base of blade beneath, drying grayish-olive. Inflorescence one or 2 cauliflorous
flowers borne on larger branches or trunk, the pedicels lepidote, 1.5 cm long.
Flowers with a musty odor, calyx bilabiately split to the base, each lobe 1.8-2.6 cm
long and 1.3-2.4 cm wide, mostly glabrous with plate-shaped glands on the
upper half of lobes, slightly lepidote at the base; corolla off-white to yellowish-
white with purplish venation on the lobes and purplish lines on the tube out-
side, tubular with a transverse fold midway across the lower side of the throat,
fleshy, 4.1-7.4 cm long and 3.1-4.5 cm wide at the mouth of the tube, the tube
2.8-4.5 cm long, the lobes triangular with the apex extended as a narrow point,
2.5-3.1 cm long, sparsely lepidote or stalked-lepidote on the tube outside and
near the mouth of tube inside, more densely so at the level of stamen insertion,
papillate-glandular on the lobes outside and sparsely so inside; stamens sub-
exserted, the anther thecae thick, partially divergent, 5-8 mm long and 2.5-3.5
mm wide, the anterior filaments 2.8-3.2 cm long and inserted 6-11 mm from
the base of the tube, the posterior filaments 2.8&-3.3 cm long, inserted 7-15 mm
from the base of the tube, the staminode 1-3 mm long, inserted 5-11 mm from
the base of the tube; pistil 4.0-4.9 cm long, the ovary rounded conical, 5-7 mm
long and 4-7 mm wide, 3-6 mm thick, lepidote, the ovules multi-seriate on 4
placentae; disc annular-pulvinate, 3-4 mm long and 8-11 mm wide. Fruit a
pepo or calabash, spherical to ovoid-elliptic, 13-20 cm in diameter to 30 cm
long, the thin hard shell smooth, lepidote-punctate; seeds small, thin, wingless,
7-8 mm long and 4-6 mm wide, scattered through the pulp of the fruit.
Crescentia cujete is extensively cultivated through most of tropical America.

Although native to Central America, such wide cultivation makes its original
range impossible to trace. Apparently wild-growing trees are sometimes encoun-
tered, but even these may have descended from cultivated trees. It flowers
irregularly throughout the year. The flowers are bat-pollinated. The fruit
floats and can sometimes be found washed up on beaches indicating a possible
means of dispersal.
The hard shell of the fruit is widely used as a utensil for dipping or holding
liquids. They are especially conspicuous as bailers for piraguas, and nearly
every piragua in Panama's Darien is equipped with its piece of Crescentia
exocarp. The pulp is edible but insipid-tasting and little used for food.
BOCAS DEL TORO: Near Almirante, Gentry 2807 (MO). Mariano Creek, von Wedel 2894
(GH, MO). CANAL ZONE: Near Los Rios, Gentry 3690 (MO). Gatun Station, Hayes 94
(BM). Balboa Heights, Sharp s.n. (SCZ). Chiva Chiva Trail, Tyson & Blum 3565 (SCZ).
CHIRIQUI: Without locality, Wagoner s.n. (M). Progreso, Cooper & Slater 288 (US). DARIEN:
Village of Paya, Gentry 4397 (MO). El Real, Gentry 4505 (MO). LOS SANTOS: S of Pedasi,
Stimson 5293 (MO, SCZ). PANAMA: Cafiitas, Gentry 1405 (MO). Rio Maestro, Gentry 2245
(MO). Rio Piragua (Tabardi ), Gentry 2523 (MO). Bella Vista, Standley 25343 (US).
Taboga Island, Standley 27885 (US). Between Matias Hernandes and Juan Diaz, Standley
31974 (US). SAN BLAS: Mountains above Puerto Obaldia, Gentry 1478 (MO).
9. CUSPIDARIA
Cuspidaria DC., Rev. Bign. (Bibl. Univ. Geneve) 8. 1838, nom. cons. -TYPE: B.
pterocarpa Cham. = Cuspidaria pterocarpa (Cham.) DC.
Lochmocydia Mart. ex DC., Prodr. 9: 177. 1845, pro syn.

Saldanhaea Bur., Adansonia 8: 354. 1868, non Saldanha Vell., 1825. TYPE: S. lateriflora
(DC.) Bur.
Blepharitheca Pichon, Bull. Soc. Bot. France 92: 224. 1945. TYPE: B. floribunda (DC.)
Pichon = Cuspidaria floribunda (DC.) A. Gentry.
Lianas, branchlets terete, with or without interpetiolar glandular fields;

pseudostipules small or lacking. Leaves simple to 5-foliolate, sometimes with a
simple tendril. Inflorescence a terminal panicle, often on a short lateral branch.
Flowers with the calyx cupular, 5-denticulate, the teeth usually long and con-
spicuous; corolla lavender, tubular-infundibuliform, puberulous outside; anthers
pubescent or glabrous, the thecae divaricate, bent at the middle and strongly
reflexed forward; pollen grains in tetrads; ovary linear-cylindric, lepidote, the
ovules 2-4-seriate in each locule. Fruit a linear or oblong capsule, the valves
more or less woody though sometimes quite thin, parallel to the septum, with
conspicuously raised more or less winged lateral margins or with 2 submarginal
ridges, in either case with a more or less evident central furrow on each valve;
seeds thin, bialate, the wings hyaline-membranaceous.
About 10 species in continental tropical America from eastern Panama to

Argentina, mostly in Brazil.
Cuspidaria floribunda (DC.) A. Gentry, Brittonia 25: 232. 1973.-FIG. 9:
Adenocalymma floribundum DC., Prodr. 9: 201. 1845, non Bignonia floribunda H.B.K.,
Arrabidaea floribunda (H.B.K.) Loes. TYPE: Bolivia, Santa Cruz, d'Orbigny 533 (P).
Arrabidaea rubrinervis Miers, Proc. Roy. Hort. Soc. 3: 192. 1863. TYPE: Brazil, So Paulo,
Wier 178 (BM, K).
A. subfastigiata Bur. ex Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 70. 1896. TYPE: Brazil,
Minas Gerais, Warming s.n. (BR).
A. macrocarpa 0. Kuntze, Rev. Gen. PI. 3(2): 242. 1898. TYPE: Bolivia, Velasco, Kuntze
s.n. (NY).
Blepharitheca floribunda (DC.) Pichon, Bull. Soc. Bot. France 92: 224. 1945.
Saldanhaea subfastigiata (Bur.) J. C. Comes, Rev. Bras. Biol. 9: 51. 1951.
S. floribunda (DC.) Sandw., Kew Bull. 22: 408. 1968.
Liana without interpetiolar glandular fields or noticeable pseudostipules;
.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ ....
FIGURE 9. Cuspidaria floribunda (DC.) A. Gentry. A. Habit (X 1/2). B. Ovary

and disc (x 6).-C. Anther (x 6). [After Duke 5386 (MO).]
branchlets subterete, flattened at nodes, sometimes with conspicuous V-shaped

interpetiolar ridges, longitudinally striate when dried, noticeably lenticellate,
appressed puberulous, sometimes hollow at the very center. Leaves 2-foliolate,
sometimes with a tendril; leaflets more or less elliptic, acute, the base broadly
cuneate to rounded, 8.2-12.2 cm long and 4.4-8.6 cm wide, membranaceous to

chartaceous, secondary veins 4-5 on a side, pubescent with scattered simple
trichomes above, beneath with simple trichomes at least on the nerves, some-
times glabrescent, with axillary recesses beneath, these fringed with short
trichomes, usually with a few large plate-shaped glands along lower part of
midvein, drying olive to reddish-brown above, greenish-brown to olive with
reddish veins and veinlets beneath; tendril simple, 10-14 cm long; petiolules
1.2-2.8 cm long, petiole 2.0-2.9 cm long, short-puberulous. Inflorescence a
terminal panicle, the branches appressed short-puberulous. Flowers with the
calyx cupular, subtruncate, 5-6 mm long and 4-5 mm wide, 5-denticulate with
teeth ca. 0.5 mm long, puberulous, usually drying pale pinkish; corolla magenta
(Stern et al.), tubular-infundibuliform, 3.6-4.5 cm long and 0.8-1.2 cm wide
on tube inside and on the lobes, tube inside mostly glabrous, pubescent at the
level of stamen insertion; stamens didynamous, the anther thecae divaricate,
glabrous or pilose, usually reflexed forward from the base, 2.0-2.5 mm long,
the connective apiculate ca. 0.5 mm, the longer filaments 1.6-1.7 cm long, the
shorter filaments 1.1-1.2 cm long, the staminode 3-4 mm long, inserted 9-10
mm from base of corolla tube; pistil 2.3-2.4 cm long, the ovary cylindrical,
densely minutely lepidote, 1.5-2 mm long and 1 mm wide, the ovules 4-seriate;
disc pulvinate, 1 mm long and 1 mm wide. Capsule elongate-linear, 50-43 cm
long and 0.9-1.0 cm wide, the valves somewhat compressed, with two submedian
ridges and an inconspicuous median furrow between them, minutely warty but
essentially glabrous; seeds thin, bialate, 0.8-0.9 cm long and 3-3.7 cm wide,
body brown, wings hyaline membranaceous at the tips, brown at the base,
indistinctly demarcated.
A liana of the tropical wet forest, this species was previously known only
from Peru, Bolivia, and southern Brazil. The Panamanian collections extend its
range northward and westward into the eastern Darien. Cuspidaria floribunda
apparently flowers during the wet season from Jurne to August.
This is one of the two Panamanian species of Bignoniaceae of which I
have not seen living plants. It can be recognized in the herbarium by the
color of its dried leaves: greenish-brown beneath with reddish brown veins
and veinlets and reddish brown above. Its dried flowers superficially resemble
those of Xylophragma seemannianum.
DARIEN: Along Rio Tuira between Rio Pucro and Rio Punusa, Duke 5386 (MO). Near
Cana, Stern et al. 488 (GH, MO, US).
10. CYDISTA
Cydista Miers, Proc. Roy. Hort. Soc. 3: 191. 1863. TYPE: C. aequinoctialis (L.)
Miers.
Levya Bur. ex Baill., Hist. Pl. 10: 28. 1888. TYPE: L. nicaraguensis Bur. ex Baill. = Cydista
aequinoctialis var. hirtella (Benth.) A. Gentry.
Lianas, stems with 8 phloem arms in cross section; branchlets terete to

tetragonal, without interpetiolar glandular fields; pseudostipules foliaceous or
absent. Leaves simple or 2-foliolate, sometimes with a simple tendril. Inflor-

escence an axillary or terminal raceme or few-flowered panicle. Flowers with
the calyx cupular, truncate or irregularly and shallowly bilabiate, lepidote to
puberulent or sub-hirsute; corolla white to magenta, tubular-infundibuliform,
somewhat lepidote and sometimes inconspicuously puberulent outside; anthers
glabrous, the thecae straight, divaricate; pollen grains single, acolpate, the exine
finely reticulate; ovary narrowly cylindric, lepidote or somewhat puberulous,
the ovules 2-seriate in each locule. Fruit a compressed, linear capsule, the valves
parallel to the septum, the midline inconspicuous; seeds flat, bialate, the wings
thin, brown, sometimes sub-hyaline at the extreme tip, not differentiated from
the seed body.
Four species ranging from Mexico and the West Indies to Brazil.
a. Pseudostipules persistent, foliaceous, conspicuous; branchlets acutely tetragonal; capsule

valves thin, uniformly flattened -2. C. diversifolia
aa. Pseudostipules absent or inconspicuously foliaceous but then early caducous and present
only on very young branchlets; branchlets not acutely tetragonal; capsule valves with
margins raised or with submarginal ridges.
b. Calyx shallowly 2-3-labiate; branchlets terete; leaves simple or 2-foliolate, the leaf or
leaflets conspicuously 3-veined from the base, beneath with glandular fields only in
the axils of the basal pair of nerves; capsule with 2 parallel submarginal ridges
--------------------------------------------------------------------3. C. heterophylla
bb. Calyx more or less truncate; branchlets terete to tetragonal; leaves 2-foliolate (except
on seedlings), the leaflets more or less pinnately veined from the base, beneath with
glandular fields in the axils of many secondary nerves; capsule with only the margins
somewhat raised -1. C. aequinoctialis
1. Cydista aequinoctialis (L.) Miers, Proc. Roy. Hort. Soc. 3: 191. 1863.
Bignonia aequinoctalis L., Sp. P1., ed. 1. 2: 623. 1753. TYPE: (not seen).
B. spectabilis Vahl, Symb. Bot. 80. 1794. TYPE: collector unknown "ex India" (C, P-JU).
B. villosa Vahl, Eclog. Amer. 2: 44. 1798, pro parte. TYPE: Colombia, Santa Marta, Von Rohr
21 (C, flower only).
B. picta H.B.K., Nov. Gen. Sp. P1. 3: 136. 1819. TYPE: Venezuela, Orinoco, Humboldt &
Bonpland 1078 (P).
B. hostmannii E. Meyer, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 779.
1825. TYPE: Martinique, Sieber s.n. (M, W).
B. variabilis Sieber ex E. Meyer, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur.
12: 779. 1825, pro syn., non Jacq.
B. sarmentosa Bertol., Fl. Guatimal. 25. 1840. TYPE: Guatemala, Escuintla, Velasquez s.n.
(BOLO, fide Sandw.).
B. incarnata Aubl. sec. Splitg., Tijdschr. Natuurl. Gesch. Physiol. 9: 7. 1842, non Aubl.
B. sarmentosa Bertol. var. hirtella Benth., Bot. Voy. Sulphur 128. 1844. TYPE: Nicaragua,
Realejo, Hinds s.n. (K).
B. nitidissima DC., Prodr. 9: 160. 1845. TYPE: Venezuela, Caracas, Vargas 244 (G-DC).
Temnocydia aequinoctialis (L.) Mart. ex DC., Prodr. 9: 155. 1845, pro syn.
T. spectabilis (Vahl) Mart. ex DC., Prodr. 9: 155. 1845, pro syn.
Cydista amoena Miers, Proc. Roy. Hort. Soc. 3: 191. 1863. TYPE: San Domingo [Hispaniola?],
Jussieu s.n. (not seen, fide Knuth, Fedde Repert. Beih. 43: 636. 1928).
C. incarnata sensu Miers, Proc. Roy. Hort. Soc. 3: 191. 1863, non Aubl.
C. sarmentosa (Bertol.) Miers, Proc. Roy. Hort. Soc. 3: 191. 1863.
C. seemannii Miers, Proc. Roy. Hort. Soc. 3: 192. 1863, nomen nudum. TYPE: Panama,
Seemann 397 (BM).
C. spectabilis (Vahl) Miers, Proc. Roy. Hort. Soc. 3: 191. 1863.
Levya nicaraguensis Bur. ex Baill., Hist. P1. 10: 29. 1888. TYPE: Nicaragua, Levy 38 (F,
fragments; K, fragments; P).
Arrabidaea guatemalensis K. Schum. & Loes., Bot, Jahrb. Syst. 23: 129. 1896. TYPE: Guate-
mala, Bernoulli & Cario 2056 (K).
A. pseudochica Krdnzl., Fedde Repert. 17: 19. 1921. TYPE: Mexico, El Platano, Langlasse
137 (K).
Anemopaegma tonduzianum Kranzl., Fedde Repert. 17: 116. 1921. TYPE: Costa Rica,
Guanacaste, Tonduz 13912 (K, P). .
Cydista pubescens Blake, Contr. U. S. Natl. Herb. 24: 23. 1922. TYPE: Honduras, Copain,
Pittier 8488 (US).
Arrabidaea isthmica Standl., Jour. Washington Acad. Sci. 15: 461. 1925, pro parte. TYPE:
Panama, Canal Zone, Pittier 2576 (leaves only) (US, holotype; GH, MO, NY, isotypes).
Liana to at least 8 cm in diameter, the bark gray to brownish gray, fibrous

with longitudinal striation; stem with 8 phloem arms in cross section; branchlets
subtetragonal to strongly tetragonal, usually drying dark, glabrous to pubescent,
the nodes without interpetiolar glandular fields, sometimes with a ridge between
opposite petioles; pseudostipules usually lacking or inconspicuous, occasionally
thick-foliaceous but early deciduous on young twigs. Leaves 2-foliolate, some-
times with a tendril; leaflets more or less ovate, acuminate to obtuse, broadly
cuneate to truncate, 5.4-16.2 cm long and 2.2-9.2 cm wide, membranaceous to
chartaceous, secondary veins (3-)4-7(-8), lepidote above and beneath, variously
pubescent with simple trichomes, especially along main veins beneath and with
glandular fields in the axils of many secondary nerves, drying olive to brownish
olive or reddish, shiny; tendril simple, 4-21 cm long; petiolules and petioles
lepidote to pubescent, the petiole apex normally puberulent even in otherwise
glabrous leaves, 0.8-3.9 cm long, petiole 0.9-4.5 cm long. Inflorescence a termi-
nal or axillary panicle, its branches lepidote to simple pubescent. Flowers sweetly
aromatic, calyx cupular, truncate to weakly 5-lobed or bilabiate, 4-10 cm long
and 4-9 cm wide, lepidote to simple pubescent, sometimes with glands; corolla
white or lavender with 13 maroon lines at the base of the lobes, 3 extending into
the upper throat, the throat opening yellow, tubular-campanulate, 2.5-7.6 cm
long and 0.6-2.2 cm wide at the mouth, the tube 1.9-5 cm long, lobes 0.6-2.6 cm
long, glandular lepidote outside, inside simple puberulous with short, thick, 1-3-
celled trichomes on the lobes, sparsely glandular-lepidote on the tube and simple-
pubescent with multicellular trichomes from the level of stamen insertion to the
base of the corolla tube; stamens didynamous, the anther thecae divaricate, 3-4
mm long, the longer filaments 1.1-1.7 cm long, the shorter filaments 0.7-1.4 cm
long, the staminode 3.5 mm long, inserted 4-5 mm from the base of the corolla
tube; pistil 1.6-3.2 cm long, the ovary narrowly cylindrical, 2-3 mm long and
1 mm wide, lepidote, the ovules 2-seriate in each locule; disc obsolete. Capsule
linear, round-tipped, flattened, 21-43 cm long and 1.4-2.4 cm wide, the margins
rounded, slightly raised, the surface longitudinally finely wrinkled, lepidote,
drying blackish; seeds 1.6-1.9 cm long and 5.0-7.3 cm wide, the wings mem-
branaceous, brown, not demarcated from the seed body.
Most abundant in tropical moist forest where it is one of the commonest

species of Bignoniaceae, C. aequinoctialis also occurs in tropical dry forest and
premontane moist forest. It is a highly polymorphic and widespread species
which ranges from Mexico and the West Indies to Brazil. It flowers and fruits
throughout the year.
Field characteristics include the more or less tetragonal twigs, glandular fields
in the axils of most main lateral nerves beneath, lack of pseudostipules (except
occasionally on very young and actively growing branches), and more or less
ovate leaflets (not strictly elliptic as in Clytostoma).
The pubescent-leaved form often referred to as C. sarmentosa or C. pubescens
should be known as C. aequinoctialis var. hirtella (Benth.) A. Gentry.
BOCAS DEL TORO: SW of Bocas at Macaw Hills, von Wedel 554 (MO). Fish Creek low-
lands, von Wedel 2388 (GH, MO, US). Fish Creek hills, von Wedel 2415, 2449 (both GH,
MO, US), 2451 (CGH, US). Isla Colon, von Wedel 2827 (CGH, MO, US). CANAL ZONE:
Barro Colorado Island, Aviles X-1, X-8 (both F, MO); Bailey & Bailey 132, 610 (both F);
Bangham 422 (A), 605 (A, F); Blum & Dwyer 2130 (MO, SCZ); Croat 4764 (MO), 5119,
5162, 5208 (all MO, SCZ), 5439 (MO), 5589, 5611 (both MO, SCZ), 5700, 6143 (both MO),
6155, 6169 (both MO, SCZ), 7176 (MO), 8310, 8552 (both MO, NY, SCZ), 8755, 9514,
9566 (all MO, SCZ), 10324 (MO), 11097 (MO, SCZ), 11610 (MO), 12268 (NY), 12645,
12957 (both MO), 13487 (MO, SCZ); Dodge 3507 (GH, US); Ebinger 288 (MO, US);
Foster 933 (PMA); Hayden 20 (MO); Kenoyer 533 (US); Knight 69-87 (MO, SCZ);
Shattuck 784 (F), 1040, 1147 (both F, MO). Mojinga swamp near mouth of Rio Chagres,
Allen 858 (F, GH, K, MO, NY, US). Near Gamboa, Allen 1964 (F, GH, MO, NY, US). Near
Madden Dam, Alston 8865, pro parte (BM), 8865 (K). Between Mt. Hope and Santa Rita
trail, Cowell 78 (NY). Mindi, Cowell 189 (NY). Summit Garden, Croat 12796 (MO). Near
beach at Ft. Kobbe, Duke 4696 (MO). Road C-21, Duke 5777 (MO). France Field U. S.
Army Tropic Test Mine Emplacement, Dwyer 8528 (MO). Pipeline Road, Gentry 1583 (MO).
Near Venado beach, Gentry 1604 (MO). Boy Scout road near Madden Lake, Gentry 2055
(MO). Pipeline Road, Gentry & Dwyer 3390 (MO). Near Margarita, Gentry & Dwyer 4327
(MO). Road to Galeta Lab, Gentry & Tyson 4854 (MO). Near Gatuln, Goldman 1861 (US).
Panama Station, Panama Railroad, Hayes 412, pro parte (BM, K). Gatuin, Hayes 587, 590
(both NY). Mt. Hope, Heriberto 40 (US). Summit Gardens, Higgins 203, 207 (both MO,
US). Road C-10, Lazor 5460 (SCZ). Near Fort Randolph, Maxon & Harvey 6533 (US).
Along Rio Indio de Gatuin, Pittier 2773 (BM, GH, NY, US). Las Sabanas, Pittier 6762, pro
parte (BM, F, GH, MO, NY, US). Near Fort Randolph, Standley 28628, 28642 (both US).
Vicinity of Fort Sherman, Standley 31183 (MO, US). Near Curundu, Tyson 1089 (MO,
SCZ). CHIRIQUI: West of San Bartolo Limite near Costa Rican border, Croat 22192, 22159
(MO). Vicinity of Puerto Armuelles, Woodson & Schery 811A (MO). COCLE: Vicinity of
El Valle de Anton, Allen 1780 (F, GH, MO, NY, US), 2503 (US); Bro. Paul s.n. (US).
Vicinity of Penonome, Williams 228 (NY, US). COLON: Miguel de la Borda, Croat 9846
(MO). Nuevo Chagres, Lewis et al. 1865 (GH, MO). Near Fort San Lorenzo, Quistgaard
58 (PMA). Without locality, Carrasquilla & Taymes 140. (MO). DARIEN: 1-5 mi. down-
stream from El Real, Duke 4922 (MO). Along Rio Pirre, Duke 4978 (GH, MO, US). Along
Rio Chucunaque between El Real and Rio Canalones, Duke 4989 (GH, MO). Santa Fe,
Duke 14256 (K, MO). Punto Sabanas opposite La Palma, Gentry 3957 (MO). S of
Jaque, Gentry 4116 (MO). Rio Tuira between Rios Paya and Aspave, Gentry 4410 (MO).
El Real, Gentry 4564, 4580 (both MO); Stern et al. 755 (GH, MO, US). Rio
Chucunaque above confluence with Rio Tuquesa, Stern et al. 882 (GH, K, MO, US).
HERRERA: E of Las Minas, Gentry 3138, 3144 (MO). S of Los Pozos, Tyson 2683 (MO,
SCZ). LOS SANTOS: Punta Mala, D'Arcy & Croat 4215 (MO). PANAMA: Vicinity of
Bejuco, Allen 2458 (US). Cerro Campana, Allen 2641 (MO). Vicinity of La Chorrera,
Allen 3646 (GH, MO, NY). Cerca de Rio Tapia como 2 km de la carretera de Tocumen,
Correa & Escobar 1788 (MO). Between Capira and Chorrera, Dodge 10715 (F, GH, MO).
Between El Llano and Rio Mamoni, Duke 5531 (MO, US). Tocumen Agronomy School
Farm, Dwyer 4045 (MO, US), 5130 (MO). San Jose Island, Erlanson 485 (GH, NY, US).
Along road to Chepo, Garner 4 (MO). Between El Llano and Caiiitas, Gentry 1414 (MO).
Mouth of Rio Pasiga, Gentry 2198 (MO). Rio Pasiga, Gentry 2267 (MO). Rio Agua Clara,
Gentry 2629 (MO). Near Rio Espave, Gentry 3711 (MO). 0-4 mi. from Rio Bayano crossing
on road to Santa Fe, Gentry 3859 (MO). W of Chepo, Gentry 4970 (MO); Gentry & Tyson
1642 (MO). Dam site on Rio Bayano, Gentry & Tyson 1663D (MO). Las Sabanas, Paul 236
(US). Near Panama City, Seemann 397 (K). Taboga Island, Standley 27944 (US).
Vicinity of Campana, Woodson et al. 1309 (A, MO, NY). Between Pacora and Chepo,
Woodson et al. 1651 (A, MO, NY). Vicinity of Campana, Woodson et al. 1679 (A, MO, NY).
SAN BLAS: Mountains above Puerto Obaldia, Gentry 1477 (MO). Outskirts of Puerto
Obaldia, Gentry 1570 (MO). VERAGUAS: Santiago, Dwyer & Kirkbride 7438 (MO). Coiba
Island, Seemann 602 (K). Island off Veraguas, Sinclair s.n. (K). Between Sonat and Santiago,
Woodson et al. 1196 (A, K, MO, NY). Without locality, Hinds s.n. (K). WITHOUT LOCALITY:
Duchaissaing s.n. (K); Duke 12268 (MO).
2. Cydista diversifolia (H.B.K.) Miers, Proc. Roy. Hort. Soc. 3: 192. 1863.
Bignonia diversifolia H.B.K., Nov. Gen. Sp. P1. 3: 133. 1819. TYPE: Mexico, Campeche,
Humboldt & Bonpland s.n. (P).
Anemopaegma vargasianum DC., Prodr. 9: 190. 1845. TYPE: Venezuela, Caracas, Vargas s.n.
(G-DC).
Bignonia sagreana DC., Prodr. 9: 148. 1845. TYPE: Cuba, Sagra s.n. (G-DC, K).
Cydista vargasiana (DC.) Miers, Proc. Roy. Hort. Soc. 3: 192. 1863.
Pleonotoma diversifolium (H.B.K.) Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 274. 1897.
Liana to at least 5 cm in diameter, the bark smooth, striate vertically; stem

with 8 phloem arms in cross section; branchlets sharply tetragonal with angles
ribbed, lepidote to simple pubescent, the nodes without interpetiolar glandular
fields; pseudostipules foliaceous, 4 at each node, 0.7-1.5 cm long and 0.6-1.7 cm
wide. Leaves 2-foliolate, at times cirrhose, sometimes simple, especially the first
leaves on a shoot which then intergrade with pseudostipules; leaflets ovate,
acuminate, basally truncate to shallowly cordate, 4.0-11.6 cm long and 2.2-8.5
cm wide, entire, membranaceous, basally palmately 5-veined, the secondary veins
3-4 on a side, the surface with scattered lepidote scales, otherwise glabrous or
simple-puberulent on main veins above and beneath, sometimes sparsely puberu-
lent overall beneath, conspicuous plate-shaped glands in the axils of basal nerves
beneath, drying olive with the venation beneath often reddish; tendril simple,
4-13 cm long; petiolules 1.1-2.2 cm long, the petiole 1.4-3.6 cm long, (to 5 cm
on simple leaves), lepidote to simple puberulous, the simple trichomes often
with curved tips. Inflorescence an axillary or terminal panicle, the branches
lepidote. Flowers with an inconspicuous sweet aroma, calyx cupular, 5-lobed to
more or less truncate, 3-5 mm long and 4-6 mm wide, lepidote, slightly puberu-
lous at least around the margin; corolla magenta, the throat white inside, tubular-
campanulate to tubular-infundibuliform 2.4-3.9 cm long and 0.8-1.8 cm wide at
the mouth, the tube 1.6-2.6 cm long, the lobes 0.7-1.2 cm long, lepidote and
puberulous outside at least on the lobes, within glandular-lepidote on the lobes
and sparsely so in the throat, glandular simple-pubescent at and below the level
of stamen insertion; stamens didynamous, the anther thecae divaricate, 2-2.5 mm
long, the longer filaments 1.1-1.4 cm long, shorter filaments 0.8-1.1 cm long, the
staminodes 3-5 mm long, inserted 2-3 mm from base of the corolla tube; pistil
1.9-2.1 cm long, the ovary cylindric, 3-5 mm long and 1.5 mm wide, lepidote and
inconspicuously puberulous, the ovules 2-seriate; disc lacking. Capsule linear,
strongly compressed, 30-40 cm long and 1.2-1.5 cm wide, inconspicuously lepi-
dote or lepidote-punctate; seeds 1.2-1.3 cm long and 4-5.5 cm wide, the wings
membranaceous, brown, not differentiated from the seed body.
An extremely common species of the tropical dry forest, this plant also occurs
in premontane moist and less commonly in tropical moist forests. In the tropical
moist forest it appears to be restricted to edaphically dry locations, e.g. the only
Panamanian locality is a dry limestone cave area near Madden Lake. The species
ranges from Mexico to Venezuela and the West Indies. It flowers mostly during
the wet season from May through September, also blooming sporadically in the
dry season. Seeds are mostly released during the early dry season and are wind
dispersed.
This species is distinctive in its conspicuously square stem with ribbed angles
and conspicuous foliaceous pseudostipules. Glandular fields in the axils of the
basal pair of lateral nerves of the leaf undersurface are also a useful character.
CANAL ZONE: Boy Scout road, Gentry 2480 (MO). Alrededores de la Represa Madden,
Kant 1 (MO, PMA). PANAMA': Archaeological site near Madden Lake, Gentry 5012 (MO).
3. Cydista heterophylla Seib., Carnegie Inst. Washington Publ. 522: 417. 1940.
TYPE: Mexico, Yucatan, Lundell & Lundell 7350 (A, F, US).-FIG. 10.
Bignonia lepidota Seem., Bot. Voy. Herald 179. 1854, non H.B.K. SYNTYPES: Panama, Isla de
Iguana, Seemann 601 (BM, K); Cuming 1262, pro parte (K).
Liana to at least 5 cm in diameter, the bark smooth, gray; stem with 8(-16)
phloem arms in cross section; branchlets terete to subterete, grayish with incon-
spicuous lenticels, lepidote when young, glabrous when older, the nodes lacking
interpetiolar glandular fields, sometimes with a ridge connecting opposite peti-
oles; pseudostipules lacking. Leaves simple or 2-foliolate, sometimes cirrhose, the
leaves of young branches characteristically simple and appearing in a whorl of 4
due to suppression of an internode; blades ovate to obovate, acute to acuminate,
truncate to subcordate, 4.5-17.5 cm long and 2.1-11.1 cm wide, membranaceous
to chartaceous, secondary veins 3-5 on a side, the basal pair strongly arcuate and
arising from the base, the surface minutely papillate, scattered lepidote above
and beneath, the scales sometimes deciduous, usually minutely scabrous on main
veins, glandular fields in the axils of the basal pair of lateral nerves beneath,
typically drying olive to yellowish-olive beneath with brownish reticulate vena-
tion at least when young, darker above; tendril simple, 8-18 cm long; petiole and
petiolules lepidote to scabrous, the petiolules 0.5-5.1 flowers, usually borne when
leafless, the branches lepidote. Flowers somewhat aromatic, calyx cupular, 4-7
mm long and 4-6 mm wide, shallowly bilabiate to 3-lobed, rarely truncate with
the tip calypterate, the edge often drying lighter, lepidote, sometimes with plate-
shaped glands; corolla lavender to deep magenta inside and on the lobes, the tube
white inside and at base of the 2 adaxial lobes with ca. 13 magenta nectar guides
in the floor, tubular-campanulate to tubular-infundibuliform, slightly bilibiate,
4-5.9 cm long and 1.1-1.7 cm wide at the mouth of tube, the tube 2.8-3.7 cm
long, the lobes 0.9-1.9 cm long, inconspicuously lepidote outside, more so inside,
simple-puberulous on the lobes, especially inside, puberulous with short some-
times gland-tipped trichomes at the level of stamen insertion; stamens didyna-
FIGURE 10. Cydista heterophylla Siebert.-A. Inflorescence ( X 1/2).-B. Ovary cross

section (x 12). [After Gentry 4967 (MO).]-C. Fruit (X 1/2). [After Gentry 4088 (MO).]
-D. Ovary (x 21/2).-E. Habit (x 1/2). [After Gentry 4088 (MO).]-F. Corolla split open
( X3/5). [After Gentry 4967 (MO).]
BSS
mous, the anther thecae divaricate, 4-5 mm long, the longer filaments 1.7-1.8
cm long, the shorter filaments 1.1-1.2 cm long, the staminode 4 mm long, inserted
4-5 mm from the base of the corolla tube; pistil 3.0-3.1 cm long, the ovary linear
lepidote, 4-5 mm long and 1.5 mm wide, the ovules 2-seriate in each locule; disc
reduced to glandular tissue at base of ovary. Capsule linear, flattened but with
2 conspicuous raised longitudinal submarginal ridges on each valve, rounded at
the tip, 11-33 cm long and 1.0-2.0 cm wide, the surface glabrous to more or less
lepidote; seeds 0.8-1.5 cm long and 2.6-5.3 cm wide, the wings membranaceous,
merging into the seed body, brown with irregularly hyaline tips.
A common vine of the tropical dry forest and drier parts of the tropical moist
forest, this species occurs also, but uncommonly, in moister parts of the tropical
moist forest. It ranges from Mexico to Colombia. It flowers during the dry
season especially in April and May. The species is deciduous during the bloom-
ing period so leaves are rare in herbaria.
A propensity to simple leaves and a strongly arcuate basal pair of lateral
nerves with glandular fields in their axils on the undersides mark this species.
An additional field character, especially useful on juvenile plants, is the two
terminal pairs of simple leaves which appear as a whorl of four.
CANAL ZONE: Barro Colorado Island, Croat 5426 (MO), 8206 (MO, SCZ), Shattuck
784, pro parte (F, MO). Victoria Fill near Miraflores Locks, Allen 1756 (GH, MO). Near
Madden Dam, Alston 8865, pro parte (BM, K); Gaillard Highway 1 mi. NW of Summit,
Croat 9776 (MO, NY, SCZ). Near Madden Dam, Dodge 16570 (MO, U). Road K-19, Gentry
1779 (MO). N of Summit, Gentry 5126 (MO). Paraiso, Hayes 126 (BM, K). Las Sabanas
Pittier 6762, pro parte (BM, GH, MO, NY). Cocoli Road, White 146 (GH, MO). DARIEN:
Isla Encanto in Golfo de San Miguel, Gentry 4020 (MO). Near Punta Alegre, Gentry 4059
(MO). Punta Sabanas across bay from La Palma, Gentry 4088 (MO). Ridge from Cerro
Pavarando to Cerro Derumba, Gentry 4206 (MO). Rio Tuira between Rios Paya and Aspave,
Gentry 4420 (MO). El Real, Gentry 4539, 4575 (both MO). HERRERA: E of Las Minas,
Gentry 3143 (MO). PANAMA: E of El Llano, Croat 14482 (MO), 14499 (MO, SCZ). San
Jose Island, Erlanson 38 (GH, US). Rio Pasiga, Gentry 2204, 2256, 2274, 2359 (all MO).
Rio Maestro, Gentry 2219 (MO). Rio Piragua (Tabardi), Gentry 2564 (MO). Icanti, Gentry
2615 (MO). Near Rio Espave, Gentry 3700 (MO). 16 km from Bayano crossing on trail to
Santa Fe, Gentry 3817 (MO). 1 mi. W of Cafnitas, Gentry 4967 (MO). San Jose Island,
Johnston 812 (BM, GH, MO, US), 831 (GH). Juan Diaz, Maxon & Harvey 6640, pro parte
(US). Chepo, Pittier 4766 (JS). Perlas Islands, El Rey Island, Svenson 417 (GH). WITHOUT
LOCALITY, Cuming 1262 (K); Seemann 601 (BM, K).
11. DENDROSICUS
Dendrosicus Raf., Sylva Tell. 80. 1838. TYPE: D. saxatilis Raf. D. latifolius
(Mill.) A. Gentry.
Amphitecna Miers, Trans. Linn. Soc., ser. 1. 26: 163. 1870. TYPE: A. macrophylla (Seem.)
Miers ex Baill. = Dendrosicus macrophyllus (Seem.) A. Gentry.
Crescentia sect. Enallagma Miers, Trans. Linn. Soc., ser. 1. 26: 174. 1870.
Enallagma (Miers) Baill., Bull. Soc. Linn. Paris 85: 679. 1887. TYPE: E. cucurbitina (L.)
Baill. = Dendrosicus latifolius (Mill.) A. Gentry.
Small to medium-sized trees. Leaves simple, alternate, borne singly on the

twigs. Inflorescence a solitary flower or a few-flowered terminal fascicle cauli-
florous on bracteate short-shoots from older twigs and branches. Flowers with
the calyx large, rupturing irregularly, usually bilabiate, sometimes subtruncate,
rarely spathaceous; corolla greenish-white, tubular-campanulate, usually with an

inconspicuous transverse fold in the throat, thick; stamens subexserted, the an-
thers glabrous, the thecae thick, somewhat divergent; pollen grains single, ecol-
pate, the exine microreticulate; ovary ovoid-elliptic, more or less lepidote, in-
completely 2-locular. Fruit a pepo or calabash, spherical to ellipsoidal,
indehiscent, with a relatively fragile woody shell and soft pulp; seeds thick, large,
more than 1.3 cm long and 1.4 cm wide, not winged, embedded in the pulp.
About ten species ranging from Mexico and the West Indies to coastal Vene-
zuela and Ecuador. The species of Dendrosicus are readily distinguished from
most other members of the family by their large, simple, mostly alternate leaves.
The use of the name Dendrosicus for this genus is discussed by Gentry (1973).
Useful references:
Gentry, A. H. Studies in Bignoniaceae VII. Dendrosicus, Enallagma, and
Amphitecna, Taxon 22: 643-646. 1973.
Williams, L. 0. Amphitecna and Enallagma. Fieldiana, Bot. 36: 21-27. 1973.
a. Fruit spherical; leaves coriaceous; occurring near sea level -3. D. latifolius
aa. Fruit ellipsoidal, apiculate; leaves chartaceous; generally occurring above 100 m ele-
vation.
b. Calyx thick, irregularly and shallowly split; flowers mostly terminal; ovary bilocular
to above the middle; leaves drying gray or brownish with darker midvein, without a
fine network of impressed veinlets, not shagreened; over 1000 m elevation
------------------------------------------------------------------------------- 4. D. sessilifolius
bb. Calyx thin, split to near the base; flowers terminal or on bracteate short shoots from
older branches; ovary unilocular except at the extreme base or bilocular to above
the middle; leaves drying gray or olive with lighter midvein, often with a fine net-
work of impressed whitish veinlets and more or less shagreened; under 1000 m
elevation.
c. Calyx bilabiately split; buds round-tipped.
d. Flowers cauliflorous on bracteate short shoots; ovary unilocular except at the
extreme base; Atlantic slope -2. D. kennedyi
dd. Flowers mostly terminal; ovary bilocular to a
--------------------------------------- -----------------------1. D. isthmicus
cc. Calyx spathaceously split; buds pointed ---5. D. spathicalyx
1. Dendrosicus isthmicus A. Gentry, Phytologia 26: 442. 197

Chiriqui, Liesner 75 (MO).
Small tree to 10 m. Leaves simple, alternate, glabrous, narrowly elliptic,

acute to acuminate, basally cuneate, 7-32 cm long and 2.8-10 cm wide, the mid-
rib raised beneath, the main veins drying whitish or yellowish beneath, leaf sur-
face gray or olive-gray above and beneath, the veinlets sometimes minutely ap-
pressed beneath; the petiole less than 5 mm long or lacking. Inflorescence of
1-2 flowers, usually terminal, the pedicel 2-4 cm long. Flowers with the calyx
bilabiately split almost to the base, 2.3-3 cm long, each lobe 1.3-1.8 cm wide,
obtuse, glabrous with a few plate-shaped glands; corolla greenish white, 3.4-6
cm long and 1.7-2 cm wide at the mouth, the tube 2.4-3.5 cm long, glabrous, the
rim lepidote papillate, the lobes fused into a reflexed rim; stamens subexserted,
the anther thecae divergent, each 4-5 mm long and 1-2 mm wide; filaments 2-3
cm long, inserted ca. 1.5 cm from the base of the tube; pistil ca. 4 cm long, the
ovary oblong, 2-3 mm long and ca. 2 mm wide, glandular lepidote, 2-locular in
the lower 2/3 with the ovules several seriate on 2 indistinct axillary placentae
each locule, above unilocular with parietal placentation; disc annular pulvinate.
Fruit a pepo or calabash, to 14 cm long and 7 cm wide, ellipsoid, apiculate at the
ends with thick seeds embedded in the pulp.
This species is apparently restricted to the tropical wet forest of eastern Costa
Rica and western Panama. It has been collected only on the Pacific side of the
Cordillera Central.
The status of this species is difficult to evaluate from the limited material
available. Most of the collections have only a single flower, and the only mature
fruit was found washed up on a beach. These specimens were originally deter-
mined as Enallagma latifolia (= Dendrosicus latifolius), although they appear
more closely related to D. sessilifolius. Dendrosicus isthmicus differs from D.
latifolius in leaf shape and texture, fruit shape, and especially in ecology. It
differs from D. sessilifolius in its generally smaller, narrowly elliptic rather than
oblanceolate to narrowly obovate leaves, as well as in the characters noted in the
key. Athough differences in calyx thickness and degree of splitting are relative,
they appear to be constant. The uneven splitting and thickness of the calyx of
D. sessilifolius usually results in a whitish edging along the split edges of its
black-drying calyx especially near the base of the splits; in D. isthmicus the thin-
ner calyx lacks a whitish edging along the lines of rupture.
The closest relative of D. isthmicus is D. kennedyi, and I am unable to sep-
arate these species reliably on the basis of sterile or fruiting material. Terminal
(noted as terminal or cauline on one collection) placement of flowers, the blunt-
tipped calyx, and geographic separation appear sufficient for separation of this
species from D. kennedyi. In addition the leaves of D. isthmicus arc usually
much smaller and the ovary (based on a single dissection for each species) is
bilocular to above the middle rather than only at the extreme base as in D.
kennedyi.
CHIRIQUI: Progreso, Cooper & Slater 277 (F, NY, US). San Bartolo de Limite, 21 km
WNW of Puerto Armuelles, rich forest, 400 m, Leisner 75 (MO). Burica Peninsula, Quebrada
de Fraile, Liesner s.n. (MO). COCLE: El Valle, vicinity of La Mesa, 900 m, tropical wet
forest, Gentry 7443 (MO).
2. Dendrosicus kennedyi A. Gentry, Phytologia 26: 441. 1973. TYPE: Panama,

Bocas del Toro, Kennedy & Dressler 1258 (MO).-FIG. llA-B.
Small tree. Leaves simple, alternate, glabrous or sparsely lepidote, oblanceo-

late to narrowly obovate, apically acute to acuminate, basally cuneate, attenuate,
the petiole not differentiated, 16-38 cm long and 5-14 cm wide, the midrib con-
spicuously raised beneath, the main veins drying whitish beneath, leaf surface
gray above and grayish beneath, the veinlets sometimes minutely impressed be-
neath. Inflorescence 1-2 flowers from a conspicuously bracteate abbreviated
short-shoot on older branches or trunk, the pedicels 1.5-3 cm long. Flowers with
the calyx bilabiately split almost to base, 1.5-3.0 cm long, each lobe ca. 1 cm
wide, conspicuously apiculate; corolla greenish white, 4.3-4.8 cm long and 1.3-1.9
N'' ~ ~ ~ ~
FIGURE(11. Dendrosicus. A-B. D. kennedyi A. Gentry.-A.5Habit
.......] .......
H For 737, Coombia( MO)/.G
FIGUIRE 11. Dendrosicus.-A-B. D. kennedyi A. Gentry.-A. Habit (X 3Ao).-B In-

florescence (x %). [After Kennedy & Dressier 1258 (MO) .]-C-C D. spathicalyx A. Gentry.
-C. Flower ( x %). [After Gentry 5769 (MO) .]-D. Fruit (X '/2). [Aftei Dressier 3943
(MO).]-E. Corolla (x %/).-F. Ovary cross section (x %).-G. Ovary and disc (x 11/i).
[After Gentry 5769 (MO)J.-H. D. latifolius (Mill.) A. Gentry, calyx (X 3/5)) [After Gentry
& Forero 7379, Colombia (MO).]I
cm wide at the mouth, the tube 2.7-3.5 cm long, the lobes fused into a reflexed
rim; stamens subexserted, the anther thecae somewhat divergent, 4-5 mm long
and 1.5-2 mm wide, the filaments 2.4-2.8 cm long, inserted 1.3-1.5 cm from the
base of the tube; pistil ca. 4 cm long, the ovary rounded conical, 2-3 mm long
and 2 mm wide, densely lepidote, the ovules several seriate on 2 parietal placen-
tae; disc annular pulvinate. Fruit a pepo or calabash, 11.5-13 cm long and 5.5-7
cm wide, ellipsoid, apiculate at the ends with thick seeds 1.3-1.5 cm long em-
bedded in the pulp.
This species is restricted to the tropical wet forest and premontane wet forest
of Bocas del Toro Province and adjacent Costa Rica. It is distinguished from its
congeners in Panama by its thin, sharply pointed, deeply bilabiate calyx. Among
its closest allies are D. isthmicus and D. macrophyllus (Seem.) A. Gentry of
Mexico and Guatemala. The former differs in the characters given in the key;
the latter most conspicuously in its 8-costate rather than smooth fruit.
It is possible that the collections cited include a second species. The Kirkbride
and Duke and the von Wedel collections have shorter leaves which are widest
near the middle and may prove separable from D. kennedyi. These two collec-
tions are somewhat similar to material from the wet Pacific lowlands in Costa
Rica and Panama described as D. isthmicus. Inadequate material prevents better
evaluation of these collections.
BOCAS DEL TORO: Changuinola Valley, Cooper & Slater 36 (US). Near Almirante,
Kennedy & Dressler 1258 (MO), 1261 (MO, PMA). Above Quebrada Huron on Cerro
Bonyic, Kirkbride & Duke 615 (MO). Fish Creek mountains, von Wedel 2258 (GH, MO).
3. Dendrosicus latifolius (Mill.) A. Gentry, Taxon 22: 644. 1973.-FIG. 1111.
Crescentia latifolia Mill., Gard. Dict., ed. 8. 306. 1768. TYPE: (not seen).
C. cucurbitina L., Mant. P1. 2: 250. 1771. TYPE: Based on C. latifolia Mill.
C. lethifera Tussac, Fl. Antill. 4: 50, tab. 17. 1827. TYPE: (not seen).
C. toxicaria Tussac, Fl. Antill. 4: 50, tab. 17. 1827. TYPE: (not seen).
Dendrosicus saxatilis Raf., Sylva Tell. 80. 1838. TYPE: ;Based on C. cucurbitina L.
Crescentia obovata Benth., Bot. Voy. Sulphur 130. 1844. TYPE: Panama, Isle of Gorgona,
Barclay s.n. (BM, K).
C. palustris Forsyth ex Seem., Bot. Voy. Herald 183. 1854, pro syn.
C. coriacea Miers, Trans. Linn. Soc. 26: 177. 1870. TYPE: Fruit and unpressed leaves, without
data, P. Browne, Jamaica? (BM).
C. cuspidata Miers, Trans. Linn. Soc. 26: 178. 1870. TYPE: Venezuela, Fendler 780 (K).
Enallagma cucurbitina (L.) Baill. ex Schum. in Engler & Prantl, Nat. Pflanzenf. 4(3b): 247.
1894.
E. obovata (Benth.) Baill. ex Schum. in Engler & Prantl, Nat. Pflanzenf. 4(3b): 247. 1894.
E. latifolia (Mill.) Small, Fl. Miami 171. 1913.
Amphitecna obovata (Benth.) L. 0. Wms., Fieldiana, Bot. 36: 25. 1973.
Small scraggly tree to 10 m tall and 18 cm d.b.h., the branches usually crooked,
the crown open; twigs subterete or 3- or 4-angled with decurrent ridges below the
nodes. Leaves simple, alternate to subopposite, obovate, the tip rounded to al-
most acute, usually apiculate, the base cuneate-attenuate, 7-19 cm long and 3.3-
10.6 cm wide, rigid-chartaceous to coriaceous, secondary veins 7-30 on a side, the
midvein depressed above, glabrous beneath, with plate-shaped glands especially
at the base, drying grayish olive to brownish, the petiole poorly defined, more or
less winged, 0.2-1.1 cm long, glabrous. Inflorescence a single flower or fascicle

of 2-3 flowers borne on the larger twigs, pedicels glabrous, 2.2-3.6 cm long.
Flowers with the calyx split into subequal anterior and posterior halves, 2.5-3.7
cm long and 1-2.5 cm wide, essentially glabrous; corolla greenish-white, tubular
with a transverse fold midway across the lower side of throat, fleshy, 3.8-6.2 cm
long and 1.2-2.4 cm wide at mouth, the tube 3.0-4.0 cm long, the lobes 0.8-1.5
cm long, upper 2 and lower 3 lobes respectively more or less fused, mostly gla-
brous, stalked-lepidote on the lobes outside and at the level of stamen insertion
inside; stamens subexserted, the anther thecae thick, only partially divergent,
5-6 mm long, the anterior filaments 2.6 cm long and inserted 1.5-1.6 cm
from base of tube, the posterior filaments 2.4-2.5 cm long, inserted 2 cm from
base of tube, staminode ca. 10 mm long, inserted ca. 10 mm from base of tube;
pistil 5.0-5.1 cm long, the ovary rounded-conical, 4-5 mm long and 3 mm wide,
lepidote, the ovules ca. 4-seriate in each locule of the bilocular ovary; disc pulvi-
nate, 2 mm long and 5 mm wide. Fruit a pepo or calabash, more or less spherical
when mature, 6-9 cm in diameter, glabrous, minutely papillate with scattered
sunken plate-shaped glands; seeds thick, corky, 1.3-1.6 cm long and 1.6-1.8 cm
wide, without wings.
A common tree of seashores, mangrove fringes, and brackish water swamps

along both coasts, D. latifolius ranges from southern Florida and Mexico through
the West Indies to Venezuela. It flowers more or less throughout the year, es-
pecially in the wet season.
The fruits are reported by campesinos to be an important food source of the
paca and the insipid pulp is eaten by humans in some localities. Fruits of D.
latifolius are commonly found washed up on beaches with still fertile seeds wh
germinate only when the fruit is broken open. Evidently seed dispersal is a
combination of water-dispersed fruit with mammal-dispersed seeds, either vector
being sufficient to disseminate the species.
The large, simple, typically alternate leaves of D. latifolius are unusual in the
Bignoniaceae. Their coriaceous texture and the presence of a definite petiole
serve to distinguish this species from D. sessilifolius. The more or less spherical
fruit is superficially similar to that of Crescentia alata but unlike the more elon-
gate fruits of other species of Dendrosicus.
BOCAS DEL TORO: Isla Colon, von Wedel 561 (GH, MO, U). Water Valley, von Wedel
988 (GH, MO). Old Bank Island, von Wedel 2101 (GH, MO, US). Johns Creek, von Wedel
2769 (CGH, MO, US). CANAL ZONE: Summit Garden, Croat 13827 (MO, SCZ). Chagres,
Fendler 210 (K). 1 mi. NE of Pifia, Lewis et al. 1842 (MO). Along Rio Indio de Gatuin,
Pittier 2782 (US). Vicinity of Fort Sherman, Standley 30919 (US); Tyson & Blum 3797
(MO, SCZ). COLON: Vicinity of Camp Pifia, Allen 3632 (MO). 3 mi. SW of Colon along
beach, Croat 14168 (MO). W of Portobelo, Gentry 1742 (MO). Aspinwall, Hayes 862 (NY).
Punta Pedro, Icacal between Salud and mouth of Rio Indio, Howell 40 (MO). Salud, Lao
& Holdridge 250 (MO). Rio Indio de Fato, Pittier 4272 (US). Plaza Colon, Rose 22028
(US), 22078 (NY). DARIEN: On Rio Tuira ca. 3 mi. NW of El Real, Duke 4820 (SCZ). S of
Jaque, Gentry 4121 (MO). Along Rio Tuira below El Real and Piriaque Island, Stern et al.
891 (GH, K, MO). LOS SANTOS: Azuero Peninsula near Raja Point, Ostenfeld 20 (C, US).
Punta Mala, Tyson 2738 (MO, SCZ). PANAMA: San Jose Island, Erlanson 4 (GH, NY, US);
Harlow 97 (P. US); Johnston 10, 1179 (both GH); Stimson 5333 (NY, PMA, SCZ); Tyson
& Loftin 5074 (SCZ). Seaside, Panamat, Hayes 826 (NY). SAN BLAS: Mulatupo, Duke 8552
(MO). Near Puerto Obaldia, Pittier 4362 (US). VERAGUAS: Isla de Coiba, Dwyer 1568
(MO); Foster 1617 (PMA); Kelsall 467 (K). WITHOUT LOCALITY: Barclay s.n. (BM, K);
Hayes 44 (CGH).
4. Dendrosicus sessilifolius (Donn. Sm.) A. Gentry, Taxon 22: 646. 1973.

Tabebuia sessilifolia Donn. Sm., Bot. Gaz. (Crawfordsville) 25: 156. 1898. TYPE: Costa
Rica, Tablazo, Tonduz 7937 (BR, CR, US).
Neotuerckheimia gonoclada Donn. Sm., Bot. Gas. (Crawfordsville) 47: 259. 1909. TYPE:
Costa Rica, San Jose, Tonduz 7384 (BM, GH, K, M, MO, NY, US).
Enallagma sessilifolia (Donn. Sm.) Standl., Publ. Field Mus. Nat. Hist., Bot. ser. 18: 1120.
1938.
Amphitecna sessilifolia (Donn. Sm.) L. 0. Wms., Fieldiana, Bot. 36: 25. 1973.
Shrub or tree to 18 m tall and 25 cm d.b.h.; twigs subterete, glabrate, usually

moss-covered. Leaves simple, alternate, oblanceolate to narrowly obovate, acute,
attenuate with the petiole undifferentiated and essentially lacking, 13-29 cm long
and 4.0-8.5 cm wide, chartaceous, secondary veins 11-16 on a side, the midvein
raised above, lepidote above and beneath, a few plate-shaped glands at base
beneath, drying dark brown. Inflorescence a terminal fascicle, the pedicels gla-
brous to inconspicuously lepidote, 2.2-7.9 cm long, commonly reduced to a single
flower. Flower with the calyx thick, fleshy, irregularly split into subequal an-
terior and posterior halves or merely ruptured apically by the expanding corolla,
one of the lobes sometimes itself split, 2.5-2.8 cm long and 1.9-2.1 cm wide,
glabrous; corolla greenish white, tubular, usually with an inconspicuous trans-
verse fold in the throat, fleshy, 4.1-5.2 cm long and 1.2-1.6 cm wide at the mouth,
the tube 2.5-3.3 cm long, the lobes fused, 0.5-1.2 cm long, glabrous; stamens
subexserted, the anther thecae thick, somewhat divergent, 5 mm long and 2-3
mm wide, the anterior filaments 2.3-2.4 cm long, inserted 13-14 mm from base
of tube, the posterior filaments 2.5-2.7 cm long, inserted 16-18 mm from base of
tube, the staminode 2.5-2.7 cm long, inserted 16-18 mm from the base of the
tube; pistil 3.8-3.9 cm long, the ovary rounded-conical, 8-9 mm long and 4-5 mm
wide, incompletely 2-locular with 2 protruding, parietal placentae at the top,
these fused above middle, the ovules then irregularly 4-seriate in each locule;
disc pulvinate, ca. 3 mm long and 9-10 mm wide. Fruit a pepo or calabash,
ellipsoidal with the apex elongated, 12-15 cm long and 6.6-7.2 cm in diameter,
evenly rough-surfaced under a lens, inconspicuously scattered lepidote; seeds
thick, 1.5-2 cm long and 1.5-2.4 cm wide, without wings.
A tree of the premontane rain forest and lower montane rain forest of Costa
Rica and Panama, this species occurs from 1200-2000 m elevation. Dendrosicus
sessilifolius flowers and fruits irregularly throughout the year.
The fruits are much softer-shelled than those of D. latifolius, probably reflect-
ing elimination of water borne dispersal. It seems that mammals disperse the
seeds by eating the pulp which surrounds them; the pulp of mature fruits is also
eaten by campesinos in Chiriqui Province.
The leaves of D. sessilifolius can be told from those of D. latifolius by their
less rigid texture and narrower shape; the ecology of the two species is also dis-
tinct. The ellipsoid, apiculate fruits of D. sessilifolius are different from the
spherical fruits of D. latifolius. Dendrosicus spathicalyx has smaller leaves and
a spathaceous calyx; D. kennedyi has a much thinner, evenly bilabiate calyx with
each lobe sharply apiculate; D. isthmicus has a thinner calyx and narrowly ellip-
tic, usually smaller, leaves.
CHIRIQUI: Central valley of Rio Chiriqui Viejo, Allen 1362 (GH, MO, NY, US). Near
Boquete, Blum & Dwyer 2570 (MO, SCZ). Bajo Chorro, Boquete District, Davidson 410
(MO). Cerro Horqueta NW of Boquete, Dwyer et al. 444 (GH, MO, US); Dwyer & Hayden
7694 (K, MO). Nueva Suissa, Gentry 5982 (MO). 2 mi. E of El Hato del Volcan, King 5305
(K, US). Around El Boquete, Pittier 3001 (NY, US). Rio Chiriqui Viejo valley near El
Volcain, White 227 (CGH, MO, US).
5. Dendrosicus spathicalyx A. Gentry, Phytologia 26: 439. 1973. TYPE: Panama,

Gentry 5769 (MO, SCZ) .-FIG. 11C-G.
Small, leaning tree 5-7 m tall, 15 cm d.b.h., the bark thickly moss-encrusted;
the twigs decurrent below the nodes. Leaves alternate, narrowly elliptic to nar-
rowly obovate-elliptic, acute to long acuminate, basally cuneate or attenuate,
essentially sessile, the extreme base swollen and woody, conspicuously jointed at
nodes, 7.5-16 cm long and 2.1-4.3 cm wide, chartaceous, the secondary nerves
6-12 on a side, slightly raised above, conspicuously so beneath, the veinlets mi-
nutely and intricately impressed above and beneath, the surface thus roughened
and shagreened. Inflorescence a single flower borne from a bracteate short-shoot
on an older twig, the pedicel glabrous, red when fresh. Flowers with the calyx
membranaceous, 4-4.1 cm long and ca. 2 cm wide, spathaceously split adaxially
to 0.8-1 cm from the base with a tendency to tardily circumcissile dehiscence,
glabrous, light green when fresh; corolla cream to whitish basally, becoming
greenish white toward the mouth, campanulate with a transverse fold in the
throat, 5.5-5.8 cm long and 1.8-1.9 cm wide at the mouth, the tube 3.7-3.8 cm
long, the lobes fused into a reflexed rim, 1.7 cm wide, glabrous with a glandular
epidermis inside and out, becoming stalked glandular-lepidote distally outside;
stamens subexserted, the anther thecae slightly divergent, 7-8 mm long and 1.5-2
mm wide, the anterior filaments 2.5-2.6 cm long, inserted 1.7-1.8 cm from the
base of the tube, the posterior filaments 2.3-2.4 cm, long, inserted 2.0-2.1 cm from
the base of the tube, the staminode 1.1-1.2 cm long, inserted 1.0 cm from the
base of the tube, bent straight out from the corolla; pistil (partially destroyed in
all flowers examined) with the ovary rounded conical, 4 mm long and 3-4 mm
wide, lepidote, the ovules widely spaced, irregularly 8-seriate on each of 2 pro-
jecting parietal placentae, the placentae almost meeting but fused only at the
extreme base; disc annular-pulvinate, 2 mm long and 9 mm wide. Fruit a pepo or
calabash, ellipsoidal, pointed.
This species is known only from the tropical wet forest on top of Cerro
Campana. Its unique, thin, dorsally split, spathaceous calyx is distinctive. Its
bracteate short-shoots relate it to D. macrophyllus, as does the intricately im-
pressed finer venation of the leaves. Except for the calyx this species appears
similar to Dendrosicus silvicola (L. 0. Wms.) A. Gentry. An additional fruiting
collection, Kennedy & Foster 2147 from the Rio Guanche, Colon Province, may
represent this species, but without buds or flowers I am unable to positively
distinguish it from D. isthmicus.
PANAMA: Cerro Campana, Croat 14765 (MO, SCZ); Dressler 3943 (MO); Gentry 5769
(MO, SCZ).
12. DISTICTELLA
Distictella 0. Kuntze in Post & 0. Kuntze, Lex. Gen. Phan. 182. 1903. LECTOTYPE:
D. mansoana (DC.) Urb.
Distictis Bur., Mon. Bignon. 46. 1864, non Mart. ex Meisn.
Lianas, rarely shrubs, branchlets terete; stems with 4 phloem arms in cross
section; branchlets terete without interpetiolar glandular fields; pseudostipules
inconspicuous, short and thick. Leaves 3-foliolate, the terminal leaflet often re-
placed by a trifid tendril. Inflorescence a terminal or axillary raceme or racemose
panicle. Flowers with the calyx cupular, truncate, thick, usually withn glandular
fields below the margin; corolla white, tubular-campanulate, densely short-pu-
bescent outside; anthers glabrous, the thecae straight, divaricate; pollen grains
ecolpate, the exine alveolate; ovary ovoid-oblong, puberulous, the ovules 4-8-
seriate in each locule; disc annular-pulvinate. Fruit a compressed or biconvex
oblong capsule, the valves parallel to the septum, thick and woody, smooth or
minutely rough-surfaced, often shortly puberulous; seeds thin but usually more
or less woody, the wings glabrous, usually brown, often more or less reduced.
About 10 species in continental tropical America ranging from Costa Rica to

Brazil. The genus has not previously been reported in Central America.
Distictella magnoliifolia (H.B.K.) Sandw., Lilloa 3: 460. 1938.-FIG. 12.

Bignonia magnoliaefolia [sic] H.B.K., Nov. Gen. Sp. P1. 3: 136. 1819. TYPE: Venezuela,
Javita, Humboldt & Bonpland 973 (P).
B. alliacea Lam. sec. Splitg., Tijdschr. Natuurl. Gesch. Physiol. 9: 5. 1842, non Lam.
Distictis racemosa Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 179. 1897. SYNTYPES: Surinam,
Wullschlagel 1033 (BR); Brazil, Para, Martius s.n. (BR).
Bignonia rusbyi Britt. ex Rusby, Bull. Torrey Bot. Club 27: 71. 1900. TYPE: Bolivia, junction
of Rios Beni and Madre de Dios, Rusby 1140 (NY).
Distictis kochii Pilg., Fedde Repert. 8: 151. 1910. TYPE: Brazil, Vaupes region, Koch-Grunberg
s.n. (B-destroyed; K, photo).
Distictella broadwayana Urb., Fedde Repert. 14: 310. 1916. TYPE: Tobago, near Menna,
Broadway 4753 (BM).
D. kochii (Pi1g.) Urb., Fedde Repert. 14: 310. 1916.
D. racemosa (Bur. & K. Schum.) Urb., Fedde Repert. 14: 310. 1916.
Liana to at least 7 cm in diameter, the bark somewhat fibrous, vertically

striate, twisting to form a bush rope; stem with 4 phloem arms in cross section
at least when young; branchlets terete, often somewhat flattened at the nodes,
appressed-puberulous, usually reddish brown; nodes without interpetiolar glandu-
lar fields, the pseudostipules short and thick, inconspicuous. Leaves 2-foliolate
with a tendril or tendril scar; leaflets more or less elliptic, acute to acuminate,
the base widely cuneate, often briefly truncate at the apex of petiolule, 13.5-29
cm long and 6.1-13.0 cm wide, chartaceous to subcoriaceous, secondary veins 7-9
on each side, lepidote beneath and less so above, beneath appressed puberulous
on the midvein and pubescent in axils of lateral nerves with some plate-shaped
V~~~~~~~~~~~~~~~~~~~~~~~.
@ X ,,/ A~~~~~~/
o A~~~~~~~
FIGURE 12. Distictella magnoliifolia (H.B.K.) Sandw.-A. Habit (X 3/%5).-B. Ovary
and disc ( X 3%).- C. Ovary cross section ( X 71/.,). [After Woytkowski 5140, Peru, and Gentry
737 (both MO).]-D. Fruit (x 3/5). [After Krukoff 6728, Brazil (MO).]
glands near the midvein, sometimes also apically, above somewhat puberulous on
the midvein, drying brownish-olive with the veins usually impressed; tendril
trifid, 6-16 cm long to branching, the arms 0.2-1.5 cm long; petiolules and petiole
appressed-puberulous and lepidote, prominently jointed, the petiolules 0.9-2.7
cm long, the petiole 1.8-5.8 cm long. Inflorescence a racemose panicle, its
branches densely subpuberulous. Flowers with the calyx cupular, thick, subtrun-
cate, 7-10 mm long and 6-8 mm wide, short-puberulous with linear fields of
plate-shaped glands descending from the rim in the upper half; corolla white,
thick, tubular-campanulate above a short, narrowed base, often bent slightly at
the middle, 4.5-6.0 cm long and 1.4-1.7 cm wide at mouth, the tube 3.24.0 cm
long, the lobes 1.0-1.2 cm long, densely short-pubescent on the tube outside and
on the lobes, the tube glabrous inside except for dense glandular trichomes which
form a ridge at the level of stamen insertion; stamens didynamous, the anther
thecae divaricate, 4-5 mm long, the longer filaments 2.1-2.3 cm long, the shorter
filaments 1.6-1.7 cm long, the staminode 6-8 mm long, inserted 7-10 mm from
the corolla base; pistil 3.8-3.9 cm long, the style appressed-puberulent, the ovary
flattened-ellipsoid, 3-4 mm long and 2 mm wide, 1.5 mm thick, densely appressed-
puberulent, the ovules more or less 6-seriate in each locule; disc pulvinate with
a narrowed neck, 1.5-2 mm long and ca. 4 mm wide. Capsule (Krukoff 6728
(MO), Amazonas, Brazil) elliptic-oblong, somewhat compressed, the valves thick
and woody, the midrib not visible, 10-15 cm long and 5-6 cm wide, 1.6-1.7 cm
thick, appressed-tomentose with numerous, small, black-drying glandular areas,
yellowish-brown in color as the trichomes, the surface somewhat roughened;
seeds 2.1-2.4 cm long and 3.94.0 cm wide, flat and thin but brown and more or
less woody.
A vine of the tropical wet forest, this species ranges from Costa Rica to Brazil.
In the field the trifid tendrils and subterete, thick, smooth, reddish twigs, some-
what flattened at the nodes, identify it vegetatively.
The material referred to here has gone under the name D. racemosa, sup-
posedly distinct from D. magnoliifolia on the basis of leaflets with veins and vein-
lets impressed above and lacking glandular fields. Both characters are variable,
and the distinction does not seem adequate for specific separation.
The description of the fruit of this species is based on extra-limital material.
All of the fruits I have seen on South American collections are quite homoge-
neous, according well with this description, but a single unattached capsule valve
collected from the forest floor at Jaque apparently belongs to Distictella and is
notably different from the South American collections I have seen. This valve is
glabrous except for dense lepidote scales and is reddish brown in color, although
agreeing in size and shape. If this capsule valve is found to be representative
of the Central American plant, then a re-examination of its taxonomic position
is in order.
CANAL ZONE: Pipeline Road, TTC transect 3, Gentry 7400 (MO). COLON: Santa Rita
Ridge, Croat 13183 (MO, SCZ); Gentry 737, 1861, 1882B (all MO). DARIEN: S of Jaque,
Gentry 4125 (MO). Slopes of Cerro Chucula, Gentry 4265 (MO).
13. GODMANIA
Godmania Hemsl., Diag. P1. Mex. Centr. Amer. 35. 1879. TYPE: G. macrocarpa
(Benth.) Hemsl. = G. aesculifolia (H.B.K.) Standl.
Small to medium-sized trees. Leaves digitately 5-9-foliolate. Inflorescence

a large terminal corymbose panicle. Flowers with the calyx small, less than 2
mm long, broadly campanulate, shortly 5-lobed; corolla yellow ventrally and
orange-brown dorsally with edges of the lower 3 lobes also brownish, urceolate,
bilabiate, less than 1.6 cm long, puberulous; anthers pubescent, the thecae divari-
cate, less than 1 mm long, the filaments pubescent; pollen grains single, 3-colpate,
the exine microreticulate; ovary linear-conical, lepidote and more or less puberu-
lent, the ovules multiseriate in each locule; disc pulvinate. Fruit a linear capsule,
dehiscing perpendicularly to the septum; seeds bialate, with long, sharply demar-
cated, irregular-ended, hyaline-membranaceous wings.
One species of continental tropical America ranging from Mexico to Brazil

and Bolivia.
Godmania aesculifolia (H.B.K.) Standl. in Standl. & Calderon, Lista Prelim.

P1. El Salvador 200. 1925.-FIG. 13.
Bignonia aesculifolia H.B.K., Nov. Gen. Sp. P1. 3: 140. 1819. TYPE: Mexico, Guerrero,
Tecoma digitata H.B.K., Nov. Gen. Sp. P1. 3: 142. 1819. TYPE: Colombia, Santa Cruz,
T. fuscata Moc. ex DC., Prodr. 9: 218. 1845. TYPE: (not seen).
T.? aesculifolia (H.B.K.) DC., Prodr. 9: 221. 1845.
Cybistax macrocarpa Benth. in Benth. & Hook., Gen. P1. 2: 1043. 1876. TYPE: Panama, near
Panama and Rio Grande Station, Hayes 61 (BM, K, P).
Godmania macrocarpa (Benth.) Hemsl., Diag. P1. Mex. Centr. Amer. 2: 35. 1879.
Tabebuia aesculifolia (H.B.K.) Hemsl., Biol. Centr. Amer. Bot. 2: 494. 1882.
T. fuscata (Moc. ex DC.) Hemsl., Biol. Centr. Amer. Bot. 2: 494. 1882.
Godmania uleana Kranzl., Notizbl. Bot. Gart. Berlin 6: 379. 1915. TYPE: Brazil, Rio Branco,
Ule 7696 (US).
Small to medium-sized tree to 13 m tall and 30 cm d.b.h; bark smooth to

longitudinally ridged; twigs puberulous to subpuberulous, terete, thick with a
large pith, foul-smelling when broken (as the inner bark) with an odor rem-
iniscent of horse urine. Leaves digitately (5-)7-9-foliolate; the leaflets more or
less obovate or oblanceolate, acute, attenuate-cuneate, membranaceous, secondary
veins 7-13 on a side, above and beneath simple puberulous especially along veins,
densely impressed-lepidote, usually with a few plate-shaped glands along the
midvein beneath, drying more or less olive above and brownish beneath, the
terminal leaflet 8.2-16.5 cm long and 3.1-7.2 cm wide, the first intermediate pair
6.7-17.5 cm long and 2.5-7.0 cm wide, the second pair 5-15.2 cm long and 2-5.6
cm wide, the third pair 2.7-6.5 cm long and 0.9-2.9 cm wide, the fourth pair
when present smaller, the terminal to basal petiolules 0.2-2.3 cm, 0.2-1.6 cm,
0.2-0.9 cm, and 0.2-0.7 cm long respectively; petiole 6.6-15.5 cm long, puberulous
as the petiolules. Inflorescence a flat-topped, terminal panicle of few to many
flowers, its branches puberulous. Flowers with a musky odor, only a few opening
B~~~~~~~~~~~~~
-7
FIGURE 13. Godmania aesculifolia (H.B.K.) Standl. A. Habit (x 1/2). [After Allen
4492 (MO).]-B. Fruit (X 1/4). [After Gentry 359, Costa Rica (MO).]
at a time; calyx widely campanulate, shortly 5-lobed, lepidote and simple-p

lent, 1-2 mm long and 2-4 mm wide; corolla yellow ventrally and orangish-brown
dorsally outside, inside the ventral throat and most of the lower 3 lobes yellow,
the dorsal 2 lobes brownish at the edges of the lower 3 lobes and the dorsal part
of the throat, urceolate, 1.0-1.6 cm long and 7-8 mm wide, the tube 9-10 mm
long, the upper 2 lobes 0.2-0.3 cm long, the 3 lower lobes 0.4-0.3 cm long, simple-
puberulent outside, short-pubescent inside on the edges of the lobes with long,
simple trichomes on the lower lobe, in the floor of throat, and sparsely at the
pubescent, less than 1 mm long, the filaments pubescent, the longer pair 0.8-1.0
cm long, the shorter pair 0.6-0.7 cm long, the staminode 0.4 cm long, inserted
1-3 mm from base of corolla tube; pistil curved to fit against the corolla roof, 1.1
cm long, the ovary linear-conical, 2 mm long, 1 mm wide, strongly lepidote and
somewhat simple-puberulous, the ovules multiseriate; disc pulvinate, inconspicu-
ous, 0.5 mm long and 1.5 mm wide. Capsule linear, corkscrew-twisted, 45-100
cm long and 0.9-1.5 cm wide, terete, longitudinally finely ridged, somewhat sim-
ple-puberulous; seeds thin, bialate, 0.9-1.5 cm long and 7.0-13.5 cm wide, the
wings hyaline-membranaceous, sharply demarcated, narrow, long, irregular-
ended.
A tree of the tropical dry forest and premontane moist forest, this species also
occurs less commonly in drier parts of the tropical moist forest. It ranges from
Mexico to Brazil and Bolivia. It flowers primarily in the dry season, and fruit
set is in middle or late dry season.
The tiny flowers (less than 1.6 cm) are the smallest of the neotropical
Tecomeae and are distinctive in shape and color pattern. The spirally twisted
fruit is unique among mainland Bignoniaceae (Spirotecoma of the West Indies
has a similar fruit). The leaves vary extensively in shape and amount of pu-
bescence, but the attenuate more or less apetiolular leaflet base is a fairly constant
distinguishing character. Another field character, especially useful when the tree
is leafless, is the rank odor of the broken twigs.
CANAL ZONE: Vicinity of Summit, Allen 4492 (GH, MO, P, U, US). Near beach at Fort
Kobbe, Duke 4244 (K, MO). NE of Summit, Fosberg 27394 (NY). W end of Thatcher
Ferry Bridge, Gentry 1394 (MO). Pipeline Road, Gentry 2096 (MO). Near City of Panama,
Hayes 61 (BM). Near Gamboa, Pittier 3703 (GH, NY, US). Las Sabanas, Pittier 6720
(C, NY). Pittier 7002 (US). Near Balboa, Standley 29238, 32130 (both US). Chiva Chiva
Trail, Tyson 1335 (MO, SCZ). Albrook Air Force Base, Tyson 2041 (SCZ). Chiva Chiva
Trail, Tyson & Blum 3569 (GH, MO, SCZ). Curundu, Tyson & Dwyer 4456 (GH, MO,
NY, SCZ, US). Near bridge between "our" island and mainland, White 105 (GH, MO, US).
CHIRIQUI: Vicinity of Boquete, Allen 1018 (MO). W of Rio Chorchita, Gentry 5851 (MO).
DARIEN: Sexton 116 (MO). HERBERA: La Cabuya, Las Minas, Lao 140 (MO). PANAMA:
Below El Valle, Gentry & Dwyer 3675 (MO). Near Panamai, Hayes 61 (BM, K, P). Pedregal,
Ciudad de Panama, Holdridge 6505 (MO, PMA). N of Panama City, Paul 455 (US). Juan
Diaz, Standley 30594 (US). Rio Tapia, Standley 30678 (US). Nuevo San Francisco, Standley
30699 (US). Between Las Sabanas and Matias Hernandez, Standley 31816, 31845 (both US).
VERAGUAS: NE of La Mesa, Blum & Tyson 654 (MO, SCZ). WITHOUT LOCALITY: Duchaissing
s.n. (CGH); Grisebach s.n. (MO).
14. HAPLOPHRAGMA
Haplophragma Dop, Bull. Soc. Bot. France 72: 889. 1925. TYPE: H. adenophyl-
lum (Wall. ex G. Don) Dop.
Trees. Leaves imparipinnate, glabrous to tomentose. Inflorescence an axillary

or terminal, many-flowered panicle. Flowers with the calyx campanulate or
tubular-campanulate, irregularly 2-5-lobed; corolla broadly campanulate above
a tubular base, pubescent outside; anthers slightly divergent, gabrous or puberu-
lous; ovary cylindrical, pubescent, the ovules multiseriate in each locule; disc
annular-pulvinate. Fruit a cylindrical capsule, more or less terete, usually spirally
twisted, dehiscent perpendicular to the septum; seeds flat, winged.
Two species native to southeast Asia.
1. Haplophragma adenophyllum (Wall. ex G. Don) Dop, Bull. Soc. Bot.

France 72: 890. 1925.-FIG. 14.
Bignonia adenophylla Wall., Cat. 6502. 1832, nomen nudum.

B. adenophylla Wall. ex G. Don, Gen. Syst. 4: 221. 1838. TYPE: (not seen).
Spathodea adenophylla (Wall. ex G. Don) DC., Prodr. 9: 206. 1845.
Heterophragma adenophyllum (Wall. ex G. Don) Seem. ex Benth. & Hook., Gen. Pl. 2: 1047.
1875.
Tree 10 m tall, 25 cm d.b.h.; branchlets thick, terete. Leaves pinnately com-

pound (3-)5-7-foliolate, the upper leaflets 14-21 cm long and 9-13 cm wide,
the lowest leaflet pair near the base of petiole, much reduced, resembling foli-
aceous pseudostipules, all leaflets elliptic to elliptic-ovate, obtuse, basally trun-
cate, stellate pubescent with mostly few-armed trichomes, especially beneath.
Inflorescence an open panicle, its branches pubescent with brown stellate tri-
chomes. Flowers with the calyx cupular-campanulate, irregularly 3-5-lobed,
3.7-5 cm long and 2.5-3.7 cm wide, densely pubescent with brown stellate tri-
chomes, these interspersed with longer, pale, fimbriate-stellate trichomes; corolla
light yellow, the tube brown outside from the pubescence, broadly campanulate
above a 1.2-1.4-cm-wide neck, the extreme base enlarged to 2-2.5 cm, 8-9 cm
long and 4-4.5 cm wide at mouth, the tube 6-7 cm long, the lobes 2-3 cm long;
stamens subexserted, the thecae parallel or slightly divergent, 6-7 mm long,
slightly pubescent near attachment to connective, the posterior filaments 3.8
cm long, the anterior filaments 4.6-4.8 cm long, the staminode 14-15 mm long
inserted 2.2-2.5 cm from the base of tube; pistil 6.5-7.1 cm long, the style base
stellate pubescent; ovary cylindrical, 6-9 mm long and 4-5 mm wide, densely
brown-pubescent with stellate trichomes, the ovules multiseriate; disc annular-
pulvinate, 3-4 mm long and 14-15 mm wide. Capsule linear, terete with 8 slightly
raised, longitudinal ridges, spirally twisted, 55-100 cm long and 1.5-2 cm wide,
conspicuously stellate-pubescent with brown trichomes, spotted with small,
lighter, glabrous lenticels, the calyx persistent; seeds flat, 0.8-1.1 cm long and
3.4-3.7 cm wide, the wings hyaline-membranaceous.
A species of southeast Asia ranging from Burma to Malaysia, H. adenophyllum

is cultivated occasionally elsewhere in the tropics. Two trees of this species are
cultivated at Summit Gardens.
CANAL ZONE: Summit Gardens, Gentry 1446A (MO, SCZ).
15. JACARANDA
Jacaranda Juss., Gen. P1. 138. 1789. TYPE: J. caerulea (L.) St. Hilaire.
Iacaranda Persoon, Syn. Mant. 2: 173. 1805, sphalma.

Kordelestris Arruda in Koster, Trav. Braz. 1: 500. 1816, fide Index Kewensis.
Rafinesquia Raf., Sylva Tell. 79. 1838, non Nutt. TYPE: R. caerulea (L.) Raf. = Jacaranda
caerulea (L.) St. Hilaire.
Etorloba Raf., Sylva Tell. 79. 1838, nom. alt. for Rafinesquia: "another beautiful G. that I
dedicate to myself if the Rafinesquia of New Flora 600 is not deemed good enough, and
I add a second name meaning 'heart pod,' if any one cavils at this again."-Rafinesque.
Pteropodium DC. ex Meissn., Gen. 300; Comm. 209. 1840. LECTOTYPE: P. glabrum DC.-
Jacaranda glabra (DC.) Bur. & K. Schum.
5'A
FIGURE 14. Haplophragma adenophyllum (Wall. ex D. Don) Dop.-A. Habit (X 1/4).

-B. Fruit (X 1/4). [After Gentry 1446A (MO).]
Mostly medium-sized to large trees (some Brazilian species are shrubs or

subshrubs). Leaves usually bipinnate (rarely pinnate or even simple in some
Brazilian species). Inflorescence a few- to many-flowered terminal or axillary
panicle, sometimes from the old wood. Flowers with the calyx short and broadly
campanulate to cupular, more or less truncate, usually 5-denticulate or acutely
5-lobed; corolla blue or blue-purple tubular-campanulate, pubescent to essentially
glabrous outside; anthers glabrous, often 1-thecate, the staminode elongate, ex-
ceeding the stamens, variously glandular-pubesc

grains 3-colpate, the exine smooth; ovary flatten
bescent (even in the same species), the ovules c
pulvinate. Fruit an oblong capsule strongly flattened perpendicularly to the
septum, dehiscing perpendicularly to the septum and parallel to the plane of
compression, the valves variously glabrous or lepidote, often with undulate mar-
gins; seeds thin, winged, the wings membranaceous, hyaline or brownish.
About 30 species of tropical America ranging from southern Mexico and the
West Indies to Argentina, widely cultivated in the Old World. Opposite bicom-
pound leaves distinguish most species of Jacaranda from other New World trees
of the Bignoniaceae.
Useful reference:
Dugand, A. Sobre Jacaranda (Bignoniaceae) de Colombia y Venezuela.
Mutisia 23: 1-16. 1954.
a. Corolla tube densely puberulous outside; fruit drying black or brown without frecklin
of paler spots, the margin not undulate.
b. Calyx tubular-cupular; inflorescence a racemose panicle; fruit oblong; leaflets larg
(more than 1.5 cm long and 0.8 cm wide); native, rarely cultivated - 2. J. copaia
bb. Calyx short, broadly campanulate; inflorescence an open panicle; fruit suborbicular;
leaflets small (less than 1.2 cm long and 0.4 cm wide); cultivated exotic, mostly at
higher elevations -3. J. mimosifolia
aa. Corolla tube glabrate outside, glandular
with a freckling of tan or whitish spots, th
------------------------------------------------------------------ - -------1. J. caucana
1. Jacaranda caucana Pittier, Contr. U.S. Natl. Herb. 18: 258. 1917. TYPE:
Colombia, Cauca, Pittier 925 (US).
J. filicifolia D. Don sec. Seem., Bot. Voy. Herald 181. 1854, non Don.
J. gualanday Cortes, Fl. Col. 99. 1897, nomen nudum.
J. trianae Kranzl., Fedde Repert. 17: 226. 1921. TYPE: Colombia, Cundinamarca, Triana s.n.
(G).
la. Jacaranda caucana Pittier subsp. sandwithiana A. Gentry.5 TYPE: Panama,

Canal Zone; cultivated at Summit Gardens, Gentry 744 (MO).-FIG. 15.
Tree to at least 39 cm d.b.h. and 25 m tall; the trunk usually somewhat flat-
tened, bark light gray, smooth; twigs lepidote, subterete, gray with somewhat
elongate whitish lenticels. Leaves pinnately bicompound, 24-43 cm long, with
8-18 pinnae, each pinna 10-13.2 cm long with winged rachis and 11-23 sessile
leaflets, these 0.8-3.0 cm long and 0.4-1.0 cm wide, in the shape of a parallelo-
gram with the midvein connecting the 2 acute angles except the rhombic terminal
one, 3-3.5 cm long and 1.3-1.8 cm wide, apically obtuse and apiculate, basally
abruptly wide-cuneate, membranaceous, secondary veins in 4-7 pairs, lepidote
above and beneath, pubescent beneath especially along the veins with long simple
trichomes and sparsely above with short simple trichomes, drying brownish-green
5Jacaranda caucana Pittier subsp. sandwithiana A. Gentry, subsp. nov. Ab subspecie

typica foliolis majusibus, pinnis plus separatis, lobis calycis abbreviatis differt.
A:;~~~~~~~~
FIGURE 15. Jacaranda cau~cana subsp. sandwithiana A. Gentry.-A. Corolla slit open
( X 3/5).-B. Leaf ( Xh).- C. Fruit ( X hi.[After Gentry 1939 (MO).] D. Staminode
(X 14/5).-E. Ovary and disc ( X 4%7).-F. Anther ( X 12).-G. Inflorescence (X ).
H. Ovary cross section ( X 74/5). [After Gentry 744 (MO).]
above, grayish beneath; petiole 4.5-7 cm long, the petiolules 4-10 mm long,
petiole and petiolules lepidote and sparsely simple pubescent as the main and
lateral rachises. Inflorescence a cluster of 3-27-flowered panicles on older twigs,
the pedicels and peduncle simple-puberulous. Flowers with a delicate sweet
fragrance; calyx reduced, broadly campanulate, subtruncate to bluntly 5-toothed,
simple-puberulous and scattered-lepidote, 1.5-3 mm long and 3-4 mm wide;
corolla purplish-blue outside and on the lobes and abaxially on throat inside,
white at base of throat inside and adaxially to the base of the 2 adaxial lobes,
tubular-campanulate above a 5-9 mm X 5-7 mm basal bulge and 3-4 mm X 1.2
mm neck which bends 900, 3.64.8 cm long and 1.1-2.0 cm wide at the mouth,
the tube 3.0-3.6 cm long, the lobes 0.7-1.0 cm long; the tube mostly glabrous
outside, glandular-puberulous on the neck, the adaxial lobes and tube densely
minute-lepidote inside, the tube also with a few long simple trichomes in the
throat and glandular-puberulous with 1-3-celled trichomes at the level of stamen
insertion and in the neck, the abaxial lobe pubescent inside with long, simple or
forked trichomes, the others often ciliate; stamens didynamous, the anthers
1-thecate, 1.5-2 mm long, the anterior filaments 1.2-1.5 cm long, the posterior
filaments 1.1-1.4 cm long, inserted 8-15 mm from the base of tube; the staminode
2.2-2.5 cm long, glandular-pubescent with long (to 2 mm) 1-6-celled trichomes
for the apical 2-3 mm and central 5-7 mm; pistil 2.0-2.6 cm long, bent to follow
the posterior contour of the corolla, the ovary flattened-cylindrical, 1.5-2 mm
long and 1.5-2 mm wide, 1-1.5 mm thick, appressed-pubescent with simple uni-
cellular trichomes, the ovules 4-6-seriate in each locule; disc pulvinate, 1-1.5 mm
long, 1-2 mm wide. Capsule compressed-oblong with undulate margins at ma-
turity, 4.6-8.5 cm long and 3.8-6.0 cm wide, lepidote, drying dark with slightly
raised lighter spots; seeds 0.8-1.6 cm long and 2.3-4.1 cm wide, the wings brown
to subhyaline, the body slightly thicker and relatively woody, not distinctly
demarcated.
In Panama this species has been collected only on the Pacific slope, mostly
in the tropical moist forest. It is widely cultivated in the lowlands and its natural
range is much obscured. The species is apparently disjunct across most of Panama
between Darien and Chiriqui Provinces. It flowers in the dry season with an
extended flowering period lasting more than a month. The fruit takes just over
a year to mature, and seed release is usually in late dry season.
Jacaranda caucana ranges from southeastern Costa Rica to the valleys of the
Rio Cauca and Rio Magdalena in Columbia. Its close relative, J. hesperia Du-
gand, occurs along the Pacific lowlands of Colombia in the departments of Valle
and Choco and northward in the Atrato Valley into Antioquia Province south of
Turbo.
On the basis of the Colombian material alone, the separation between J. hes-
peria and J. caucana is easily justified. Some of the most important differences
by which J. hesperia is separated from J. caucana include larger leaflets and more
widely separated pinnae, much larger fruits with straight' margins, reduction of
the glandular pubescence at the corolla base to glandular-lepidote scales, and a
different ecology. The Panamanian material is intermediate between the two

Colombian species. It has the smaller, undulate-margined fruits with light spot-
ting and the glandular pubescent corolla tube of J. caucana but the larger leaflets
and widely spaced pinnae of J. hesperia. The Panamanian plants also differ
notably from both J. caucana and J. hesperia in the short-lobed, sometimes sub
truncate calyx.
All but one of the Panamanian and Costa Rican specimens come from
areas of tropical moist forest, the single exception being from a transitional
area at Caldera, Chiriqui. Thus the Darien plants represent not a con-
tinuation of the range of J. hesperia but a distinct population separated by the
wet forest area near the Colombian border. While the habitat of J. hesperia is
different from that of J. caucana as it occurs in Central America, the valleys of
the Magdalena and Cauca Rivers in which typical J. caucana occurs arte quite
dry in places despite their greater elevation. Thus the Central American plants
agree with typical J. caucana in everything but their leaves and calyces.
If variations in leaf dimensions are considered not to merit specific distinc-
tion, then only shorter calyx lobes remain to differentiate the Central American
plants from J. caucana. There is sufficient variation in calyx lobe length, es-
pecially in specimens from the Darien, that I would consider this character, too,
unworthy of specific recognition. The Panamanian plant should rather be
regarded as a distinctive subspecies of J. caucana differentiated by its shorter
calyx lobes and larger leaflets. I have named this variety for the late N. Y. Sand-
with in recognition of his early understanding of its correct relationship. This is
clearly documented by the correspondence between Sandwith and Dugand, which
is filed in the herbarium at Kew. It is evident from this correspondence that
Sandwith, though accepting J. hesperia, recognized the difficulty of assigning
the Central American plants to it. In a marginal note next to Dugand's discussion
of differences in fruit size he reminded himself to check the Sutton Hayes speci-
men in the carpological collection at Kew. After" doing this he added in a second
note that the Hayes fruit matched J. caucana rather than J. hesperia. Although
Sandwith had access to only a few Central American collections he clearly under-
stood the problem and his use of "Jacaranda aff. caucana" in identifying such
material further underscores his grasp of the situation, although he failed to give
the Central American plants formal taxonomic recognition.
CANAL ZONE: Summit Garden, Croat 11201 (MO); Gentry 1939 (MO, SCZ), 3691, 4961
(both MO). Near Paraiso Station, Panama Railroad, Hayes 12 (BM, BR, K). Fort Clayton,
Tyson 3463 (SCZ). Vicinity of Miraflores Locks, White & White 190 (GH, US). CHIRIQUI:
Quebrada Quanabanito W of Puerto Armuelles, Croat 22080 (MO). Quebrada Melliza,
Burica Peninsula, Liesner 473 (MO). Caldera (Boquete) near banks of Rio Chiriqui Viejo,
Maurice 866 (US). Vicinity of San Lorenzo, Seemann 1127 (BM, K). DARIEN: Rio Chico,
Allen 4596 (MO). Cerro Piriaque, Duke 8129 (MO). Rio Tuira between Rios Paya and
Pucro, Duke & Kirkbride 14055 (MO, SCZ). Isla Cartagena near La Palma, Gentry 3934
(MO). Rio Tuira between Rios Paya and Aspave, Gentry 4411 (MO). Below Boca de Cupe
on Rio Tuira, Gentry 4487 (MO). El Real, Gentry 4531 (MO). Rio Tuira between Sumacate
and Chepigana, Pittier 6622 (US). Vicinity of El Real, Stern et al. 743 (GH, K, US). Cerro
Piriaque, Tyson et al. 3847 (MO). PANAMA: Vicinity of Arraija'n, Allen 1763 (F, GH, NY,
US). Along highway between Chorrera and Bejuco, Croat 14413A (MO).
2. Jacaranda copaia (Aubl.) D. Don, Edinburg Phil. Jour. 9: 267. 1823.

Bignonia copaia Aubl., Hist. P1. Guiane Fr. 2: 650, tab. 262, fig. 1, tab. 265. 1775. TYPE:
French Guiana, Aublet s.n. (P-AD, 12304).
B. procera Willd., Sp. P1. 3: 307. 1801. TYPE: Based on B. copaia Aubl.
Jacaranda procera (Willd.) R. Br., Bot. Mag. tab. 2327. 1822.
J. spectabilis Mart. ex DC., Prodr. 9: 229. 1845. TYPE: Brazil, Amazonas, Martius, 1819 (M).
J. superba Pittier, Bol. Soc. Venez. Ci. Nat. 6: 19. 1940. TYPE: Venezuela, Bolivar, Llewelyn
Williams 11537 (US).
Tree to 45 cm d.b.h. and 45 m tall, the trunk usually unbranched for

of its length and often somewhat flattened, young trees unbranche
cluster of giant leaves at top, bark gray, smooth; twigs lepidote, subtet
drying brown or blackish, the pith large, sometimes hollowed by ants.
pinnately bicompound, 15-165 cm long, with 5-20 pinnae, each pinna 5-3
long with essentially wingless rachis and 5-25 sessile leaflets, these 1
long and 0.8-2.5 cm wide, asymmetrically rhombic-elliptic, acute to acum
asymmetrically attenuate, membranaceous, secondary veins 3-5 on one s
4-6 on the other, lepidote with scabrous to puberulous midvein above an
neath; drying dark brown above, olive beneath, petiole 3.5-29 cm lon
petiolules 2.5-4 cm long, petiole and petiolules lepidote to subpuberulous
the main and lateral rachises. Inflorescence a narrow terminal panic
branches lepidote and subpuberulous to puberulous. Flowers with th
cupular, more or less truncate with 5 subequal teeth, pubescent with sim
branched trichomes, 5-7 mm long and 3-5 mm wide, eglandular; corolla p
blue outside and on the lobes and abaxially on the throat inside, white a
on the throat inside and at the base of 2 adaxial lobes, tubular-campanulat
a 7-10 mm x 3-4 mm basal constriction, 3.2-5.0 cm long and 0.8-1.9
at the mouth, the tube 2.3-3.7 cm long, the lobes 0.3-1.4 mm long, densel
ulent outside with simple, forked, and mostly short dendroid trichomes, t
dendroid pubescent inside, the tube glabrous adaxially and pubescent with
simple or forked trichomes abaxially, pubescent with gland-tipped trich
the level of stamen insertion; stamens didynamous, the anthers 1-thecate
mm long, the anterior filaments 1.1-1.3 cm long, the posterior filaments
cm long, inserted 7-10 mm from the base of the tube, the staminode 2.
long, bifurcate at the tip, glandular-pubescent with long simple trichome
tip and central portion; pistil 1.5-1.8 cm long, the ovary flattened-cylin
2-2.5 mm long and 2-2.5 mm wide, 1.5 mm thick, glabrous, the ovules m
less 8-seriate in each locule; disc pulvinate, 1 mm long and 2.5 mm wide.
compressed-oblong, the margins straight, not wavy, 6.2-12.7 cm long and
cm wide, somewhat lepidote to glabrous, drying brown or blackish; seed
bodied with a surrounding suborbicular wing, 1.0-2.0 cm long and 1.7
wide, the body 3-5 mm long and 3-4 mm wide, the wing hyaline-m
naceous, with radial brownish striations, clearly demarcated from the see
This is a widespread tree of the tropical moist forest, premontane wet

and tropical wet forest. It ranges from British Honduras to Brazil, Peru,
Guianas. It flowers during the dry season from February to April, a
release is in mid wet season in late July and early August. Juvenile t
extremely abundant in second growth situations and are one of the characteristic
elements of wet forest second growth. Their terminal tuft of leaves on a long
slender stem (a habit shared with juveniles of Schizolobium parahybum (Vell.)
Blake, which are common in the same associations) provides part of the typical
physiognomy of young second growth. In mature forest young trees survive only
in tree-fall clearings, although seedlings sprout under the forest canopy.
The cupular calyx of this species distinguishes it from the other Central
American species of Jacaranda. Its larger leaflets also distinguish it vegetatively
from J. caucana and the cultivated J. mimosifolia.
BOCAS DEL TORO: Region of Almirante, Cooper s.n. (F). CANAL ZONE: Barro Colorado
Island, Croat 6782 (MO, NY, SCZ), 7888 (MO, SCZ), 10187, 10820, 11331, 12196, 15585,
16211A, 16518A (all MO); Foster 1429 (MO, PMA); Gentry 425 (MO, WIS); Kenoyer 525
(US); Shattuck 197 (A, MO), 672, 781 (both MO); Standley 41025 (US); Wetmore &
Abbe 104 (A, GH, MO); Zetek 5085 (MO). Near Pedro Miguel, Allen 729 (A, -GH, MO,
US), 4465 (NY), 4466 (GH, U). Vicinity of Madden Dam, Allen 4467 (K, MO). Monte
Lirio, Christopherson 121 (NY, US). Las Cruces trail in Madden Forest, Gentry 1377 (MO),
1957 (MO, SCZ). Pipeline Road, Gentry 1798 (MO). Paraiso Station, Panama Railroad,
Hayes 627 (BM, BR, K). Summit, Higgins 309 (US). N of Frijoles, Standley 27513 (US).
COLON: Santa Rita Ridge, Gentry 1867 (MO). DARIEN: Vicinity of Pinogana, Allen 4431
(K, MO, US). Near Santa Fe, Duke 12281 (MO). S of La Palma, Gentry 4294 (MO). Near
Cerro Pirre on Rio Parasenico, Gentry 4717 (MO). Piriaque at River Lara, Tyson et al. 4759,
4768 (both SCZ). PANAMA': Vicinity of National Airport E of Panama City, Allen 4460
(BM, BR, MO). Cerro Campana, Gentry 1828 (MO). Cerro Jefe, Gentry 2105 (MO). Rio
Maestra, Gentry 2217, 2218 (both MO). Rio Pasiga, Gentry 2255, 2269 (both MO). Rio
Agua Clara, Gentry 2603 (MO). Along trail between Rio Espave and Rio Bayano, Gentry
7752 (MO). 10-15 km from Rio Bayano crossing on trail to Santa Fe, Gentry 3820 (MO).
Between Rio Sulugandi and Rio Espave, Gentry 4982 (MO). W of El Llano on road to
Carti, Gentry 5089 (MO). San Jose Island, Johnston 635 (GH). SAN BLAS: Perme, Cooper
631 (BM, K, NY, US). Mountains above Puerto Obaldia, Gentry 1490 (MO). Mainland
opposite Playon Chico, Gentry 6370 (MO).
3. Jacaranda mimosifolia D. Don, Bot. Reg. Tab. 631. 1822. TYPE: (not seen).
J. ovalifolia R. Br., Bot. Mag. tab. 2327. 1822. TYPE: (not seen).
J. chelonia Griseb., P1. Lorentz. 175. 1874, non Symb. Fl. Argent. 258. 1879. TYPE: (not seen).
Tree to at least 15 m tall and 45 cm d.b.h., the bark gray; branchlets terete.
Leaves pinnately bicompound, 15-30 cm long, with 13-31 pinnae, these 1.3-2.1
cm apart on the rachis, each pinna 5-10 cm long with winged rachis and 13-41
sessile leaflets, these 3-12 mm long and 1-4 mm wide, narrowly elliptic, sharply
acuminate, the base cuneate, chartaceous, glabrous or slightly puberulous along
midrib and margin, the margin slightly revolute. Inflorescence an open terminal
panicle, the branches shortly puberulous. Flowers with the calyx reduced, broadly
campanulate, 5-denticulate, 1 mm long and 1.5-2 mm wide, subpuberulous at
least along margin; corolla purplish-blue with the tube white inside, tubular-
campanulate above a slightly narrowed neck which is inconspicuously curved
and slightly enlarged toward the base, 3-4 cm long and 0.7-1.2 cm wide at the
mouth of tube, the tube 2.4-5.2 cm long, the lobes 0.3-0.5 cm long pubescent
outside, especially toward the base, pubescent inside at the level of stamen
insertion and sparsely so with long trichomes in the tube; stamens didynamous,
the anthers 1-thecate, the second theca reduced to a minute appendage, each
theca 2 mm long, the anterior filaments 1.4-1.8 cm long, the posterior filaments
1.1-1.4 cm long, the filaments pubescent below the middle, inserted 1.1-1.2 cm
from the base of tube, the staminode 2.0-2.5 cm long, the middle portion and tip
glandular-pubescent, inserted 0.9-1.0 cm from the base of the tube; pistil 2.2-2.5
cm long, the ovary flattened-cylindrical 3-3.5 mm long and 1.5-2 mm wide, gla-
brous, the ovules 6-8-seriate in each locule; disc short, pulvinate. Capsule com-
pressed-orbicular, often shallowly emarginate at the tip and truncate to broadly
but shallowly subcordate basally, 3.2-5.8 cm long and 3.7-5.5 cm wide, drying
reddish brown, the margins not undulate, seeds thin, 0.9-1.2 cm long and 1.1-1.7
cm wide, the wing more or less surrounding and clearly demarcated from the
seed body, hyaline-membranaceous with brownish streaks.
This species is commonly cultivated in the Costa Rican uplands and probably
also in Chiriqui Province, although I have seen no Panamanian specimens. As
interpreted by Sandwith (Kew Bull. 1953: 455. 1954) it has a restricted natural
range in northwestern Argentina. However, it is widely cultivated throughout
the tropics and subtropics of the world. It is closely related to the earlier-de-
scribed J. acutifolia H.B.K., from which it can be distinguished by its larger
fruits, shorter calyx, and greater number of pinnae per leaf.
The naming of this species represents a fantastic coincidence in which the
same plant was described in two different publications under two different names
on the same day (June 1, 1822). Don was the first to unite J. ovalifolia with his
J. mimosifolia so his choice of names must be accepted.
16. KIGELIA
Kigelia DC., Rev. Bign. (Bibl. Univ. Geneve) 19. 1838, nom. cons.
Medium-sized trees with spreading crowns. Leaves imparipinnately com-

pound, usually 7-9-foliolate. Inflorescence a pendulous racemose panicle with
the central axis greatly elongated, hanging flagelliflorously below lower branches.
Flowers with the calyx large, cupular, irregularly 5-toothed; corolla brownish
maroon, campanulate above a tubular base; stamens more or less didynamous,
the anther thecae not divergent; ovary cylindrical, lepidote; disc large, annular-
pulvinate. Fruit very large, indehiscent, cylindrical-oblong, fibrous and woody,
to 12 cm or more in diameter, commonly to 50 cm long; seeds flat, wingless, the
cotyledons folded in half along their central axis.
Probably only one species of tropical African savannas.
1. Kigelia pinnata (Jacq.) DC., Rev. Bign. (Bibl. Univ. Geneve) 24. 1838.
-FIG. 16.
Crescentia pinnata Jacq., Collect. 3: 203, tab. 18. 1791. TYPE: (not seen).
Tanaecium pinnatum (Jacq.) Willd. in L. Sp. P1., ed. 4. 3: 312. 1802.
Tree to at least 15 m tall and 50 cm d.b.h.; crown spreading; twigs pubescent,

often hollow. Leaves imparipinnately compound, mostly 7-9-foliolate, the leaflets
opposite or subopposite, each leaflet elliptic to elliptic-obovate, rounded at the
tip and base, 1.9-11.6 cm long and 1.4-6.8 cm wide, proximal leaflets smallest
FIGURE 16. Kigelia pinnata (Jacq.) DC.-A. Inflorescence (X 1/2). -B. Leaf (X 1/2).
[After Tyson & Blum 3454 (MO).]-C. Fruit (x %). [After Gentry s.n., Florida (MO).]
and the terminal leaflet largest, the surface strongly minutely papillose and ir
regularly lepidote, pubescent with simple trichomes on main veins above and
beneath and to some extent over the undersurface, petiole 3.3-4.0 cm long.
Inflorescence a pendent simple panicle with the central axis greatly elongated,
flagelliflorous, hanging below lower branches. Flowers with the calyx cupular,
irregularly 5-dentate, 2.5-4.5 cm long and 1.6-2.7 cm wide, glabrous or sub-
puberulous, usually with glandular fields in the upper part; corolla deep maroon
inside, maroon streaked with green outside, campanulate above a tubular base,
glabrous with a glandular epidermis, pubescent at the level of stamen insertion,
7-9 cm long and 4-6 cm wide at the top of the tube; stamens didynamous, subex-
serted, the anther thecae slightly divergent, 7-9 mm long and 1.5 mm wide, the
filaments basally pubescent, the longer pair 4.9-5.1 cm long, the shorter pair ca.
4.3-4.5 cm long, the staminode 4-5 mm long, inserted 1.2-1.3 cm above base of
tube; pistil 6-6.5 cm long, the ovary cylindrical, 10-11 mm long and 3 mm wide,
more or less glandular-lepidote, unilocular with 2 bifid parietal placentae at the
top, bilocular with 2 axile placentae in each locule for most of the length, the
ovules ca. 10-seriate in each locule; disc very large, annular-pulvinate, 3 mm
long and 12-13 mm wide. Fruit cylindric with rounded ends, large, pendent,
indehiscent, to at least 80 cm long and 8 cm wide, rarely set in Panama.
Native to the wooded savannas of East Africa, this species is widely-cultivated

as an exotic ornamental throughout the tropical and subtropical regions of the
world. The pendulous inflorescence with its large off-maroon flowers makes the
"sausage tree" distinctive.
CANAL ZONE: Summit Garden, Croat 14539 (MO). Balboa near firehouse, Standley
26895 (MO). Ancon, Gentry 6053 (MO); Tyson & Blum 3454 (MO, SCZ).
17. LUNDIA
Lundia DC., Rev. Bign. (Bibl. Univ. Geneve) 11. 1838, nom. cons. TYPE: L.
glabra DC.
Phoenicocissus Mart. ex Meisn., Gen. Comm. 209. 1840.

Craterotecoma Mart. ex DC., Prodr. 9: 215. 1845. TYPE: C. parviflora DC. = L. erionema DC.
Lianas; stems with 4 phloem arms in cross section; branchlets terete with
interpetiolar glandular fields; pseudostipules inconspicuous or lacking. Leaves
3-foliolate, the terminal leaflet often replaced by a simple or trifid tendril.
Inflorescence an axillary or terminal panicle. Flowers with the calyx closed and
pointed in bud, splitting calypterately, or sometimes bilabiately, cupular and
truncate (or bilabiate) at maturity; corolla white to magenta, tubular-infundib-
uliform to tubular-campanulate, pubescent outside; anthers densely pubescent,
the thecae straight, divaricate, the filaments glabrous or pubescent; pollen grains
single, 3-colpate, the exine microreticulate; ovary oblong-conical, densely pubes-
cent, the ovules 2-6-seriate in each locule, the style usually more or less pu-
bescent; disc lacking. Fruit an elongate-linear more or less compressed capsule,
the valves parallel to the septum, pubescent, the median nerve conspicuously
raised, and also to some extent the margins or an area near them; seeds thin,
bialate, the wings membranaceous, irregularly hyaline at least at tips.
About 12 species of tropical America ranging from Guatemala and British

Honduras to Brazil and Bolivia.
Flowering Lundia can be readily recognized by the pubescent anthers. The
pubescent fruit is also a good character. When sterile, the slightly ciliate, widely
ovate, truncate leaves distinguish Lundia from most other Bignoniaceae vines.
Its much less pronounced pubescence, blunt (not acuminate) pseudostipules, and
FIGURE 17. Lundia.-A. L. corymbifera (Vahl) Sandw., hab

5565 (MO).] B-H. L. puberula Pittier.-B. Corolla ( X 3/5).-C
-D. Fruit (X1/) [After Gentry 4230 (MO).] E. Calyces (X 3/5).-F. Anther (X 41/C).
-G. Ovary cross section ( X 15).-H. Ovary ( X 6). [After Gentry & Dressler 1986 (MO).]
mostly wet forest habitat differentiate this genus from the vegetatively similar
Arrabidaea mollissima.
a. Corolla pink to magenta; style shortly puberulous at the apex or throughout; calyx dry-
ing dark, often bilabiate --------------------------------------------2. L. puberula
aa. Corolla white; style glabrous except at the extreme base; calyx drying tan, almost always
evenly truncate ---- 1. L. corymbifera
1. Lundia corymbifera (Vahl) Sandw., Recueil Trav. Bot. Neerl. 34: 229.
1937.-FIG. 17A.
Bignonia corymbifera Vahl, Eclog. Amer. 2: 45, pl. 17. 1798. TYPE: Trinidad, von Rohr 6 (C).
B. umbrosa H.B.K., Nov. Gen. Sp. P1. 3: 138. 1819. TYPE: Colombia, Caripa, Humboldt &
Bonpland 279 (P).
Lundia umbrosa (H.B.K.) Bur., Adansonia 8: 282. 1868.
Arrabidaea corymbifera (Vahl) Bur. ex K. Schum. in Engler & Prantl, Nat. Pflanzenf. 4(3b):
213. 1894.
Cuspidaria corymbifera (Vahl) Baill. ex Schum. in Engler & Prantl, Nat. Pflanzenf. 4(3b):
216. 1894.
Lundia phaseolifolia Rusby, Mem. Torrey Bot. Club 6: 99. 1896. TYPE: Bolivia, Tipuani-
Guanae, Bang 1650 (A, GH, K, MO, NY, US).
L. truncata Rusby, Mem. New York Bot. Gard. 7: 356. 1927. TYPE: Bolivia, Rurrenabaque,
Cardenas 2036 (IGH, K, NY, US).
L. valenzuelae Dugand, Mutisia 10: 7. 1952. TYPE: Colombia, Cundinamarca, Fernandez &
Mora 1369 (COL).
Liana to 2.5 cm in diameter; stems with 4 phloem arms in cross section;

branchlets terete, drying longitudinally striate, simple puberulous, the nodes with
interpetiolar glandular fields; pseudostipules inconspicuous, blunt-conical. Leaves
3-foliolate or 2-foliolate with a tendril or tendril scar; leaflets ovate to elliptic-
obovate, acute to acuminate, the base truncate, 6.5-11.5 cm long and 3.3-7.6 cm
wide, membranaceous, palmately veined basally, the secondary veins 4-6 on a
side, short puberulous at least on main veins above and below and in axils of the
lateral nerves beneath, sometimes also with scattered trichomes over the surface,
usually more or less lepidote, some plate-shaped glands near the midvein be-
neath, lepidote glandular basally, drying dark green to brown above, grayish
green beneath, never with a macroscopically conspicuous white fringe when
young although microscopically ciliate; tendril simple 4-14 cm long; petiolules
1.2-3.2 cm long, petiole 3.2-5.3 cm long, conspicuously simple puberulous. In-
florescence a contracted axillary or terminal panicle, the branches strongly simple
puberulous. Flowers with the calyx in bud terminating in 5 elongate, fused teeth
which form an apical point, calypterate with the apical third dehiscent, thus
cupular and evenly truncate in mature bud or flower, 3-6 mm long and 4-5 mm
wide, puberulous with 1-3-celled simple trichomes; corolla white with the 2
throat ridges yellow, tubular-infundibuliform, 2.5-4.3 cm long and 0.8-1.3 cm
wide at the mouth, the tube 1.8-2.8 cm long, the lobes 0.4-1.2 cm long, densely
simple puberulous outside, the lobes ciliate margined, glandular-lepidote and
puberulous with simple trichomes, the throat mostly glabrous inside, short-puber-
ulous with gland-tipped trichomes at and below the level of stamen insertion;
stamens didynamous, the anther thecae divergent to divaricate, 2-3 mm long, the
longer filaments 1.3-1.6 cm long, the shorter filaments 0.8-1.2 cm long, the
staminode 2-3 mm long, inserted 3-7 mm from base of corolla tube; pistil 1.5-2.4
cm long, the style glabrous except for short appressed trichomes near the base,
the ovary tapered cylindric, 2-3 mm long and 1-1.5 mm wide, densely sericeous,
the ovules 4-seriate in each locule; disc inconspicuous, 0.5 mm long and 1.5 mm
1.973] GENTRY-FLORA OF PANAMA (Family 172. Bignoniaceae) 869
wide. Capsule linear, more or less compressed, velvety

and 1.5-1.9 cm wide, both median nerve and margins distinctly raised; seeds
thin, bialate, 1.0-1.3 cm long and 3.4-3.9 cm wide, wings hyaline at the tips,
brown at the base, poorly demarcated.
Locally common in tropical wet forest, L. corymbifera occurs also in premon-

tane wet forest and parts of the tropical moist forest. It is apparently restricted
to the Pacific slope in Central America and ranges from Costa Rica to Bolivia and
Brazil. It flowers mostly during the wet season from April or June to October.
Lundia corymbifera is vegetatively different from closely related L. puberula
in a shorter puberulence, the leaves grayish green below, and branchlets usually
drying tannish (not dark). The two species are extremely close and can be dis-
tinguished most easily by flower color. The paucity of material of both species
makes an understanding of specific delineations difficult.
Lundia valenzuelae Dugand of Colombia is apparently a pubescent variant
of this species.
CHIRIQUI: Monteverde 2.5 kin W of Puerto Armuelles, Liesner 54 (MO). Rio Chiriqui
to Remedios, Woodson et al. 1178 (A, K, MO, NY). Vicinity of Puerto Armuelles, Woodson
& Schery 811 (GH, MO). Vicinity of Bartolome, Peninsula de Burica, Woodson & Schery
915 (GH, MO, US). DARIEN: Between Rios Punusa and Pucro, Duke 14641 (MO). PANAMA:
Rio Maestro, Gentry 2236 (MO). 2 mi. E of El Llano, Gentry 5565 (MO).
2. Lundia puberula Pittier, Contr. U. S. Nail. Herb. 18: 258. 15 Sept. 1917.
TYPE: Panama, Darien, Pittier 5499 (F, K, MO, US).-FIG. 17B-H.
L. dicheilocalyx Blake, Contr. Gray Herb. 52: 94. 28 Sept. 1917. TYPE: British Honduras,
Toledo, Peck 495 (GH, K).
L. schumanniana Kranzl., Fedde Repert. 17: 120. 1921. TYPE: Mexico, Campeche, (not seen).
L. colombiana Dugand, Caldasia 4: 236. 1946. TYPE: Colombia, Meta, Dugand & Jaramillo
3111 (COL, NY).
Liana to at least 2.5 cm in diameter; stems with 4 phloem arms in

tion; the branchlets terete, puberulous, nodes with interpetiolar gland
pseudostipules inconspicuous, blunt. Leaves 3-foliolate or 2-foliolate with a ten-
dril or tendril scar; leaflets ovate, long-acuminate, basally truncate to asymmetri-
cally cordate, 7.4-9.2 cm long and 4.6-5.7 cm wide, membranaceous, palmately
veined at the base, the secondary veins 3-5 on a side, sparsely subpuberulous at
least on the main veins above and beneath with a few longer trichomes in the
axils of lateral nerves beneath and scattered lepidote scales mostly beneath, the
margins strongly ciliate, plate-shaped glands near the midvein beneath, lepidote-
glandular basally, drying olive brown to reddish-brown; tendril simple, 5-10 cm
long; petiolules 2-2.3 cm long, the petiole 2.4-2.9 cm long, puberulous. Inflores-
cence an axillary or terminal panicle, the branches puberulous. Flowers with the
calyx in young bud conical-pointed, at length truncate, the tip calypterate and
evenly dehiscent, or 2 (-3) -labiate and the tip splitting, 4-6 mm long and 3-4
mm wide, subpuberulous; corolla pale pink or lavender to magenta, tubular-
campanulate or infundibuliform, 3.1-5.8 cm long and 0.7-1.5 cm wide at the
mouth, the tube 2.5-4.0 cm long, the lobes 0.8-2.1 cm long, densely simple puber-
ulous outside, the lobes ciliate-margined, glandular-lepidote and puberulous with
simple trichomes inside, the throat glabrous inside or with a few scattered gland-
tipped trichomes at and below the level of stamen insertion; stamens didynamous,
the anther thecae divaricate, 3-4 mm long, the longer filaments 1.5-2.1 cm long,
the shorter pair 1.0-1.3 cm long, the staminode 1-2 mm long, inserted 11-15 mm
from the base of corolla tube; pistil 2.9-3.2 cm long, the style somewhat puber-
ulous at least toward the apex, the ovary cylindric, 2.5-3.0 mm long and 1 mm
wide, strongly appressed puberulous, the ovules 4-seriate in each locule; disc
absent. Capsule linear, the valves somewhat compressed, the midvein conspicu-
ously raised, the margins raised but rounded, velvety pubescent, drying blackish,
25-45 cm long and 1.5-1.6 cm wide; the seeds thin, bialate, 0.9-1.2 cm long and
3.3-3.5 cm wide, the wings membranaceous, basally brown, dirty-hyaline at the
tip, not clearly demarcated from the seed body.
A seldom-collected species of tropical wet forest and wetter parts of the

tropical moist forest, L. puberula ranges from British Honduras and southern
Mexico to Venezuela. It has been collected in flower in November, January,
February, March, and August.
This species is similar to L. corymbifera. Although the style of L. puberula
tends to be pubescent toward the top while that of L. corymbifera is pubescent
only at the extreme base, this character is variable. The dried calyx of this species
is usually dark and bilobed, that of L. corymbifera light brown or tan and
truncate, but neither of these characters is absolutely constant. Difference in
flower color is the best character for separating the two species.
CANAL ZONE: Navy reservation N of Gamboa, Dressler 3140 (MO). Pipeline Road,
Gentry & Dressler 1986 (MO). COLON: Santa Rita Ridge, 1.6 mi. NE of Carretera
Transisthmica, Nee & Mori 3678 (MO); Gentry & Dwyer 4826 (MO, PMA). DARIEN: Ridge
from top of Cerro Pavarando to Cerro Venado and Cerro Derumba, Gentry 4230 (MO). Near
La Palma, Gentry 3933 (MO); Pittier 5499 (F, K, MO, US).
18. MACFADYENA
Macfadyena A. DC., Prodr. 9: 179. 1845. TYPt: M. uncinata (G. Meyer) A.

DC. = M. uncata (Andr.) Sprague & Sandw.
Doxantha Miers, Proc. Roy. Hort. Soc. 3: 189. 1863. TYPE: D. unguis-cati (L.) Miers -
Macfadyena unguis-cati (L.) A. Gentry.
Microbignonia Kranzl., Notizbl. Bot. Gart. Berlin 6: 380. 1915. TYPE: M. auristellae Kranzl.
Lianas; stems irregularly many-armed in cross section, the phloem arms

basically 8 but the xylem irregularly fissured as well; roots usually with con-
spicuously swollen tubers; branchlets terete; interpetiolar glandular fields present
or lacking, even in the same species; pseudostipules small, lanceolate to ovate;
juvenile plants distinctive, growing closely appressed against a supporting tree,
climbing by means of their recurved tendrils; the leaves much smaller and the
tendril relatively larger; frequently rooting from the nodes with the adventitious
roots helping to affix the slender vine to its support. Leaves 2-foliolate, often
with a trifid, uncate tendril. Inflorescence a contracted axillary few-to several-
flowered cyme or panicle, often reduced to a single flower. Flowers with the
calyx membranaceous, loosely campanulate, truncate to subspathaceous, often
irregularly lobed; corolla yellow, tubular-campanulate, glabrous outside; anthers

glabrous, the thecae straight, divaricate; pollen grains 3 colpate, the exine smooth
to microreticulate; ovary linear-oblong, minutely lepidote to subpuberulous or
glabrate, the ovules 2-4-seriate in each locule; disc more or less annular-pulvinate.
Fruit a narrow, linear, compressed capsule, the valves parallel to the septum,
smooth, the median nerve thin, slightly raised; seeds thin, bialate, the wings not
sharply demarcated from seed body, short and brown, or longer and irregularly
hyaline at tips.
Two or three species in tropical America ranging from Mexico and the West
Indies to northern Argentina and Uruguay. The two Panamanian species can be
told from the other Central American genera by their trifid, uncate tendrils which
do not become thick and woody with age. Melloa quadrivalvis is the only other
species with trifid uncate tendrils and its tendrils become conspicuously woody
with age. The juvenile plants of both M. uncata and M. unguis-cati have a growth
form quite different from the adults. The juvenile leaves are much smaller and
the plants grow strongly appressed to the bark of trees, climbing straight up by
means of hooked tendrils.
a. Calyx subspathaceously split, often with an incurved apicule; leaflets long-acuminate;

pseudostipules subulate-lanceolate; interpetiolar glandular fields conspicuous; seeds with
short wings, shiny and dark blackish brown; swamps mostly below 50 m elevation
1. M. uncata
aa. Calyx truncate or subtruncate, without an apicule; leaflets acute or short-acuminate;
pseudostipules more or less ovate, usually striate; interpetiolar glandular fields absent or
small and inconspicuous; seeds with wings longer and irregularly hyaline at tip; well-
drained sites, mostly above 50 m elevation -2. M. unguis-cati
1. Macfadyena uncata (Andr.) Sprague & Sandw., Recueil Trav. Bot. Neerl.
34: 215. 1937.-FIG. 18F-J.
Bignonia uncata Andr., Bot. Repos. tab. 530. 1808. TYPE: (not seen).
B. uncinata G. Meyer, Prim. Fl. Essequeb. 210. 1818. TYPE: Surinam (not seen).
Spathodea uncata (Andr.) Spreng., Syst. Veg. 2: 835. 1825.
S. uncinata (G. Meyer) Spreng., Syst. Veg. 2: 835. 1825.
Bignonia pachyptera DC., Rev. Bign. (Bibl. Univ. Geneve) 8. 1838, nomen nudum.
Dolichandra fenzliana Miq., Linnaea 18: 251. 1844. TYPE: Surinam, Focke 661 (U).
Macfadyena uncinata (G. Meyer) A. DC., Prodr. 9: 180. 1845.
Pachyptera puberula DC., Prodr. 9: 175. 1845. TYPE: Brazil, Cujaba, Manso 10 (G-DC).
Spathodea hispida DC., Prodr. 9: 205. 1845. TYPE: Brazil, Cujaba, Manso 105A (G-DC).
S. mollis Sond., Linnaea 22: 561. 1849. TYPE: Brazil, Regnell 11.192. (not seen).
Macfadyena fenzliana (Miq.) Miq., Stirp. Surinam. Select. 125. 1850.
M. hispida (DC) Seem., Jour. Bot. 1: 227. 1863.
M. mollis (Sond.) Seem., Jour. Bot. 1: 227. 1863.
M. pubescens S. Moore, Trans. Linn. Soc., Bot., ser. 2. 4: 418. 1895. TYPE: Brazil, Iter
Mattogrossensis, Moore 1021 (BM).
M. guatemalensis Blake, Contr. U. S. Natl. Herb. 24: 24. 1922. TYPE: Guatemala, Izabal,
Blake 7845 (US).
Liana to at least 2.5 cm in diameter, the bark fibrous, stem with phloem
irregularly many-armed in cross section; dimorphic, the juvenile plant with small
leaflets, ca. 1.5-3 cm long, climbing straight up tree trunks, often rooting at
nodes; branchlets terete, glabrous to puberulous, interpetiolar glandular fields
usually present on young twigs, rarely on fertile branches; pseudostipules subu-
FIGURE 18. Macfadyena.-A-E. M. unguis-cati ( L. ) A. Gentry.-A. Habit (X3/).

[After Woodson et al. 736 (MO).] B. Juvenile habit ( X 3/5). [After Gentry 715B (MO).]
-C. Fruit ( X 2/5). [After Croat 14062 (MO).] D. Ovary cross section ( X 15).-E. Anther
( X 4%75) .-F-J. M. uncata ( Andr. ) Sprague & Sandw.-F. Corolla split open ( X 3/5 ) .G.
Corolla (X 3/i).-H. Calyx in bud (X 3/5j).-I. Ovary cross section ( X 12).-J. Pistil and
disc ( X -3/5). [After Gentry 1745 (MO).]
late-lanceolate, ca. 2 mm long. Leaves 2-foliolate, often with a tendril, especially

when young; leaflets when young narrowly ovate to lanceoate, mucronate, basally
rounded, when mature narrowly ovate to ovate or elliptic, long acuminate, more
or less cuneate to rounded, 5.1-18 cm long and 2.0-9.2 cm wide, membranaceous,
secondary veins 5-8 on a side, minutely papillose and lepidote above and be-
neath, sometimes simple-puberulous beneath especially along main veins, and
more sparsely so above; gray green when dry, the main veins reddish or blackish
beneath; tendril trifid, uncate, 0.8-2.0 cm long to branching, the 3 arms thick-
ened, 0.5-1.1 cm long; mature petiolules 0.4-2.7 cm long, petiole 0.8-3.6 cm long,
petiole and petiolules glabrous to puberulous. Inflorescence of flowers borne
singly or in reduced few-flowered panicles in leaf axils. Flowers inconspicuously
scented, calyx membranaceous, glabrous to somewhat lepidote basally, often with
scattered glands, usually spathaceously split with apiculate incurved tip, 1.5-3.0
cm long and 0.6-1.4 cm wide, 1-2 cm from base to base of split; corolla yellow
with the longitudinal pleats in throat darker, tubular-campanulate, 4.2-8.8 cm
long and 0.8-2.0 cm wide at the mouth, the tube 3.8-6 cm long, lobes 0.7-1.7 cm
long, glabrous outside, slightly lepidote on lobes inside, sparsely pubescent with
simple trichomes along throat ridges and strongly pubescent with long, gland-
tipped trichomes and a few simple trichomes at the level of stamen insertion;
stamens didynamous, the anther thecae divaricate, 2.5 mm long, the longer fila-
ments 2.1-2.4 cm long, the shorter filaments 1.6-1.9 cm long, the staminode 1.0
cm long, inserted 15-16 mm above base of corolla tube; pistil 3.5-3.7 cm long,
the ovary linear, 2.5-3.0 mm long and 1.0 mm wide, minutely lepidote to glabrate,
the ovules 2-seriate in each locule; disc annular-pulvinate, 1 mm long and 3-4
mm wide. Capsule linear, acute, compressed, 16-26 cm long and 1.5-1.9 cm wide,
the wings woody but thin, dark brown or blackish, not demarcated from the
seed body.
Found mostly near the coast in swampy habitats of the tropical wet forest
and wetter parts of the tropical moist forest, it is especially common in the fresh
wate-r sub-mangrove association just inland from the Rhizophora-Avicennia-La-
guncularia association. It ranges from Mexico to Brazil. This species flowers
irregularly throughout the year. The light, short-winged seeds are probably
partly wind-dispersed and partly water-dispersed, being light enough to be blown
easily and thick enough to withstand continued immersion.
The spathaceously split calyx is the traditional definitive characteristic of
the species. Narrow, subulate pseudostipules, conspicuous interpetiolar glandu-
lar fields and especially its usually swampy wet forest habitat distinguish this
species from Macfadyena unguis-cati. Macfadyena uncata is less dimorphic than
M. unguis-cati, and its juvenile leaves are larger.
The supposed distinctions between Macfadyena uncata and M. mollis (an
earlier name is M. hispida) do not have specific value. Ecology of these two
so-called species seems identical. Although there may be two taxa, I cannot
at present justify recognition of more than one in Central America.
BOCAS DEL TORO: Region of Almirante, Cooper 191 (F, NY). Banks of Rio Changuinola,
Dunlap 513 (F, US). Hillside above Almirante, Gentry 2696, 2757 (both MO). Isla Colon,
von Wedel 57 (MO). Nances Cay Island, von Wedel 484 (GH, MO). Water Valley, von
Wedel 654 (MO), 1911 (GH, MO, US). Vicinity of Chiriqui Lagoon, von Wedel 1251 (GH,
MO, US). Old Bank Island, von Wedel 2110 (GH, MO, US). Fish Creek lowlands, von
Wedel 2380 (GH, MO, US). CANAL ZONE: Chagres, Fendler 207 (GH, K, MO, US), 200
(K). Along Trinidad River, Pittier 4003 (BM, NY, US). CHIRIQUI: Without locality, Hart
CLi15 (K). COLON: Camino a Portobelo despues del Rio Guanche, Correa & Dressler 1765
(MO, PMA). Vicinity of San Miguel de la Borda, Croat 9899 (MO, SCZ). W of Portobelo,
Gentry 1745 (MO). Puente de Rio Salud, Salud, Taymes & Carrasquilla 142 (MO). DARIEN:
Rio Ucurganti, Bristan 1132 (MO). Rio Jaque to base of Cerro Pavarando, Gentry 4198 (MO).
Vicinity of El Real, Gentry 4315 (MO), Gentry 4574 (MO). PANAMA: San Jose Island,
Johnston 794 (GH). SAN BLAS: Near Mandinga, Duke 8883 (MO). Near Puerto Obaldia,
Gentry 1558 (MO).
2. Macfadyena unguis-cati (L.) A. Gentry, Brittonia 25: 236. 1973.-FIG.

18A-E.
Bignonia unguis-cati L., Sp. P1., ed. 1. 2: 623. 1753. TYPE: (not seen).
B. unguiculata Vell., Fl. Flum. 234. 1825; 5: tab. 33. 1827. TYPE: Brazil (not seen).
B. exoleta Vell., Fl. Flum. 248. 1825; 6: 30. 1827. TYPE: Brazil (not seen).
B. pseudounguis Desf., Cat. Hort. Paris., ed. 3. 397. TYPE: (not seen).
B. gracilis Lodd., Bot. Cab. tab. 1705. 1825. TYPE: (not seen).
B. triantha DC., Rev. Bign. (Bibl. Univ. Geneve) 21. 1838, nomen nudum.
B. tweediana Lindl., Bot. Reg. 26: tab. 45. 1840, non Griseb. TYPE: Strangways s.n., cultivated
in England, imported from Buenos Aires (not seen).
Batocydia unguis (L. emend. DC.) Mart. ex DC., Prodr. 9: 146. 1845.
B. exoleta (Vell.) Mart. ex DC., Prodr. 9: 148. 1845.
Bignonia unguis L. emend. DC., Prodr. 9: 146. 1845.
B. unguis var. gracilis (Lodd.) A. DC., Prodr. 9: 146. 1845.
B. acutistipula Schlecht., Linnaea 26: 375. 1853. TYPE: Mexico (not seen).
Doxantha acutistipula (Schlecht.) Miers, Proc. Roy. Hort. Soc. 3: 190. 1863.
D. adunca Miers, Proc. Roy. Hort. Soc. 3: 188. 1864. TYPE: Based on Bignonia unguis Vell.
D. exoleta (Vell.) Miers, Proc. Roy. Hort. Soc. 3: 190. 1863.
D. lanuginosa Miers, Proc. Roy. Hort. Soc. 3: 190. 1863, nomen nudum.
D. mexicana Miers, Proc. Roy. Hort. Soc. 3: 190. 1863, nomen nudum. TYPE: Mexico, Coulter
1004 (not seen).
D. praesignis Miers, Proc. Roy. Hort. Soc. 3: 190. 1863, nomen nudum. TYPE: San Domingo,
Imray s.n. (not seen).
D. serrulata Miers, Proc. Roy. Hort. Soc. 3: 190. 1863, nomen nudum. TYPE: Guadeloupe,
Duchassaing s.n. (not seen).
D. tenuicula Miers, Proc. Roy. Hort. Soc. 3: 190. 1863; nomen nudum. TYPE: Hispaniola,
Swartz s.n. (not seen).
D. unguiculata (Vell.) Miers, Proc. Roy. Hort. Soc. 3: 190. 1863.
D. unguis (L. emend. DC.) Miers, Proc. Roy. Hort. Soc. 3: 190. 1863.
Bignonia inflata Griseb., Symb. Fl. Argent. 256. 1879. TYPE: Argentina (not seen, CORD
fide Fabris, Revista Mus. La Plata, Secc. Bot. 9: 391. 1965).
B. lanuginosa Hemsl., Biol. Centr. Amer. Bot. 2: 491. 1882. TYPE: Mexico, Sierra San Pedro
Nolasco, Tolaya, etc., Jurgenson 478 (K, holotype; BM, isotype).
B. unguis-cati L. var. guatemalensis K. Schum. & Loes., Bot. Jahrb. Syst. 23: 130. 1896.
TYPE: Guatemala, Retaluleu, Bernoulli & Cario 2057 (K).
B. unguis-cati L. var. serrata Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 283. 1896. TYPE:
(not seen).
B. californica Brandeg., Zoe 6: 170. 1903. TYPE: Mexico, Baja California, Purpus 249
(MO, US).
Bignonia unguis-cati var. exoleta (Vell.) Sprague, Bull. Herb. Boiss., ser. 2. 5: 84. 1905.
Doxantha unguis-cati (L.) Miers emend. Rehder, Mitt. Deutsch. Dendrol. Ges. 1913: 262.
1913.
Bignonia dasyonyx Blake, Contr. Gray Herb. 52: 93. 1917. TYPE: British Honduras, Toledo,
Peck 919 (GGH, holotype; K, isotype).
Doxantha dasyonyx (Blake) Blake, Jour. Bot. 61: 192. 1923.
D. unguis-cati var. dasyonyx (Blake) Seib., Carnegie Inst. Washington Publ. 522: 423. 1940.
D. unguis-cati var. exoleta (Vell.) Fabris, Revista Mus. La Plata, Secc. Bot. 9: 394. 1965.
Liana to 6 cm in diameter, the bark dark, fibrou

to supporting tree by adventitious rootlets; phlo
armed in cross section; dimorphic, the juvenile p
cm long and 4-8 mm wide, climbing straight up
nodes; branchlets terete, puberulus to glabrous, t
petiolar glandular fields; pseudostipules ovate, longitudinally striate. Leaves
2-foliolate, often with a tendril especially when young; leaflets when young
narrowly ovate to lanceolate, mucronate at the tip, basally rounded, when
mature narrowly ovate to ovate, acute to acuminate, cuneate to truncate, 5.0-16
cm long and 1.2-6.9 cm wide, membranaceous, secondary veins 4-6 on a side,
scattered lepidote above and beneath, otherwise glabrous above and beneath
or puberulous especially along main veins, a few plate-shaped glands near
midvein beneath, drying dark green or dark olive to almost black, the main veins
often reddish below; tendril trifid, uncate, 0.1-3.5 cm long to branching, the
3 arms thickened, 0.3-1.7 cm long; mature petiolules 0.5-2.5 cm long, petiole
1.1-4.7 cm long, glabrous or subpuberulous. Inflorescence an axillary panicle
usually reduced to 1 or 3 flowers, rarely to 15 flowers in Panama, the branches
more or less glabrous. Flowers with an odor similar to the leaves of many
Rutaceae, calyx membranaceous, cupular with a sinuous margin, subtruncate,
glabrous to slightly lepidote, often with scattered glands, 5-18 mm long and
8-18 mm wide; corolla yellow with ca. 9 orange lines in the throat, tubular-
campanulate, 4.5-10.0 cm long and 1.2-2.4 cm wide at the mouth, the tube
3.3-6.9 cm long, the lobes 1.2-3.1 cm long, glabrous outside, glabrous or
slightly simple puberulous along throat ridges inside, pubescent with simple,
jointed trichomes at the level of stamen insertion; stamens didynamous, the
-anther thecae divaricate, 3 mm long, the longer filaments 1.9-2.4 cm long, th
shorter filaments 1.2-1.8 cm long, the staminode 0.5-1.5 cm long, occasionally
developed into a fifth stamen, inserted 4-8 mm above the base of the tube;
pistil 2.8-3.8 cm long, the ovary linear, 6-7 mm long and 1.2 mm wide, minutely
lepidote to subpuberulous, the ovules 4-seriate, but appearing 2-seriate in most
cross sections; disc annular, 1-1.5 mm long and 3-4 mm wide. Capsule elongate-
linear, tapering at the ends, flattened, 26-95 cm long and 1.0-1.9 cm wide,
the surface inconspicuously lepidote, drying blackish with small whitish lenticels,
midrib not raised; seeds 1.0-1.8 cm long and 4.2-5.8 cm wide, the wings mem-
branaceous, blunt-ended, mostly brown, the extreme tips irregularly hyaline,
not sharply demarcated from seed body.
A common vine of the tropical dry forest, this species becomes progressively
less common in premontane moist forest and tropical moist forest and rare in
premontane wet forest and tropical wet forest. The species is widespread,
ranging from Mexico (including Baja California) and the West Indies to
Argentina. It flowers mostly during late dry season and early wet season from
March to June. The single collection from the premontane wet forest of
Panama (lower slopes of Cerro Jefe) was in flower in mid wet season (October).
The seeds are dispersed in mid dry season.
Seedlings of Macfadyena unguis-cati can persist for extended periods and
are one of the commonest seedlings encountered in tropical dry forest. They
normally have one or two pairs of relatively large leaves on a short upright
stem followed by production of the small-leaved juvenile growth form, which
trails across the ground until it reaches a tree upon which it can climb. This
juvenile form is also very abundant. The shift to adult growth form occurs
only after the vine escapes the shade of the forest floor and is probably a
light-triggered reaction.
When sterile this species is differentiated from M. uncata by its ovate,
longitudinally striate pseudostipules and the habitat difference. Presence or
absence of interpetiolar glandular fields is not a reliable character. Seedlings
which lack tendrils and are difficult to recognize can often be identified by
a faint sweetish odor in the crushed leaves. The thickened tap root of even
tiny seedlings with only four leaves is another character of some use in
identification.
CANAL ZONE: Barro Colorado Island, Croat 9141, 10110, 10224 (all MO), 12498 (MO,
SCZ), 12588 (MO), 14062 (MO, SCZ), 14081, 14645 (both MO); Shattuck 264 (F, MO),
944 (MO); Wetmore & Vestal 75 (F). Pipeline Road, Gentry 2094 (MO). S of Pedro
Miguel, Gentry 4865 (MO). Boy Scout road near Madden Lake, Gentry 5059 (MO). Madden
Dam, Gentry 5066 (MO). Ancon Hill, Gentry 5125 (MO). Fort Sherman, Gentry & Tyson
4837 (MO). Vicinity of Ancon, Piper 6023 (US). Corozal, Sharp s.n. (SCZ). Las Cascades
Plantation, Standley 29693 (US). DARIEN: S of Jaque, Gentry 4145 (MO). El Real, Gentry
4532 (MO). Trail up Cerro Pirre, Gentry 4591, 4606 (both MO). PANAMA: Finca Indio
on slopes of Cerro Jefe, Gentry 2148 (MO). Rio Pasiga, Gentry 2286, 2328, 2366 (all MO).
Near Rio Espave, Gentry 3697 (MO). Ca. 16 km from Bayano crossing on trail to Santa Fe,
Gentry 3807 (MO). Near Madden Lake, Gentry 5051 (MO). E of Pacora, Woodson et al.
736 (A, MO, NY). W of Campana, Woodson et al. 1308 (A, MO, NY). Isla Taboga,
Woodson et al. 1450 (A, MO).
19. MARTINELLA
Martinella Baill., Hist. PI. 10: 30. 1888. TYPE: M. martinii (DC.) Baill. ex K.
Schum. M. obovata (H.B.K.) Bur. & K. Schum.
Lianas; stems with 4 phloem arms in cross section, the pith large; branch-
lets terete, without interpetiolar glandular fields, with a conspicuously swollen
interpetiolar ridge, this less pronounced but still present when dried; pseudo-
stipules absent. Leaves 2-foliolate, often with a trifid tendril, the tendril arms
somewhat flexuous. Inflorescence an axillary raceme. Flowers with the calyx
tubular-campanulate, bilabiate or sometimes irregularly 3-4-labiate; corolla deep
wine-colored, tubular-campanulate above a narrow tubular base, glabrous to
inconspicuously lepidote outside; anthers glabrous, the thecae straight, divari-
cate; pollen grains single, 3-colpate, the exine finely reticulate; ovary linear-
cylindric, sparsely lepidote or puberulous, the ovules (2-) 4-seriate in each
locule. Fruit a strongly compressed linear capsule, the valves parallel to the
septum, thin, smooth, the median nerve indistinct; seeds thin, bialate, the
wings poorly demarcated, membranaceous, mostly pale brown, lighter at the
tips.
About two species of continental tropical America ranging from southern

Mexico to Brazil and Bolivia.
1. Martinella obovata (H.B.K.) Bur. & K. Schum. in Mart., Fl. Bras. 8(2):
161, pl. 84. 1896.-FiG. 19.
Spathodea obovata H.B.K., Nov. Gen. Sp. P1. 3: 147. 1819. TYPE: Colombia, Turbaco, Hum-
boldt & Bonpland 1391 (P).
Bignonia longisiliqua Bert. ex Spreng., Syst. Veg. 2: 830. 1825, non Vell. TYPE: Colombia;
Rio Magdalena (not seen).
B. obovata (H.B.K.) Spreng., Syst. Veg. 2: 830. 1825.
B. fockeana Miq., Linnaea 18: 609. 1844. TYPE: Surinam, Paramaribo, Focke 924 (U).
Tabebuia cordata Benth., Bot. Voy. Sulphur 129. 1844. TYPE: Panama, Darien, Barclay
s.n. (K).
Bignonia martinii DC., Prodr. 9: 152. 1845. TYPE: French Guiana, Martin s.n. (U, US).
Tabebuia latifolia DC., Prodr. 9: 213. 1845, non Rich. TYPE: French Guiana, collector un-
known (G-DC, ex P, 1821).
Macfadyena fockeana (Miq.) Miers, Proc. Roy. Hort. Soc. 3: 200. 1863.
M. obovata (H.B.K.) Miers, Proc. Roy. Hort. Soc. 3: 200. 1863.
Martinella martinii (DC.) Baill. ex K. Schum. in Engler & Pranti, Nat. Pflanzenf. 4 (3b):
216. 1894.
Anemopaegma leptosiphon Rusby, Mem. New York Bot. Gard. 7: 354. 1927. TYPE: Bolivia,
Ixiamos, Cardenas 1926 (NY).
Martinella iquitosensis Samp., Ann. Acad. Brasil Sci. 7: 122. 1935. TYPE: Peru, Loreto,
Kuhlmann 1492 (not seen).
Arrabidaea duckei Samp., Bol. Mus. Nac. Rio de Janeiro, Bot. 12: 81. 1936. TYPE: Brazil,
Ducke s.n. (R No. 28,626) (not seen).
Periarrabidaea duckei (Samp.) Samp., Ann. Acad. Bras. Sci. 12: 91. 1936.
Liana to at least 7 cm in diameter, the bark smooth, green on the younger

branches; stem with 4 phloem arms in cross section, the pith large; branchlets
terete to subterete, puberulous with short-gland-tipped trichomes to glabrate,
green and smooth when fresh, usually dark when dry, the nodes without inter-
petiolar glands or pseudostipules, usually conspicuously thickened laterally
with a raised ridge connecting opposite petioles. Leaves 2-foliolate, sometimes
with a tendril; the leaflets ovate to elliptic, acuminate to obtuse, truncate to
asymmetrically subcordate, 7.2-21 cm long and 3.3-15.5 cm wide, chartaceous,
secondary veins 5-8 on a side, the surface microscopically roughened, sometimes
indistinctly and sparsely puberulous along veins, plate-shaped glands irregularly
scattered along the midvein beneath, glossy green when fresh, drying grayish
green to olive with main veins brownish to tan beneath; tendril usually trifid,
4-23 cm long to branching, the 3 arms 0.2-2.4 cm long; petiolules and petiole
often twisted slightly, glabrous to subpuberulous, the petiolules 3.0-6.6 cm long,
the petiole 2.9-7.5 cm long. Inflorescence an axillary raceme of 1-21 flowers,
the branches slightly puberulous. Flowers without noticeable odor, calyx tubular-
campanulate, 2-4-lipped, 1.1-2.1 cm long and 0.7-1.2 cm wide, inconspicuously
puberulous with simple, glandular, or rarely thick-stellate trichomes and/or
lepidote scales, without glands; corolla deep wine-colored; campanulate above
a narrowed 1.7-2.4-cm-long and 0.4-0.6-cm-wide base, 4.1-6.0 cm long and 0.8-
2.1 cm wide at the mouth, the tube 3.5-5 cm long, the lobes 0.6-1.5 cm long;
more or less glabrous to glandular-lepidote on the tube outside, simple-puberulous
at the base and margin of the lobes outside, stalked-lepidote in the floor of
the tube within, the lobes papillate-lepidote inside, pubescent at the level of
stamen insertion with short, 1-3-celled, gland-tipped trichomes; stamens didyna-
mous, the anther thecae divaricate, 2.5-3 mm long, the longer filaments 1.5-1.7
FIGURE 19. Martinella obovata (H.B.K.) Bur. & K. Schum. A. Habit (X 1/2). [After
Gentry 4981 (MO)] B. Fruit (X 1/2). [After Tyson 5460 (MO)] C. Ovary cross section
(X 10).-D. Pistil and disc ( x ?/2). E. Anther ( x 5). [After miscellaneous fresh material.]
cm long, the shorter filaments 1.1-1.4 cm long, the filaments inserted 14-18 mm
from the base of the corolla tube, the staminode 1-2 mm long, inserted 10-11
mm from base of tube; pistil 3.5-3.7 cm long, the ovary linear, 4 mm long and
1.5 mm wide, sparsely lepidote or simple-puberulous, the ovules (2-)4-seriate
in each locule; disc annular-pulvinate, 1.5 mm long and 4-S mm wide. Capsule
linear, acute, strongly flattened, 55-130 cm long and 1.2-1.8 cm wide, the
surface even, finely ridged under a lens, glabrous to inconspicuously subpuberu-
lous, especially near the margins; seeds very thin, 0.9-1.8 cm long and 3.0-5.4
cm wide, the wings symmetrical, blunt-ended, membranaceous but brownish,
lighter at the tips, poorly demarcated from the seed body.
A widespread but relatively uncommon vine of tropical wet forest, pre-

montane wet forest, and wetter parts of the tropical moist forest, M. obovata
ranges from British Honduras to Brazil and Bolivia. It flowers sporadically
through the year especially in the wet season.
When sterile this vine is recognized by its glossy, long-petioluled leaves with
prominent linear interpetiolar swellings, trifid tendrils, and the tendency of
the petioles and petiolules to bend or twist. The flower color is distinctive and
the fruits are perhaps the longest (to 190 cm, fide Dugand, Caldasia 7(31): 13.
1955) in the family.
CANAL ZONE: Barro Colorado Island, Croat 10079, 14064 (both MO), 14105 (MO, NY,
SCZ), 14658 (MO). Boy Scout Road near Madden Lake, Dwyer & Elias 7489 (MO, SCZ).
Cerro Gordo near Culebra, Pittier 3735 (F, US). Chilibre River below Chilibre, Seibert 1518
(MO, US). CHIIQUI: Near Rio Tinta, Woodson et al. 403 (MO). COLON: Santa Rita
Ridge, Gentry 1800 (MO). DARIEN: Rio Piriaque, Duke 8141 (MO). Rio Sabana above Santa
Fe, Duke 14111 (MO). El Real, Gentry 4534 (MO). Without locality, Barclay s.n. (K).
PANAMA: San Jose Island, Erlanson 97, 338 (both GH, NY, US). Rio Piragua (Tabardi),
Gentry 2528 (MO). Rio Sulugandi, Gentry 3744 (MO). 0.4 km from Rio Bayano crossing
on road to Santa Fe, Gentry 3853 (MO). Rio Espave near junction with Rio Bayano, Gentry
3898 (MO). Between Rio Sulugandi and Rio Espave, Gentry 4981 (MO). Near Madden
Lake, Gentry & Tyson 5037 (MO). San Jose Island, Johnston 771 (BM, GH, MO, US). Road
to Cerro Azul, Lazor 5548 (SCZ). Vicinity of Cerro Jefe, Lewis & Dressler 7566 (MO). Near
Rio Pacora, Maurice 815 (US). Macapale Island in Madden Lake, Tyson 5460 (MO, SCZ).
SAN BLAS: Near stream above Puerto Obaldlia, Gentry 1539 (MO).
20. MELLOA
Melloa Bur., Adansonia 8: 379. 1868. TYPE: Ml. populifolia (DC.) Britt. = M.
quadrivalvis (Jacq.) A. Gentry.
Lianas, stems with phloem irregularly many-armed in cross section;

branchlets terete, with or without interpetiolar glandular fields; pseudo-
stipules ovate to almost subulate-acuminate. Leaves 2-foliolate, usually with a
short, terminal, trifid tendril with recurved claw-like arms, the tendril becoming
thickened and woody with age. Inflorescence usually a large, open, axillary or
terminal panicle, sometimes reduced to a few-flowered cyme, its dichotomies
subtended by caducous foliaceous bracts. Flowers with the calyx membra-
naceous, broadly campanulate, obliquely truncate to subspathaceous with a small
more or less recurved apicule; corolla yellow, tubular-infundibuliform, glabrous
outside; anthers glabrous, the thecae straight, divaricate; pollen grains single,
3-colpate, the exine almost smooth; ovary flattened-ovoid, ribbed, glabrous, the
ovules multiseriate in each locule; disc apparently double. Fruit a slightly
compressed elliptic-oblong capsule, the valves parallel to the septum, woody,
very thick (ca. 8 mm), each splitting in the middle at maturity; seeds thin,
bialate, the wings hyaline-membranaceous with brown streaks, well demarcated
from the seed body.
One species of continental tropical America ranging from Mexico to Argentina.
1. Melloa quadrivalvis (Jacq.) A. Gentry, Brittonia 25: 237. 1973.-FIG. 20.
Bignonia quadrivalvis Jacq., Fragm. Bot. 37, tab. 40, fig. 3. 1800-1809. TYPE: Venezuela,
Caracas (not seen).
Spathodea fraxinifolia H.B.K., Nov. Gen. Sp. P1. 3: 147. 1819. LECTOTYPE: Venezuela, Rio
Guarico, Humboldt & Bonpland 780, flowers only (P).
Bignonia populifolia DC., Prodr. 9: 159. 1845. TYPE: Brazil, Cujaba, Manso s.n. (G-DC).
Phryganocydia pisoniana Mart. ex DC., Prodr. 9: 209. 1845, pro syn.
Spathodea pisoniana DC., Prodr. 9: 209. 1845. TYPE: Brazil, Bahia, Martius, 1818. (M).
Tabebuia pisoniana (DC.) Miers, Proc. Roy. Hort. Soc. 3: 199. 1863.
Tecoma pisoniana DC. ex Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 312. 1897, sphalma for
Spathodea?
Melloa populifolia (DC.) Britt., Ann. New York Acad. Sci. 7: 188. 1893.
Liana; twigs terete, gray or brown with pale lenticellate flecks, the nodes
without interpetiolar glandular fields but often with a ridge connecting opposite
petioles; pseudostipules small, deltoid, apiculate. Leaves 2-foliolate, sometimes
with a tendril or tendril scar; leaflets elliptic, acute to rounded, cuneate to
truncate at the base, 5-10 cm long and 2-5.5 cm wide, membranaceous, mostly
glabrous, scattered lepidote, inconspicuously scabrous at the extreme base of
the midvein above, secondary veins 3-6 on a side, drying olive; tendril trifid,
the 3 arms uncate, becoming woody with age; petiolules 0.6-2 cm, the petiole
1.1-3.5 cm long, inconspicuously puberulous, usually drying dark brown. Inflor-
escence an open, few-flowered panicle sometimes reduced to a single dichotomy,
each dichotomy subtended by caducous foliaceous bracts 7-21 mm long and
3-6 mm wide, the pedicels 1-3 cm long, drying dark as the inflorescence rachis.
Flowers with the calyx broadly campanulate, membranaceous, more or less
subspathaceous, with an apicule ca. 1 mm long, 10-21 cm long and 9-13 cm
wide, glabrous (puberulous in Spathodea pisoniana); corolla yellow, tubular,
4-7 cm long and 0.8-1.5 cm wide at the mouth, the tube 3.5-5 cm long, the
lobes 0.8-1.5 cm long, mostly glabrous, the lobes more or less ciliate with simple
trichomes, conspicuously so in bud, the tube pubescent inside with short tri-
chomes at the level of stamen insertion; stamens didynamous, the thecae divari-
cate, 4-5 mm long, the longer filaments 2.1-2.3 cm long, shorter filaments
1.4-1.6 cm long, the staminode 5-7 mm long, inserted 5-7 mm from the base
of the corolla tube; pistil 3.8-4.0 cm long, the ovary flattened-oblong, conspicu-
ously longitudinally ridged, glabrous, the ovules multiseriate in each locule;
disc on a short, thick stalk (hence sometimes described as "double"), 1.5-2 mm
long and 3.5-4 mm wide. Capsule elliptic, the valves woody, thick (5-8 mm)
at maturity breaking evenly in 2 at the midline, drying black or dark brown
with tan, lenticellate flecks, minutely tuberculate (sandpaper-finished), 11-15
A~~~
FIGuRE 20. Melloa quadrivalvis (Jacq.) A. Gentry.-A. Habit (X 1/2). [A

et al. 10319, Mexico (MO).]-B. Fruit (x 1/2). [After Irwin et al. 19137, Br
cm long and 3.9-5.2 cm wide; seeds thin, bialate, 1.1-1.6 cm lon

cm wide, the body brown, the wings hyaline-membranaceous with
streaks well demarcated from the seed body.
Though widespread, ranging from Mexico to Brazil and Argentina, this

liana is of only sporadic occurrence through most of its range and is poorly
represented in herbaria. It is one of the few species I have not seen in the
field, and its ecological parameters remain undertermined. Although not yet
collected in Panama, it almost certainly occurs there, having been collected
once in Nicaragua, twice in Costa Rica, and once or twice in northern Colombia.
The affinity of M. quadrivalvis is with Macfadyena, especially with M.
unguis-cati, from which it is sometimes difficult to distinguish when in flower.

In fact it has been suggested that if Doxantha is to be sunk into Macfadyena,
then perhaps Melloa should be also. However, the remarkably different fruit
of Melloa seems sufficient to maintain its generic separation.
21. MUSSATIA
Mussatia Bur. ex Baill., list. P1. 10: 32. 1888. TYPE: M. prieurei (DC.) Bur.
& K. Schum.
Lianas; branchlets acutely tetragonal with the angles ribbed, interpetiolar

glands wanting; pseudostipules large, foliaceous. Leaves 2-foliolate, sometimes
with a simple tendril. Inflorescence a terminal panicle. Flowers with the calyx
short, shallowly campanulate, truncate, more or less lobed or split; corolla yellow
with purplish to brownish stripes, infundibuliform, bilabiate, glandulaf-lepidote
outside; anthers glabrous, the thecae short, divergent to divaricate; pollen grains
single, 3-colpate, the exine reticulate; ovary oblong, lepidote, the ovules 4-6-
seriate in each locule; disc cupular-pulvinate. Fruit an oblong to linear-oblong
capsule, the valves parallel to the septum, woody and convex, rugose tuberculate
or slightly and inconspicuously short muricate; the seeds thin, bialate, the wings
whitish to brownish membranaceous, poorly demarcated.
Two species from southern Mexico to Amazonia.
Useful reference:
Sandwith, N. Y. Notes on tropical American Bignoniaceae. Recueil Trav.
Bot. Neerl. 34: 205-232. 1937.
1. Mussatia hyacinthina (Standl.) Sandw., Recueil Trav. Bot. Neerl. 34: 218.
1937.-FIG. 21.
Bignonia prieurei DC. sec. Seem., Bot. Voy. Herald 179. 1854, non DC.
Tynnanthus hyacinthinus Standl., Carnegie Inst. Washington Publ. 461: 87. 1935. TYPE:
B3ritish Honduras, Jacinto Creek, Schipp S-661 (F, holotype; BM, G, K, isotypes).
Liana to 18 cm in diameter, the bark on smaller stems smooth or somewhat

longitudinally striate, light gray to reddish, papery-fibrous in texture, older
stems twisted bush ropes with numerous spiral-longitudinal furrows; stem
with 8(-16) phloem arms in cross section; branchlets strongly tetragonal with
raised ribs on the angles, subscabrous or lepidote, the nodes without glandular
fields; pseudostipules foliaceous, more or less ovate, usually acute, 4 at each
node, 1.2-2.1 cm long and 0.5-1.7 cm wide. Leaves 2-foliolate, sometimes cir-
rhose; leaflets ovate, acute to acuminate, basally truncate to broadly cuneate,
12.4-23 cm long and 6.2-11.5 cm wide, entire, membranaceous, secondary veins
6-7 on each side, scattered lepidote above and beneath, somewhat puberulous
on the depressed midvein above, more or less scabrous on main veins beneath,
conspicuously pubescent with long simple trichomes in axils of secondary veins
beneath, drying greenish-olive with veins brownish; tendril simple, 12-23 cm
long; petiolules 2.4-4.5 cm long, the petiole 5.9-8.1 cm long, lepidote. Inflores-
cence a terminal panicle, its branches lepidote. Flowers with the calyx broadly
FIGURE 21. Mussatia hyacinthina (Standl.) Sandw. A. Inflorescence (X '/2) [After

Tun Ortiz 992, Guatemala (MO).]-B. Leaves (X 1/2). [After Gentry 2736 and 4527 (both
MO).]-C. Fruit (X '/2). [After Gentry 2736 (MO).]
campanulate, short, 1-2 mm long and 2-4 mm wide, slightly lobed, glabrous,
the margin thin; corolla yellow, finely mottled with purple outside, streaked
with red or brown inside, infundibuliform, 15-19 mm long and 5-7 mm wide
at the mouth, the tube 10-12 mm long, the lobes 5-8 mm long, narrow; the
tube conspicuously glandular-lepidote outside, lobes glandular-lepidote, pubes-

cent at the tips and with a ciliate fringe, the tube lepidote within, lepidote but
otherwise glabrous at the level of stamen insertion; stamens didynamous, the
anther thecae slightly curved, somewhat divergent to divaricate, 1.0-1.5 mm
long, the longer filaments 1.0-1.4 cm long, the shorter filaments 0.8-1.0 cm long,
the staminode 5-6 mm long, inserted 1-2 mm from base of corolla tube; pistil
1.4-1.5 cm long, the style bent to follow the posterior curve of corolla, the
ovary oblong, densely lepidote, 3 mm long and 1.0-1.5 mm wide, 1.0 mm thick,
the ovules 4(-6)-seriate in each locule; disc shallowly pulvinate-cupular, 0.5 mm
long and 1.5 mm wide. Capsule linear-oblong, obtuse at both ends, the valves
woody, flattened but slightly convex, 17.5-29 cm long and 4.4-5.5 cm wide,
the surface rough, irregularly warty-tuberculate; seeds 2.2-2.7 cm long and
5.4-6.1 cm wide, the wings wholly light brown, thinly woody at the base to
membranaceous at tips, not distinct from the seed body.
A liana of the tropical wet forest and premontane wet forest, it is locally
common and reaches gigantic size. It ranges from British Honduras to British
Guiana and Peru. The flowers are rarely collected and little known.
The conspicuously square stem with foliaceous pseudostipules is an excellent
character for the identification of Mussatia. Pubescent leaf axils beneath and
relatively thick branchlets distinguish it from Cydista diversifolia (H.B.K.)
Miers, which has similar stems and pseudostipules. The fruit is similar only
to that of Callichlamys, which has a smooth capsule valve rather than the slightly
tuberculate valve of Mussatia.
BOCAS DEL TORO: Hillside above Almirante, Gentry 2736, 2754 (both MO). COLON:
W of Portobelo, Gentry 1770 (MO). DARIEN: Slopes of Cerro Chucula (drainage of Rio
Pavarando), Gentry 4254 (MO). Between Rios Cube and Punusa, Gentry 4470 (MO). El
Real, Gentry 4527 (MO). Near Rio Parasenico, Gentry 4740 (MO). PANAMA: Near Rio
Espave', Gentry 3702 (MO). 5-10 km from Rio Bayano crossing on trail to Santa Fe, Gentry
3844 (MO). Near Rio Bayano dam site, Gentry & Tyson 1661 (MO). VERAGUAS: Remedios,
Seemann 1128 (K, P).
22. ON'OHUALCOA
Onohualcoa Lundell, Contr. Univ. Michigan Herb. 7: 52. 1942. TYPE: 0. seleri
(Loes.) Lundell= 0. verrucifera (Schlecht.) A. Gentry.
Bayonia Dugand, Caldasia 4: 62. 1946. TYPE: B. helicocalyx (0. Kuntze) Dugand -
Onohualcoa verrucif era (Schlecht.) A. Gentry.
Lianas, stems with 8 phloem arms in cross section; branchlets terete, glabrate,
conspicuously lenticellate, without interpetiolar glandular fields; pseudostipules
inconspicuous, when present short and blunt-conical. Leaves 3-foliolate, the
terminal leaflet often replaced by a trifid tendril. Inflorescence a 3-6-flowered
corymb in the axil of a young leaf or leaf primordium, these aggregated into
many-flowered narrow terminal panicles. Flowers with the calyx cupular to
tubular-campanulate, 5-ribbed with the ribs ending in elongate (to 5 mm)
subulate teeth, puberulous and lepidote; corolla lavender, tubular-campanulate,
simple puberulous and glandular-lepidote outside; anthers glabrous, the thecae
straight, divaricate; ovary cylindric, more or less glandular-papillate and incon-
spicuously scabrous, the ovules 2-seriate in each locule. Frui

pressed linear-oblong capsule, the valves parallel to the sept
tuberculate with tubercles to 5 mm long, seeds thick-bodied
branaceous wings basally brown and hyaline at the tips.
One species ranging from southern Mexico to British Guiana and possibly
also to Peru and Brazil (fide Sandwith, 1954).
1. Onohualcoa verrucifera (Schlecht.) A. Gentry, comb. nov.-FIG. 22.

Bignonia verrucifera Schlecht., Linnaea 26: 655. 1853. TYPE: Venezuela, Curucati, Wagener
307 (not seen).
Adenocalymma verruciferum (Schlecht.) Miers, Ann. Mag. Nat. Hist., ser. 3. 7: 393. 1861.
A. helicocalyx 0. Kuntze, Rev. Gen. 2: 478. 1891. SYNTYPES: Trinidad, Kuntze 1153 (K,
NY); Venezuela, Kuntze 1734 (US).
A. moritzii Urb., Fedde Repert. 14: 308. 1916. TYPE: Venezuela, prope Palmar, Moritz s.n.
(not seen).
Setilobus vicentinus Kranzl., Fedde Repert. 17: 122. 1921. TYPE: St. Vincent, Caley s.n. (
Adenocalymma fissum Loes., Verh. Bot. Vereins Prov. Brandenberg 65: 102. 1923. TYPE:
Mexico, Yucatan, Seler 4034 (not seen).
A. seleri Loes., Verh. Bot. Vereins Prov. Brandenberg 65: 101. 1923. TYPE: Mexico, Yucatan,
Seler 9901 (not seen).
A. micradenium Standl., Trop. Woods 45: 16. 1936. TYPE: Colombia, La Mojama, Dugand
611 (F).
Onohualcoa seleri (Loes.) Lundell, Contr. Univ. Michigan Herb. 7: 52. 1942.
Bayonia fissa (Loes.) Dugand, Caldasia 4: 65. 1946.
B. helicocalyx (0. Kuntze) Dugand, Caldasia 4: 63. 1946.
Onohualcoa fissa (Loes.) Sandw., Kew Bull. 1946: 88. 1947.
0. helicocalyx (0. Kuntze) Sandw., Kew Bull. 1946: 87. 1947.
Adenocalymma perezii Standl. & L. 0. Wms., Ceiba 3: 61. 1952. TYPE: Honduras, Cortes,
Perex s.n. (US).
Liana to at least 8 cm in diameter, the bark smooth, gray with vertical

of raised brownish lenticels; wood somewhat rank-smelling; stem with 8
arms in cross section; twigs subterete, glabrous to subpuberulous, when dry
longitudinally ridged and dark with pale lenticels, the nodes without inter-
petiolar glandular fields; pseudostipules inconspicuous, blunt-conical. Leaves
3-foliolate or 2-foliolate with tendril or tendril scar; leaflets ovate-elliptic, acute,
basally more or less cuneate to broadly subcordate, 7.1-13.5 cm long and 3.8-6.3
cm wide, membranaceous, 3-nerved from the base, the secondary veins 4-8 on
a side, glabrous to puberulous above and beneath, usually puberulous at least
at the base of the midvein with glandular fields at the base and apex of the
midvein beneath, also with a few scattered plate-shaped glands, more or less
olive when dry (young leaves drying black); tendril trifid, 8-14 cm long to
branching, the arms 1-1.4 cm long; terminal petiolule 2.5-4.2 cm long, the
lateral petiolules 1.5-2.2 cm long, the petiole 3.7-8.7 cm long, puberulous.
Inflorescence a 2-6-flowered corymb in the axil of a young leaf or leaf primor-
dium, these aggregated into many-flowered narrow terminal panicles, the
branches puberulous. Flowers with sweet aroma, calyx cupular, the 5 main ribs
ending in narrow, elongate (to 5 mm) teeth, puberulous and lepidote, glandular,
5-7 mm long (without teeth) and 5-6 mm wide, the surface minutely roughened,
subverrucose; corolla pinkish lavender with a white throat; tubular-campanulate
above an 8 mm by 3 mm tubular base, 5.4-8.1 cm long and 1.0-1.9 cm wide at
FIGuRE 22. Onohualcoa verrucifera (Schlecht.) A. Gentry.-A. Habit (X 1/2). B. Fruit

(X 3/1o). [After Pittier 11100, Venezuela (VEN).]-C. Anther (X 4).-D. Calyx and pistil
( X /2).-E. Ovary cross section ( X 121/2).-F. Ovary and disc (x 3). [After Gentry &
Tyson 5025 (MO).]-G. Inflorescence (x '/2). [After Pittier 14288, Venezuela (VEN).]
the mouth, the tube 3.5-6.0 cm long, the lobes 1.2-2.5 cm long; simple puberulous
and glandular-lepidote outside, less so on the lobes, mostly glabrous within
but somewhat glandular-lepidote on lobes and pubescent at the level of stamen
insertion with long, simple, in part gland-tipped, multicellular trichomes; stamens
didynamous, the anther thecae divaricate, 3.5-5 mm long, the longer filaments
2.3-2.4 cm long, the shorter filaments 1.1-1.6 cm long, the staminode 4-5 mm
long, inserted 8-10 mm from the base of the corolla tube; pistil 3.5-4.2 cm long,
the ovary cylindric, 5 mm long and 1-1.5 mm wide, more or less glandular-
papillate and inconspicuously scabrous, the ovules 2-seriate in each locule; disc
pulvinate, 1 mm long and 3-4 mm wide. Capsule linear-oblong, 17-30 cm long
and 2.3-2.6(-3.8) cm wide, the surface strongly tuberculate; seeds 1.5-2.5 cm
long and 4-7.8 cm wide, bialate, the wings hyaline-membranaceous at the ends,
basally brown and coriaceous, merging with the thick woody seed body.
An uncommon but widespread plant of the tropical wet forest and wetter
parts of the tropical moist forest, this species ranges from southern Mexico to
Venezuela and British Guiana.
The Mexican population has heretofore been regarded as geographically
separated and specifically distinct from the plants of northern South America.
However, recent collections of this species from Honduras, Costa Rica, and
Panama remove the range disjunction and are more or less intermediate morpho-
logically as well. The variation in tubercle length is clinal, and the plants are
best treated as a single species.
I have followed a suggestion of Sandwith (1954) in adopting the basionym
Bignonia verrucifera Schlecht., which is older than 0. helicocalyx (0. Kuntze)
Sandw., which has been in general use for this plant. I have not been able
to locate type material of Bignonia verrucifera, but the description agrees with
this species and does not fit any of the other species known from the type
locality in northern Colombia.
CANAL ZONE: Pipeline Road, TTC transect 1, Gentry 7391 (MO). COLON: Above
Portobelo Road, Gentry 1763 (MO). PANAMA: Near NMadden Lake, Gentry 5050 (MO);
Gentry & Tyson 5025 (MO). SAN BLAS: Mainland opposite Playon Chico, Gentry 6372
(MO).
23. PACHYPTERA
Pachyptera DC. ex Meisn., Gen. 1: 299; 2: 207. 1840. LECTOTYPE: P. foveolata

DC. = P. kerere (Aubl.) Sandw.
Pseudocalymma Samp. & Kuhlm., 0 Campo (Rio de Janeiro) 4(11): 15. 1933; Bol. Mus.
Nac. Rio de Janeiro 10: 101. 1934. TYPE: P. laevigatum (Bur. & K. Schum.) Samp.
& Kuhlm. = Pachyptera hymenaea (DC.) A. Gentry.
Lianas; stems with 4 or 8 phloem arms in cross section; branchlets terete to

subtetragonal, outer cortex thin and tending to peel; often with conspicuous
interpetiolar glandular fields; pseudostipules subulate, usually vertically 2-3-
seriate. Leaves 2-3-foliolate, the petiole usually with an apical glandular field,
the terminal leaflet often replaced by a trifid tendril or a simple tendril with
a peltate disc at its tip. Inflorescence an axillary or terminal panicle or raceme.
Flowers with the calyx cupular to campanulate, truncate, often with plate-shaped
glands in the upper part; corolla white to purple, tubular to campanulate, pub-
erulous outside at least on the lobes; anthers glabrous or villous, the thecae
divaricate, curved or straight; pollen grains 3-colpate, the exine finely reticulate;
ovary cylindric, pustular-lepidote to papillate, the ovules 2(?-4)-seriate. Fruit
a pale brown capsule, the valves parallel to the septum, extremely flat and
compressed to thick and convex with a conspicuously raised midline; the seeds
thin with pale translucent papery wings or thick, corky, and essentially without
wings.
About seven species in continental tropical America ranging from Mexico to

Brazil.
a. Tendrils simple, disc-tipped; leaflets less than 3 cm long -3. P. parvifolia

aa. Tendrils trifid, not disc-tipped; leaflets more than 4 cm long.
b. Vegetative parts without onion odor; pseudostipules acute, vertically 3-seriate;
leaves mostly 3-foliolate; inflorescence a short-pedicelled raceme; corolla puberulous
outside on tube and lobes; anther thecae villous.
c. Capsule valves thick, woody, convex, the median nerve conspicuously raised;
seeds thick, corky, essentially wingless -2a. P. kerere var. kerere
cc. Capsule valves thin and perfectly flat or slightly raised along the margin, median
nerve not obvious; the seeds thin, bialate, with broad papery wings (not yet seen
from Panama) -2b. P. kerere var. incarnata
bb. Vegetative parts with conspicuous onion or garlic odor; pseudostipules short, blunt,
relatively inconspicuous; leaves 2-foliolate; inflorescence a panicle or few-flowered
open raceme; corolla puberulous outside only on lobes; anther thecae glabrous.
d. Calyx large, more than 13 mm long and 13 mm wide, red violet when fresh; in-
florescence an elongate, open, few-flowered panicle or raceme; one pair of anthers
exserted; fruits strongly compressed, the median nerve obscure; leaflets elliptic,
with glandular fields in the axils of basal lateral nerves beneath - 4. P. standleyi
dd. Calyx small, less than 7 mm long and 7 mm wide, greenish when fresh; inflores-
cence usually a more or less contracted panicle; anthers included; fruits only
somewhat compressed, the median nerve conspicuous, raised; leaflets ovate to
elliptic, with or without concentration of glands in axils of basal lateral nerves
beneath -1. P. hymenaea
1. Pachyptera hymenaea (DC.) A. Gentry, Brittonia 25: 236. 1973.
Bignonia hymenaea DC., Prodr. 9: 158. 1845. TYPE: Brazil, Bahia, Blanchet 1434 (G-DC,
P).
Anemopaegma pachypus K. Schum. in Engler & Prantl, Nat. Pflanzenf. 4( 3b): 215. 1894,
sphalma platypus. TYPE: Peru, San Martin, Spruce 4475 (K).
Bignonia laevigata K1. ex Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 113. 1896, pro syn.
Adenocalymma laevigatum Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 113. 1896, non Mart.
ex DC. SYNTYPES: Brazil, Rio de Janeiro, Sellow L.317 (not seen); Brazil, Langsdorf
s.n. (not seen), Glaziou 9529 (F, K, MO).
A. pachypus (K. Schum.) Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 110. 1896.
A. pohlianum Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 114. 1896. TYPE: Southeast Brazil,
Pohl 1817 (K).
A. macrocarpum Donn. Sm., Bot. Gaz. (Crawfordsville) 40: 9. 1915. LECTOTYPE: El Salva-
dor, San Vicente, Shannon 5055 (US).
A. obovatum Urb., Fedde Repert. 14: 307. 1916. TYPE: Haiti, Mackenzie, Duss 1913 (not
seen).
A. ciliolatum Blake, Contr. Gray Herb. 52: 90. 1917. TYPE: Nicaragua, Leon, Baker 2424
(GH, holotype; MO, NY, US, isotypes).
A. hosmeca Pittier, Contr. U. S. Nat. Herb. 18: 256. 1917. TYPE: Costa Rica, Guanacaste,
0. Jimenez 374 (US).
Petastoma langlasseanum Kranzl., Fedde Repert. 17: 56. 1921. TYPE: Mexico, Nusco,
Langlasse 738 (K, P, US).
P. tonduzianum Kranzl., Fedde Repert. 17: 56. 1921. TYPE: Costa Rica, Guanacaste, Tonduz
13830 (BM, CR, K, P, US).
Adenocalymma alboviolaceum Loes., Verh. Bot. Vereins Prov. Brandenburg 65: 100. 1923.
TYPE: Mexico, Oaxaca, Seler 1664 (GH, US).
Pseudocalymma laevigatum (Bur. & K. Schum.) Samp. & Kuhl., Bol. Mus. Nac. Rio de
Janeiro 10: 101, f. 1934.
P. hymenaeum (DC.) Sandw., Candollea 7: 247. 1937.
P. langlasseanum (Kranzl.) Sandw., Recueil Trav. Bot. Neerl. 34: 210. 1937.
P. macrocarpum (Donn. Sm.) Sandw., Recueil Trav. Bot. Neerl. 34: 210. 1937.
P. pachypus (K. Schum.) Sandw., Recueil Trav. Bot. Neerl. 34: 210. 1937.
P. pohlianum (Bur. & K. Schum.) Sandw., Recueil Trav. Bot. Neerl. 34: 211. 1937.
P. alliaceum var. microcalyx Sandw., Kew. Bull. 1953: 467. 1954, pro parte, typo excluso.
Liana to at least 5 cm in diameter, bark smooth, gray; stem with 4-8 phloem
arms in cross section; all parts with strong garlic odor; branchlets subterete,
glabrous or sparsely lepidote, the nodes with interpetiolar glandular fields;
pseudostipules small, flattened-conical. Leaves 2-foliolate, usually with a tendril
or tendril scar; leaflets ovate to widely ovate, obtuse to acute, basally (Panama)
more or less truncate to shallowly cordate, 4.8-9.4 cm long and 3.3-7.5 cm wide,
membranaceous, more or less 3-veined from the base, the secondary veins 3-6
on a side, mostly glabrous above and beneath, slightly lepidote at least at the
base, often with a glandular field in the axil of the basal nerve pair beneath,
drying olive; tendril shortly trifid, 10-14 cm to branching, the 3 arms 0.1-0.4 cm
long; petiolules 1.0-2.6 cm long, petiole 1.3-3.5 cm long, petiole and petiolules
lepidote, sometimes puberulous at their junction, the petiole sometimes with an
apical glandular field. Inflorescence a usually more or less contracted panicle
terminal on a young shoot, the branches puberulous. Flowers with the calyx
cupular, truncate or minutely 5-denticulate with the extreme edge membra-
naceous, 4-6 mm long and 4-6 mm wide, slightly lepidote to puberulous, drying
with glandular light spots; corolla lavender to magenta or almost white, tubular-
campanulate, 4.1-5 cm long and 0.8-1.5 cm wide at the mouth, the tube
3.0-3.6 cm long, the lobes 0.6-1.3 cm long, the lobes puberulous, tube mostly
glabrous inside and out, inconspicuously pubescent at the level of stamen inser-
tion; stamens didynamous, included, the anther thecae somewhat divergent,
3 mm long, the longer filaments 1.8-1.9 cm long, shorter filaments 1.2-1.4 cm
long, the staminode ca. 2 mm long, inserted 5-7 mm from base of corolla tube;
pistil 2.4-2.6 cm long, the ovary linear-tetragonal, 5-7 mm long and 1 mm wide,
lepidote, the ovules 2-seriate in each locule; disc annular-pulvinate, 1 mm long
and 2 mm wide. Capsule linear, long-acuminate, flattened, 15-25 cm long and
1.7-2.0 cm wide, the surface smooth, glabrous to inconspicuously lepidote with
a conspicuously raised central rib, drying tan with scattered small blackish dots;
seeds thin, bialate, 1.1-1.5 cm long and 3.1-3.5 cm wide, the wings membra-
naceous, pale brown at the base, whitish to subhyaline at the tip.
A common liana of the tropical dry forest occurring also sporadically in the
tropical moist forest, it ranges from Mexico to Brazil. This species flowers in
the dry season from February through March. The seeds are dispersed at the
end of the dry season.
The characteristic pungent odor has led to its inclusion in the native pharma-
copeia. The Cuna Indians of Panama (Duke 10854) put it on their heads to
prevent bad dreams. It is one of the few Bignoniaceae vines to receive a com-
mon name other than the general term "bejuco" and is known variously as
olmeca, "hosmeca," "Jumeca, ago, or pedo de padre."
In the field this plant is unmistakable because of its strong onion or garlic
odor, and even in the herbarium some of this odor is often maintained. The
leaves are also recognizable by their dark green color with pale main veins and
their usual wide-ovate shape with a truncate or subcordate base. The fruits can
be identified by the combination of long, tapering apex, tan color, and strongly
raised midline.
PANAMA: Rio Agua Clara at Icanti, Gentry 2596 (MO). San Jose Island, Johnston 1342
(GH, MO, US). SAN BLAS: Rio Ailigandi, Duke 10854 (MO). VERAGUAS: Slopes of Cerro
Tute W of Santa Fe, Allen 4442 (GH, MO, NY). WITHOUT LOCALITY: Hayes 610 (K).
2. Pachyptera kerere (Aubl.) Sandw., Recueil Trav. Bot. Neerl. 34: 219. 1937.
-FIG. 23A-E.
Bignonia kerere Aubl., Hist. PI. Guiane Fr. 2: 644, tab. 260. 1775, non Lindl. TYPE: French
Guiana, Aublet s.n. (BM, P-AD no. 12271, W).
B. incarnata Aubl., Hist. PI. Guiane Fr. 2: 645, tab. 261, 262, fig. 1-8. 1775. TYPE: (not
seen).
B. heterophylla Willd., Sp. PI. 3: 298. 1801. TYPE: Based on B. kerere Aubl.
Adenocalymma brachybotrys DC., Prodr. 9: 202. 1845. TYPE: French Guiana, Perrotet s.n.
(P).
Pachyptera foveolata DC., Prodr. 9: 175. 1845. TYPE: French Guiana, Poiteau s.n. (G-DC,
fragment).
Adenocalymma stridula Miers, Ann. Mag. Nat. Hist., ser. 3. 7: 392. 1864. TYPE: Based on
Bignonia kerere Aubl.
A. foveolatum (DC.) Baill., Hist. PI. 10: 7, fig. 9-16. 1888.
A. kerere (Aubl.) Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 119. 1896, pro parte.
Bignonia benensis Britton ex Rusby, Bull. Torrey Bot. Club 27: 70. 1900. TYPE: Bolivia,
junction of Beni and Madre de Dios Rivers, Rusby 1143 (NY, US).
Adenocalymma symmetricum Rusby, Descr. 300 Sp. S. Amer. PI. 122. 1920. TYPE: Venezuela,
Lower Orinoco, Rusby & Squires s.n. (NY).
Tanaecium zetekii Standl., Contr. Arnold Arbor. 5: 140, tab. 19. 1933. TYPE: Panama, Barro
Colorado Island, Woodworth & Vestal 363 (F, holotype; A, MO, US, isotypes).
Pachyptera kerere var. erythraea Dugand, Caldasia 7(31): 16. 1955. TYPE: Colombia,
Santander, Romero-Castaiieda 4727 (COL).
Liana to at least 5 cm in diameter, the bark gray, smooth; stem with 4

phloem arms in cross section; twigs subtetragonal, longitudinally striate, glab-
rous to subpuberulous, interpetiolar glands conspicuous; pseudostipules ver-
tically 3-seriate, acute. Leaves 3-foliolate or 2-foliolate with a tendril or tendril
scar; leaflets ovate, long-acuminate to acute, broadly cuneate to narrowly sub-
cordate, membranaceous, secondary nerves 6-10 on a side, scattered lepidote
below, less so above, puberulous along main veins above and beneath, sunken
plate-shaped glands usually conspicuous on both surfaces, olive to brownish
when dry, usually darker above; tendril trifid, 8-16 cm long to branching,
the three arms 0.2-4.0 cm long, often unequal; terminal leaflet 9-24 cm long
and 3.4-10 cm wide, the laterals 7.8-22 cm long and 2.4-9.2 cm wide; terminal
petiolule 1.5-6.1 cm long, the laterals 0.5-3.2 cm long, the petiole 0.9-7.1 cm
FIGuRE 23. Pachyptera.-A-E. P. kerere (Aubl. emend Splitg.) Sandw.-A. Inflores-

cence ( X 1/2).-B. Fruit ( X 1/2). [After Gentry 4401 (MO).]-C. Leaves (>X 1/2). [After
Gentry 4335 (MO).]-D. Pistil and disc (X 1/2).-E. Anther (x 5).-F-I. P. standleyi
Steyerm.-F. Corolla (X 1/2).-G. Calyx ( X 1/2).-H. Pistil and disc ( X 1/2).-I. Ovary
cross section ( X 121/2). [After Gentry & Tyson 5027 (MO).]
long, above with a conspicuous apical glandular field, scabrous as the petiolules.
Inflorescence a short axillary or terminal raceme of ca. 10 flowers opening one
at a time, the rachis and pedicels puberulous. Flowers with the calyx campanu-
late, irregularly truncate, puberulous, with scattered glands, 7-12 mm long and
5-9 mm wide; corolla white (Panama), tubular-campanulate, 3.6-7.3 cm long
and 0.7-1.4 cm wide at the mouth of tube, the tube 3.0-6.3 cm long with a
narrowed base 2.3-2.6 cm long and 0.2-0.6 cm wide, lobes 0.5-1.8 cm long,
simple puberulous outside with plate-shaped glands at the base of the lobes,
inside glandular-lepidote on the lobes and glandular-pubescent below the level
of stamen insertion; stamens didynamous, the anther thecae pubescent, each
2-4 mm long, usually bent, the longer filaments 1.9-2.1 cm long, shorter
filaments 1.4-1.6 cm long, the staminode 0.3-9.8 cm long, inserted 2.3-2.5 cm
above the base of corolla tube; pistil 4.5-4.8 cm long, ovary flattened-cylindric,
2-3 mm long and 1.5-2 mm wide, 1-1.5 mm thick, minutely papillose; disc
pulvinate, 1-2 mm long and 3-4 mm wide. Capsule (Panama) fusiform-oblong,
acuminate at both ends, only slightly compressed, 10.2-22 cm long and 2.4-4.0
cm wide, 1.0-2.4 cm thick, the surface smooth, somewhat glandular and slightly
lepidote, the midrib raised; seeds 1.8-2.4 cm long and 2.8-3.8 cm wide, rela-
tively thick and corky, without wings.
A vine restricted almost entirely to lake shores, stream margins, and coastal
swamps, P. kerere is often common, though seldom collected, in this specialized
habitat in the tropical wet forest and wetter parts of the tropical moist forest.
It ranges from British Honduras to Amazonian Brazil. This species flowers
sporadically throughout the year. The corky, wingless seeds of typical P.
kerere are water-dispersed.
Striking diagnostic characters include the 3-seriate pseudostipules and the
glandular field on the upper surface of the petiole apex. The pubescent anthers
are a good floral character.
Two color forms of this species are known in South America, but only
the white-flowered form has been recorded ihi Central America. The red-
flowered form has been designated as P. kerere var. erythraea Dugand. A wind-
dispersed form, P. kerere var. incarnata (Aubl.) A. Gentry, has been collected
in Honduras and the Choco of Colombia and will probably be found in Pan-
ama. It differs from the typical variety only in its longer, much more com-
pressed fruit and thin, winged seeds.
BOCAS DEL TORO: Hillside above Almirante, Gentry 2687 (MO). Vicinity of Chiriqui
Lagoon, von Wedel 1057 (GH, MO), 1314, pro parte (GH, MO, US), 1340, 1553, 1754 (all
GH, MO). Water Valley, von Wedel 806 (MO), 1660, pro parte (MO). CANAL ZONE:
Barro Colorado Island, Aviles 45 (F, MO); Bailey & Bailey 655 (F); Brown 12 (F); Croat
4664 (MO), 4753 (MO, SCZ), 5226 (SCZ), 5406, pro parte (MO), 6071 (MO, SCZ),
6088, 6227, 6843, 7085, 7126, 7906, 8209, 8434, 8704, 11085, 12846, 15074 (all MO); Foster
1308 (PMA); Gentry 442 (MO, WIS), 1734 (MO); Kenoyer 527 (US); Shattuck 270
(A, F, MO), 512 (F, MO, US); Wetmore & Abbe 28 (A, F), 28A (A, MO); Woodworth
& Vestal 363 (A, F, MO, US). Chagres, Fendler 206 (K, MO, US). COLON: W of
Portobelo, Gentry 1766 (MO). Rio Indio de Fato, Pittier 4258 (US). DARIEN: Rio
Ucurganti, Bristan 1157, pro parte (MO), 1169 (MO). Rio Pirre, Croat & Porter 15497
(MO). Rio Balsas between Quebrado Chusomocater and Rio Areti, Duke 8711 (MO). Rio
Jaque to base of Cerro Pavarando, Gentry 4199 (MO). Near mouth of Rio Paya on Rio
Tuira, Gentry 4335 (MO). Rio Tuira between Rios Paya and Cube, Gentry 4351 (MO).
Rio Tuira just above Rio Cube, Gentry 4369 (MO). Rio Paya from Rio Tuira to village of
Paya, Gentry 4381 (MO). Rio Tuira between Rios Paya and Aspave, Gentry 4401 (MO).
Between Rio Paya and Rio Paca, Gentry 4484 (MO). El Real, Gentry 4577 (MO). Near
Cerro Pirre on Rio Parasenico, Gentry 4688 (MO). PANAMA: Rio Sulugandi from mouth to
trail crossing, Gentry 3743 (MO). SAN BLAS: Mainland opposite Playon Chico, Gentry 6356
(MO). WITHOUT LOCALITY: Bristan 196 (MO).
3. Pachyptera parvifolia A. Gentry, Phytologia 26: 448. 1973. TYPE: Colombia,

Sur de Santander, Haught 2179 (MO).
Vine, the branchlets subterete, the smallest striate and more or less angulate,
inconspicuously capitate-puberulent or lepidote, glabrescent, the nodes without
interpetiolar glandular fields; pseudostipules minute, less than 1 mm long,
rhombic, acuminate, the margin much thinner when young. Leaves 2-foliolate,
often with tendril; leaflets elliptic, rounded or emarginate to obtuse, small,
to 3 cm long and 1.5 cm wide, membranaceous, basally asymmetrically rounded
and more or less 3-nerved, the secondary veins obscure especially when young,
conspicuously glandular-lepidote or lepidote-punctate, especially beneath, the
glands larger and more crowded at the base; tendril simple, terminating in a
woody, peltate disc, to 8 mm in diameter; petiolules 1-8 mm long, petiole
5-8 mm long, petiolules and petiole puberulous above. Inflorescences axillary
to fallen leaves, short, ebracteate, 2-3-flowered, the peduncle 5-6 mm long,
the pedicels 1.2-1.5 mm long, inconspicuously lepidote. Flowers with the
calyx campanulate, truncate or shallowly bilabiate, the margin sometimes
turned under, 5-6 mm long and 5-9 mm wide, inconspicuously lepidote or
with a few scattered simple trichomes; corolla light purple or white, tubular-
campanulate, 3.5-5 cm long and 0.8-1.2 cm wide at mouth, the base narrowed,
the tube 2.5-4 cm long, the lobes 1-1.3 cm long, scurfy puberulous outside,
mostly glabrous within, pubescent at the level of stamen insertion; stamens
didynamous, the longer filaments 1.6-2.5 cm long, the shorter filaments 1.1-1.8
cm long, the anther thecae divaricate, 2-4 mm long, glabrous, the connective
extended 0.5 mm, the staminode ca. 2 mm long," inserted 5-10 mm from the
base of the corolla tube; pistil 2.8-2.9 cm long, the ovary oblong, subtetragonal,
3-4 mm long and 1 mm wide at the base, slightly wider at the top, densely
and minutely lepidote, the ovules 4-seriate in each locule but appearing 2-seriate
in most cross sections; disc almost spherical, 2 mm long and 3 mm wide.
Capsule linear-oblong, woody, ca. 25 cm long and 2.5 cm wide, tan; seeds
thin, the wings brownish, not sharply differentiated from the body.
The distinctive juvenile stage of this species has been collected a number
of times from Honduras to Bolivia, but I have seen only two flowering collec-
tions, the type from Colombia and a specimen from Amazonian Brazil. It is
apparently restricted to the tropical wet forest.
The juvenile stage is a tenuous vine with extremely small bifoliolate leaves,
each bearing a short tendril terminated by a peltate disc. These plants grow
closely appressed against the trunks of supporting trees, attached by the
tendril-discs.
A note on the fruit of this species is in order. I collected fallen capsule
valves and seeds at two localities on the Rio Brazo Viejo, Choco, Colombia,
in an area where juvenile vines were common. The specimens were subse-
quently destroyed by the Summit Herbarium fire (June 1972) before measure-
ment or critical examination. However, some previously planted seeds germi-
nated, giving rise to the typical juvenile form and establishing the identity
of the fruiting material with this species. My general impression of the fruit
was that it was similar to that of Ceratophytum in size, color, and form,
although much more compressed. Destruction of the collection prevents a
more complete description.
Although inclusion of P. parvifolia in Pachyptera significantly enlarges the
concept of the genus, it seems better than the alternative of erecting a new
monotypic genus. This emphasizes the apparent affinities of the species
rather than its peculiarities.
BOCAS DEL TORO: Water Valley, von Wedel 687 (MO). Top of fila above Almirante,
Gentry 2769 (MO). DARIEN: Between upper Rio Membrillo and Camp 7 on the construction
road to San Blas, 100-800 m, Duke 10875 (MO).
4. Pachyptera standleyi (Steyermark) A. Gentry, Brittonia 25: 236. 1973.-

FIG. 23F-J.
Pseudocalymma standleyi Steyermark, PubA. Field Mus. Nat. Hist., Bot. Ser. 23: 235. 1947.
TYPE: Guatemala, Quetzaltenango, Steyermark 33533 (F).
P. alliaceum var. macrocalyx Sandw., Kew Bull. 1953: 468. 1954. TYPE: British Guiana,
Demerara River, Jenman 6752 (K, holotype; NY, isotype).
Liana to at least 5 cm in diameter, bark smooth, pale gray, the outer bark
thin; stem with 8 phloem arms in cross section; all parts with strong garlic
odor; branchlets subterete to subtetragonal, glabrous to scabridulous or incon-
spicuously lepidote, the nodes with interpetiolar glandular fields, the pseudo-
stipules small, blunt-conical, with distinct upper and lower segments. Leaves
2-foliolate, sometimes with a tendril or tendril scar; leaflets elliptic, acute to
short-acuminate, basally widely cuneate to rounded, 8.6-20 cm long and 3.7-22
crm wide, membranaceous, the secondary veins 7-10 on a side, glabrous or
slightly lepidote especially at the base, beneath with glandular fields in axils
of basal nerve pair; drying green to olive-green above and beneath; tendril
trifid, 9-12 cm long to branching, the 3 arms 0.4-2.0 cm long; petiolules 0.7-2.0
cm long, petiole 0.9-3.2 cm long, glabrous or slightly lepidote as the petiolules,
often with an apical glandular field. Inflorescence an axillary or terminal
panicle (often reduced to a raceme), the branches glabrous or slightly lepidote,
each flower subtended by 2 large, membranaceous, caducous, red-violet bracts,
0.8-1.9 by 0.6-1.1 cm. Flowers with odor of garlic, calyx thin, cupular, un-
dulately 5-lobed, 9-21 mm long and 8-16 mm wide, mostly glabrous with raised
glandular areas, red-violet when fresh; corolla rose-red on lobes and upper
tube outside, the base of the tube yellow inside and out, upper tube within
yellow with pinkish hue; tubular-campanulate above a 1.5 cm by 0.4 cm tubular
base, 3.8-8.4 cm long and 0.8-1.9 cm wide at the mouth, the tube 3.5-6.0 cm
long, the lobes 0.6-2.7 cm long; the tube glabrous inside and out except for
multicellular, partially gland-tipped simple trichomes inside at the level of
stamen insertion, the lobes simple-puberulous; stamens didynamous, the abaxial

pair often exserted, the anther thecae straight, divaricate, 3-4 mm long, the
longer filaments 2.4-3.0 cm long, shorter filaments 1.1-2.2 cm long, the stam-
inode 1-2 mm long, inserted 11-16 mm from the base of corolla tube; pistil
4.2-4.9 cm long, the ovary linear-cylindric, 5 mm long and 1 mm wide, pustular-
lepidote, the ovules 2-seriate; disc annular-pulvinate, 1 mm long and 4 mm
wide. Capsule (including extralimital material) elongate-linear, flat, acute at
both ends, drying tan, glabrous but more or less punctate, the median nerve
not visible, 35-75 cm long and 2.6-3.8 cm wide; seeds thin, bialate, 2-2.8 cm
long and 7.5-11 cm wide, the wings pale white, subhyaline, poorly demarcated.
A plant of the tropical wet forest and wetter parts of the tropical moist
forest, this species ranges from Guatemala to Venezuela and Peru. It evidently
flowers somewhat sporadically, having been collected in flower twice in April
and once in September.
This species has been confused with P. hymenaea (DC.) A. Gentry and
P. alliacea (Lam.) A. Gentry but is both morphologically and ecologically
distinct.
BOCAS DEL TORO: Daytonia Farm, Cooper 450 (F). Western River, von Wedel 2700
(GH, MO, US). CHIRIQUI: Burica Peninsula near Punta de Piedra, Croat 22447A (MO).
PANAMA: Archaeological site near Madden Lake, Gentry & Tyson 5027 (MO, SCZ).
24. PARAGONIA
Paragonia Bur., Bull. Soc. Bot. France 19: 17. 1872. TYPE: Bignonia lenta Mart.
ex DC. = Paragonia pyramidata (L. C. Rich.) Bur.
Lianas; stems with 4 phloem arms in cross section; branchlets terete, lenti-
cellate, interpetiolar glands lacking or rare; pseudostipules subconical, basally
thick and pointed at the tip with the inner side appressed against the twig.
Leaves 2-foliolate, often with a minutely bifid, rarely minutely trifid, tendril,
the petiole with a glandular field on the upper side near the apex. Inflorescence
a large, terminal, many-flowered panicle. Flowers with the calyx more or less
cupular, truncate to somewhat lobed, lepidote, usually ciliate; corolla pale
lavender to magenta, tubular-campanulate, velvety moniliform-pubescent out-
side; anthers glabrous, the thecae straight, divaricate; pollen grains single,
3-colpate, the exine almost smooth; ovary narrowly cylindric, the walls rela-
tively thick, lepidote, the ovules 2-seriate in each locule. Fruit an elongate-
linear capsule, the valves convex, finely tuberculate with the texture of coarse
sandpaper; seeds thin, bialate, the wings chocolate brown, not demarcated from
seed body.
One species of continental tropical America ranging from Mexico to Argen-

tina; also reported on Guadeloupe.
1. Paragonia pyramidata (L. C. Rich.) Bur., Vidensk. Meddel. Dansk

Naturhist. Foren. Kjobenhavn 1893: 104. 1894.-FIG. 24.
Bignonia pyramidata L. C. Rich., Act. Soc. Hist. Nat. Paris 1: 110. 1792. TYPE: French
Guiana, LeBlond 292 (P-LA).
B. laurifolia Vahl, Eclog. Amer. 2: 44. 1798. TYPE: Trinidad, van Rohr s.n. (C).
B. ehretioides Chaam., Linnaea 7: 704. 1832. TYPE: Brazil, Sellow s.n. (fide Kunth, Fedde
Rep. Beih. 43: 635. 1928; not seen).
B. striata DC., Rev. Bign. (Bibl. Univ. Geneve) 22. 1838, nomen nudum.
B. rupestris Gardn., Jour. Bot. (Hooker) 1: 179. 1842. TYPE: Brazil, Rio de Janeiro, Gardn
78 (K).
B. sinclairii Benth., Bot. Voy. Sulphur 129. 1844. TYPE: Panama, Sinclair s.n. (K).
Zeyheria? surinamensis Miq., Linnaea 18: 250. 1844. LECTOTYPE: Surinam, Focke 230 (U,
holotype, excluding leaves of Cydista aequinoctialis; K, isotype).
Bignonia lenta Mart. ex DC., Prodr. 9: 159. 1845. TYPE: Brazil, Amazonas, Martius 2977
(M; G-DC, fragments).
B. martiusiana DC., Prodr. 9: 156. 1845. TYPE: Brazil, Para, Martius s.n. (1817) (BR).
Pachyptera dasyantha DC., Prodr. 9: 176. 1845. TYPE: Brazil, Rio Sao Francisco, Blanchet
2903 (G-DC, K).
P. perrottetii DC., Prodr. 9: 176. 1845. TYPE: French Guiana, Perrottet 2851 (G-DC).
Pachyptera striata DC., Prodr. 9: 176. 1845. TYPE: Brazil, Sao Paulo, Lund 783 (G-DC).
P. umbelliformis DC., Prodr. 9: 186. 1845. SYNTYPES: Brazil, Sao Paulo, Martius s.n. (M,
not seen); Rio Paraiba, Neuwied s.n. (M, not seen). "
Pithecoctenium reticulare DC., Prodr. 9: 197. 1845. TYPE: Brazil, without data (G-DC,
fragment).
Tabebuia pyramidata (L. C. Rich.) DC., Prodr. 9: 214. 1845.
Temnocydia elliptica Mart. ex DC., Prodr. 9: 156. 1845, pro syn.
T. lenta Mart. ex DC., Prodr. 9: 159. 1845, pro syn.
Arrabidaea dichasia Donn. Sm., Bot. Gaz. (Crawfordsville) 20: 6. 1895. TYPE: Honduras,
Santa Barbara, Thieme 5393 (NY, US).
Adenocalymma densiflora Rusby, Descr. 300 New Sp. S. Amer. P1. 122. 1920. TYPE: Boliv
Cataracts of Bopi River, Rusby 484 (NY, US).
Petastoma leiophyllum Kranzl., Fedde Repert. 17: 58. 1921. TYPE: Brazil, Parana, Dusen
8633 (K).
P. macrocalyx Kranzl., Fedde Repert. 17: 59. 1921. TYPE: Brazil, Sao Paulo,
not seen; K, photo).
Liana with somewhat swollen nodes, to 5 cm in diameter, the grayish

usually thick and soft-corky ridged, the ridges sometimes flattened and
lescent; stem with 4 phloem arms in cross section; branchlets terete with
warty lenticels, somewhat lepidote, sometimes slightly puberulous whe
especially at the nodes, interpetiolar glands mostly lacking, an interpetiolar
ridge often present; pseudostipules present, of 2-3 scales, basally thick and
pointed apically with the inner side appressed against the twig. Leaves 2-
foliolate, sometimes with a tendril (rarely incompletely 3-foliolate); leaflets
elliptic to elliptic-obovate or elliptic-ovate, acute, cuneate to rounded, 6.9-22
cm long and 2.3-12.5 cm wide, membranaceous to chartaceous, secondary veins
(3-) 4-6 on a wide, strongly ascending, generally lauraceous in appearance,
sometimes puberulous on the midvein beneath, lepidote above and beneath, a
few plate-shaped glands scattered near the midvein beneath, the main veins
often whitish when fresh, usually a uniform gray or olive when dry, when
fresh with a sweet smell when crushed; tendril simple or bifid, rarely trifid,
at the extreme tip, 5-17 cm long; petiolules 0.7-4.7 cm long, petiole 0.9-4.5 cm
long, above with an apical glandular field, lepidote to puberulous. Inflorescence
a many-flowered terminal panicle, the branches lepidote to puberulous. Flowers
with a sweet slightly spicy aroma, calyx cupular, irregularly truncate to some-
what bilabiate, lepidote, usually ciliate around the mouth with simple trichomes,
(3-)4-7 mm long, 4-10 mm wide; corolla pale lavender to magenta, inside
with the top and base of the tube white, tubular-campanulate, 2.7-5.5 cm
B '"~~~~~~~~~~~~~~~~~~~~~"
FIGuRE 24. Paragonia pyramidata (L. Rich.) Bur. A. Habit (X 1/2). [After Gentry
2903 (MO).] B. Fruit (X 1/2). [After Croat 5641 (MO).] C. Corolla split open (X 1/2).
-D. Pistil and disc ( X 1/2). E. Anther ( X 2). F. Ovary cross section ( x 81/2). [After
Gentry 1789 (MO).]
long and 0.8-2.2 cm wide at the mouth, the tube 2.1-4.0 cm long, the base
somewhat thickened, the lobes 0.5-1.5 cm long, thickly simple-pubescent with
moniliform trichomes outside and on the lobes inside, the tube mostly glabrous
inside, densely simple-pubescent with long multi-cellular, jointed trichomnes at
the level of stamen insertion; stamens didynamous, the anther thecae divaricate,
2-2.5 mm long, the filaments slightly short-puberulous near the base with a
conspicuous tuft of long, jointed trichomes at the extreme base, the longer
pair 1.6-1.9 cm long, the shorter pair 1.2-1.5 cm long, the staminode 4-6 mm
long, inserted 3-5 mm from the base of the corolla tube; pistil 1.9-2.0 cm long
the ovary linear, 4-5 mm long and 1.5 mm wide, strongly lepidote, the ovules
2-seriate; disc annular-pulvinate, 1.5-2.0 mm long and 4 mm wide. Capsule
linear, tapering at the ends, not strongly flattened, the valves convex, 32-61
cm long and 1.2-1.4 cm wide, to 0.7 cm thick, surface finely tuberculate,
resembling coarse sandpaper in texture; seeds 0.8-1.2 cm long and 2.1-4.4 cm
wide, the wings membranaceous, chocolate-brown, not sharply demarcated
from the seed body.
An abundant liana of the tropical wet forest and premontane wet forest,
P. pyramidata is also common in the tropical moist forest. Ecologically it is
equally well adapted to well-drained hillsides or swamps. It ranges from
Mexico to Brazil and Bolivia. This species flowers erratically throughout the
year but mostly during the dry season.
Distinguishing characteristics include the sweet smell of the broken leaves,
the typical pseudostipules, and the large lauraceous-looking leaflets. Vegeta-
tively it is most easily confused with Ceratophytum tetragonolobum (Jacq.)
Sprague & Sandw., which has similar leaves and pseudostipules but also con-
spicuous interpetiolar glands and trifid tendrils. Paragonia lacks interpetiolar
glandular fields and has simple or minutely bifid tendrils.
BOCAS DEL TORO: Changuinola on Rio Changuinola, Dwyer 2892 (MO). Hillside above
Almirante, Gentry 2694 (MO). Vicinity of Chiriqui Lagoon, von Wedel 1345, pro parte (GH,
MO). Fish Creek Lowlands, von Wedel 2377 (GH, MO, US). CANAL ZONE: Barro Colorado
Island, Brown 186 (F); Croat 4762, 4805 (both MO, NY, SCZ), 4917 (MO, SCZ), 5591
(MO, NY, SCZ), 5615 (MO), 5641 (MO, NY), 5645 (MO, SCZ), 5705 (NY, SCZ), 6538
(MO, NY, SCZ), 6584, 7131 (both MO), 7695 (MO, NY, SCZ), 7744, 7895 (both MO),
7896 (MO, SCZ), 8136, 8309A, 8370, 9106, 9227, 9510 (all MO), 10161 (MO, SCZ), 10212
(MO), 10358 (MO, SCZ), 11932, 14626, 14940 (all MO); Foster 689, 1090 (both PMA);
Gentry 430 (MO, WIS), 433, 451, 719 (all MO); Standley 40941 (US); Starry 150 (F, MO);
Wetmore & Abbe 210 (F); Wetmore & Woodworth 1 (F), 847 (A); Woodworth & Vestal
454 (A, F, MO), 576 (A, F). Near mouth of Rio Chagres, Allen 898 (K, MO, NY, US).
Matachin, Cowell 193 (NY). Pipeline Road, Gentry 1417, 1580 (both MO). Road K-19,
Gentry 1789 (MO). Pipeline Road, Gentry 1899, 2459 (both MO). Panama Station, Panama
Railroad, Hayes 413 (BM). Rio Gatuin, Hayes 915, 1043 (both NY). Las Cruces trail crossing
in Madden Forest, Harvey 5285 (F). Gamboa road, Jones 306 (US). Banks of Chagres River
below Gatuin, Maxon 4795 (BM, NY, US). Near Cruces, Seemann 400 (K). Chilibre River
to 1 mi. below Chilibre, Seibert 1513 (US). CHIRIQUI: W of Rio Chorchita, Gentry 5849
(MO). COCLE: El Valle de Anton, Allen 1751 (F, GH, MO, NY, US); Alston 8715 (BM);
Gentry & Dwyer 3572 (MO). Bismark above Penonome, Williams 263 (NY, US). COLON:
Rio Gatuncillo in vicinity of Santa Rosa, Allen 4166 (MO). W of Portobelo, Gentry 1764
(MO). DARIEN: Rio Ucurganti, Bristan 1157, pro parte (MO). Santa Fe, Duke 14251 (K,
MO). Rio Areti, Duke & Nickerson 14910 (MO). Vicinity of Jaque, Gentry 4100, 4183,
4187 (all MO). Cerro Chucula, drainage of Rio Pavarando, Gentry 4258 (MO). Rio Tuira
between Rios Paya and Aspave, Gentry 4406 (MO). El Real, Gentry 4501 (MO). Near
Cerro Pirre on Rio Parasenico, Gentry 4690 (MO). Near mouth of Pavarando on Sambui
River, Pittier 5577 (US). HERRERA: S of Ocui, Gentry 3129 (MO). LOS SANTOS: Vicinity of
Tonosi, Stern et al. 1875 (MO, US); Tyson et al. 2941 (MO, SCZ), 3102 (MO, PMA).
PANAMA: Lagarto Bay (Rio Maestro?), Barclay 2777 (BM). Cerro Campana, Gentry 1835
(MO). Rio Maestro, Gentry 2210 (MO). Rio Pasiga, Gentry 2311 (MO). Rio Piragua
(Tabardi ), Gentry 2527 (MO). Rio Piragua above confluence with Rio Bayano, Gentry
2588 (MO). Rio Corona, Gentry 2903 (MO). Near Rio Espave, Gentry 3710 (MO). Trail
from Rio Espave toward Rio Agua Clara, Gentry 3759 (MO). Along Rio Bayano just above
mouth of Rio Espave, Gentry 3782 (MO). Near last river before Rio Parti on trail from Rio
Bayano, Gentry 3801 (MO). Near Madden Lake, Gentry 5002 (MO). Rio Chame, Moran 29
(PMA). SAN BLAS: Near Puerto Obaldia, Gentry 1444 (MO). Mainland opposite Playon
Chico, Gentry 6384 (MO). Ailigandi River, Lewis et al. 171 (MO, SCZ). VERAGUAS: Vicinity
of Santa Fe, Gentry 2942, 3030 (both MO). WITHOUT LOCALITY, Cuming 1179 (BM);
Sinclair s.n. (K).
25. PARMENTIERA
Parmentiera DC., Rev. Bign. (Bibl. Univ. Geneve) 19. 1838. TYPE: P. edulis
DC. = P. aculeata (H.B.K.) Seem.
Zenkeria Reichb., Nom. 236. 1841, fide Index Kewensis.
Shrubs to quite large trees; often armed with a spine below each node.
Leaves opposite or subopposite, mostly palmately 3-5-foliolate, often with a
slightly winged petiole. Inflorescence a single flower or fascicle of 2-3 flowers,
usually borne cauliflorously on the old wood or trunk, sometimes terminally.
Flowers with the calyx spathaceous, split ventrally to near its base, inconspicu-
ously lepidote; corolla white or greenish-white, large, broadly campanulate to
infundibuliform, usually with a transverse fold across the lower side of the
throat; stamens subexserted, anthers glabrous, the thecae thick, straight, some-
what divergent; pollen grains 3-colpate, the exine finely granular; ovary cylin-
drical, the epidermis glandular and appearing more or less lepidote when dry,
the ovules multiseriate; disc annular-pulvinate. Fruit elongate-linear to oblong,
indehiscent, often longitudinally ridged, with a firm, fleshy outer cortex and
a fibrous-fleshy core; seeds between cortex and coxe, small, flattened, encircled
by a narrow vestigial wing.
Eight species of Central America ranging from southern Mexico to Colombia.
a. Small trees, the branchlets usually with 2 short incurved spines at most nodes; fruit
thick, conspicuously costate and curved, less than 17 cm long; cultivated - 1. P. aculeata
aa. Small to large trees, the branchlets without spines; fruit slender or thick, cylindric, not
conspicuously curved, more than 30 cm long; native but also cultivated.
b. Shrubby trees, usually several-branched near the base; petiole winged; fruits slender,
less than 2.4 cm wide, even when fresh, terete, not ribbed; moist forest, central
Panama and in cultivation -2. P. cereifera
bb. Erect trees or treelets, branched only in the crown; petiole unwinged (except in
seedlings); fruits cylindrical, thick, more than 4 cm wide when fresh, suboctagonal
in cross section, conspicuously ribbed; wet forest, central and western Panama
-----------------------------------------------------------------3. P. macrophylla
1. Parmentiera aculeata (H.B.K.) Seem. Bot. Voy. Herald 183. 1854.
Crescentia edulis Desv., Jour. Bot. (Desvaux) 4: 112. 1814. TYPE: (not seen; not Parmentiera
edulis DC.).
C. aculeata H.B.K., Nov. Gen. Sp. P1. 3: 158. 1819. TYPE: Mexico, Campeche, Bonpland s.n.
(P.).
C. edulis Moc. ex DC., Prodr. 9: 244. 1845, pro syn., non Desv.
Parmentiera edulis DC., Prodr. 9: 244. 1845. TYPE: Mexico, based on a Sesse and Mocifio
drawing (not seen).
Crescentia musaecarpa Zaldivar ex Heller, Reisen in Mexico 414. TYPE: Mexico (not seen).
Parmentiera foliolosa Miers, Trans. Linn. Soc. 26: 166. 1870. TYPE: Mexico, Schiede 1207 (K).
P. lanceolata Miers, Trans. Linn. Soc. London 26: 167. 1870. TYPE: Based on Crescentia
edulis Desv.
Small tree, branchlets terete, the nodes armed with a thick thorn subtending
each leaf, the leaves opposite or subopposite, mostly 3-foliolate, a few simple,
the leaflets elliptic-obovate, acute to obtuse, the base cuneate or attenuate, ter-
minal leaflet 3.4-4.7 cm long, lateral leaflets 2.1-4.7 cm long, with barbate cham-
bers in the nerve axils beneath; petiolules lacking, petiole 1.3-3.5 cm long,
strongly grooved on top but not winged. Inflorescence a few-flowered fascicle
or reduced to a single flower, terminal or axillary near the branch tips, or from
nodes of old wood. Flowers with the calyx basically spathaceous, sometimes
additionally split, ca. 3 cm long; corolla white, campanulate, 5-6 cm long and
2-2.5 cm wide at the mouth; stamens subexserted, the anther thecae somewhat
divergent, thick, ovary oblong-linear, lepidote, the ovules multiseriate in each
locule. Fruit to 17 cm long, more than 3 cm in diameter when fresh, curved,
costate with thick ribs.
No collections have been seen from Panama and the above description is
taken from Mexican material. Native to southern Mexico and northern Central
America, the species is occasionally cultivated outside its range. Reported by
Standley as cultivated in lowland Costa Rica, it is probably cultivated sporad-
ically in Panama. It is readily distinguished from the native species by its
axillary spines.
2. Parmentiera cereifera Seem. in Hook., Jour. Bot. (Hooker) 3: 302. 1841.

TYPE: Panama: Rio Chagres, Seemann s.n. (K). -FIG. 25A-E.
Small tree to 20 cm d.b.h. and 7 m tall, usually several-branched near the

base, the main branches strongly ascending, the crown open; twigs terete,
subpuberulous, the nodes often double with a pair of small leaves in the axil
of a larger pair, foliaceous pseudostipules sometimes present, 6-10 mm long
and 3-4 mm wide. Leaves opposite, 3-foliolate; leaflets elliptic to elliptic-
rhomboid, acute to acuminate, basally attenuate, the terminal leaflet 3.6-9.5
cm long and 1.4-4 cm wide, the lateral leaflets 3.0-6.5 cm long and 1.4-3.6
cm wide, membranaceous, secondary veins in 4-6 pairs, somewhat lepidote at
least beneath, simple-puberulous along the midvein above and beneath, espe-
cially in axils of lateral nerves, usually with a few plate-shaped glands along
the mid-vein beneath, drying olive to blackish; petiolules not differentiated,
petiole conspicuously winged, 2.4-6.2 cm long, the wing to 0.4 cm wide, widest
apically and basally attenuate, the midrib slightly scabrous. Inflorescence a
single cauliflorous flower or a fascicle of 2-3 flowers borne on abbreviated
short-shoots on larger branches or trunk, the pedicels lepidote, 0.9-1.6 cm long.
AA
FIGURE 25. Parmentiera.-A-E. P. cereifera Seem.-A. Inflorescence (X 1/2).-B. Ovary

cross section ( X 9 ).-C. Fruit ( X 3/8 ) .D. Fruit cross section ( X 1/2 ) .E. Leaves ( X 1/2 )
[After Gentry 5610 (MO).] F-J. P. macrophylla Standl. F. Leaves (X 1/2).-G. Corolla
(X x /2) -H. Ovary and disc (x X /4).-I. Calyx (X 1/2).-J. Fruit cross section ( X 1/2).
Flowers with musky odor, the calyx spathaceously split on the ventral side,
1.9-4.9 cm long and 0.9-2.9 cm wide, inconspicuously lepidote, at least at the
base; corolla pure white, tubular with a horizontal pleat midway across the
lower side of throat, slightly fleshy, 3.7-6.4 cm long and 1.8-2.9 cm wide at
the mouth, the tube 2.5-4.0 cm long, the lobes basally fused, 1.0-1.1 cm long,
mostly glabrous throughout, the epidermal cells apparently glandular, a few
stalked-lepidote trichomes at the level of stamen insertion inside and on lobes
outside; stamens subexserted, the anther thecae thick, partially divergent, 5-8
mm long, the anterior filaments 2.6-3.5 cm long, inserted 1.1-1.8 cm from base
of corolla tube, the posterior filaments 2.7-3.2 cm long, inserted 1.2-1.7 cm
from the base of the corolla tube, the staminode 1.0-1.9 cm long, inserted
0.9-2.3 cm from the base of corolla tube; pistil 3.1-4.8 cm long, the ovary linear-
cylindrical, 7-8 mm long and 1.5-2 mm wide, glabrous with glandular epi-
dermis, the ovules irregularly 8-10-seriate; disc annular-pulvinate, 1.5-2 mm
long and 6 mm wide. Fruit elongate-linear, subterete in cross section, 39-54
cm long and 1.0-2.4 cm wide, glabrous to sparsely lepidote, waxy yellow,
divided into a fleshy part surrounding the seeds and a fibrous central core
surrounded by the seeds; seeds small, thin, 3-4 cm long and 3-4 cm wide,
with a narrow but distinct mucilaginous wing.
A locally common small to medium-sized tree, P. cereifera is apparently

restricted in nature to the tropical moist forest of the Atlantic slope of Central
Panama. It is widely cultivated as a curiosity throughout the tropics. Flow-
ering throughout the year, it is pollinated by bats. The flowers do not open
until 4:00 or 5:00 in the evening. Fruit set is abundant and the waxy-fleshy
fruits fall beneath the parent tree at maturity. The fruit pulp is relished by
a number of mammals. According to Standley (Contr. U. S. Natl. Herb.
27. 1928) it is an important food for cattle in some parts of Panama. Dispersal
of the small, slippery, slime-coated seeds is probably effected by mammals
which eat the fruit. The common names "palo de velas" or "candle tree" refer
to the candle-like appearance of the fruit.
The elongate-fleshy fruits of Parmentiera separate it from other Crescentieae.
The compound leaves of Parmentiera are opposite in contrast to the likewise
3-foliolate but fasciculate leaves of Crescentia alata. Parmentiera cereifera is
distinguishable from other species of the genus by its tendency to divided trunks
and its narrower fruits. It is the only native Crescentieae of the non-coastal
tropical moist forest.
CANAL ZONE: Summit Garden, Correa et al. 1689 (PMA). Near Salamanca Hydrographic
Station in gorge of Rio Pequeni, Dodge et al. 16963 (MO). Boy Scout road near Madden
Lake, Gentry 2053, 2488, 5060 (all MO). Barbacoa, Hayes s.n. (BM). Matachin, Hayes 604
(NY). Dirt road near parking area for Madden Dam, Kirkbride & Elias 267 (NY, SCZ).
Vicinity of Madden Dam, Lewis et al. 21 (GH, MO, US). Vicinity of Gatuncillo, Piper 5611,
5641, 6009 (all US). Between Gorgona and Mamei, Pittier 2237 (BM, C, NY, US). Rio
Chagres, Seemann s.n. (K). Chilibre River below Chilibre, Seibert 1503 (MO, US). Rio
Pedro Miguel near E Paraiso Standley 29979 (US). Vicinity of Fort Sherman, Standley
31132 (MO, US). Mamei and Gorgona, Wagoner 35 (M). Madden Dam, Woodson et al.
1554 (A, MO, NY). Vicinity of Salamanca Hydrographic Station on Rio Pequeni, Woodson
et al. 1607 (A, MO, NY). Without locality, Gaillard s.n. (US), Gaillard s.n. (NY, US);
Johansen 16 (US).
3. Parmentiera macrophylla Standl., Publ. Field Columbian Mus., Bot. Ser.

4: 263. 1929.-TYPE: Panama, Bocas del Toro, Cooper 402 (F, K, US).-FIG.
25F-J.
P. macrophylla Standl., Trop. Woods 16. 12. 1928, nomen nudum.
Small to medium-sized tree to 30 cm d.b.h. and 25 m tall, the trunk usually

erect and unbranched for much of its length, the crown open; twigs terete,
glabrous to slightly and inconspicuously scabridulous. Leaves mostly opposite,
sometimes subalternate, often ? fasciculate, 3-foliolate; leaflets rhomboid-
elliptic, acuminate to acute, basally attenuate, the terminal leaflet 6.8-13.2 cm
long and 2.6-5.9 cm wide, the lateral leaflets 4.7-10.7 cm long and 1.8-4.9 cm
wide, membranaceous, secondary veins 5-7 on a side, somewhat lepidote or
deciduous-lepidote, especially beneath, otherwise glabrous except for na ciliate
fringe of short simple trichomes along the margins, well-developed chambers
at the vein axils beneath, usually with a few plate-shaped glands along the
midvein beneath, drying dark olive above, pale olive beneath; petiolules winged
but usually differentiated from the decurrent leaflet bases, the terminal petiolule
to 3.5 cm long, the lateral petiolules to 1.1 cm long, the petiole unwinged except
on immature plants, furrowed above, glabrous, 3.6-7.2 cm long. Inflorescence
of one or 2 flowers borne along the trunk, the pedicels lepidote. Flowers with
a musky odor, calyx spathaceously split and ventral side, 2.6-5.0 cm long and
0.8-1.2 cm wide, basally lepidote, otherwise mostly glabrous with a conspicu-
ously glandular epidermis when fresh; corolla pure white with a faint greenish
tinge outside; tubular-campanulate with an inconspicuous saccate bulge in the
floor of the throat, fleshy, 4.9-6.3 cm long and 1.5-2.5 cm wide at the mouth,
the tube 3.6-4.5 cm long, the 3 lower lobes 0.8-1.0 cm long, the upper 2
fused half their length; glabrous throughout except for a few glandular-lepidote
enations at the level of stamen insertion, the epidermis conspicuously glandu-
lar; stamens subexserted, the anther thecae thick, 1.5-2.0 mm wide, partially
divergent, 5-6 mm long, the anterior filaments 3.3-3.8 cm long, inserted 1.5-1.7
cm from the base of the tube, the posterior filaments 3.3-3.8 cm long, inserted
1.7-2.1 cm from the base of the tube, the staminode 4-16 mm long, sometimes
totally fused to the corolla tube; pistil 5.7-5.9 cm long, the ovary narrowly
cylindric, octagonal in cross section, 6-7 mm long and 2 mm wide, glabrous
with a glandular epidermis, the ovules ? 6-seriate; disc annular pulvinate,
1.5 mm long and 5 mm wide. Fruit cylindric with conspicuous longitudinal
ribs, suboctagonal in cross section, 44-50 cm long and ca. 6 cm wide when
fresh and 2.8 cm wide when dried, waxy yellow when fresh, drying black,
divided into a fleshy part outside the seeds and a fibrous core inside the
seeds; seeds small, thin, 3-6 cm long and 8-9 cm wide, with a distinct, narrow
mucilaginous wing when fresh.
A tree typical of wet upland areas, P. macrophylla is found in wet forest

and lower montane rain forest, mostly at elevations above 500 meters. It is
known only from Costa Rica and Panama. It flowers irregularly through most
of the year especially during the wet season. The fleshy fruit is eaten by
mammals.
Parmentiera macrophylla is quite variable in habit and ranges from a
straight-trunked tree, 30 cm in diameter and 25 m tall, to sexually mature
saplings, only 2.5 cm thick and 3 m tall. It is separated from Parmentiera
cereifera by its different ecological requirements, its much thicker, longitudi-
nally-ridged fruit, and at maturity by its unwinged petiole. Parmentiera trunci-
flora Standl. & L. 0. Wms. of Nicaragua is perhaps a form of P. macrophylla.
BOCAS DEL TORO: Region of Almirante, Cooper 402 (F, K, US). Chinguinola Valley,
Island Potrero, Dunlap 421 (F). Above Almirante, Gentry 2802, 2815 (both MO). Vicinity
of Chiriqui Lagoon, von Wedel 1101 (GH, MO). Water Valley, von Wedel 1421 (GH, MO,
US). Darkland, von Wedel 2619 (GH, MO, NY, US). COCLE: N rim of El Valle de Anton,
Allen 1782 (F, GH, MO, NY, US); Duke 12174 (MO); Dwyer et al. 4550 (MO); Gentry &
Dwyer 3663, 3658 (both MO); Hunter & Allen 307 (MO); Kirkbride 1089; Lewis et al.
1780 (MO); Luteyn 1214 (MO).
26. PHRYGANOCYDIA
Phryganocydia Mart. ex Bur., Bull. Soc. Bot. France 19: 18. 1872. TYPE: P.
corymbosa (Vent.) Bur.
Phrygiobureaua 0. Kuntze in Post & Kuntze, Lexic. 433. 1903.
Lianas; stems with 8 (sometimes 4) phloem arms in cross section; branch-

lets terete without interpetiolar glandular fields at the nodes; pseudostipules
inconspicuous or lacking, sometimes thick-foliaceous but early caducous on
young twigs. Leaves simple or bifoliolate, often with a simple tendril, more
or less lepidote but otherwise glabrous. Inflorescence a few-branched bifurcat-
ing panicle, sometimes reduced to a single flower. Flowers with the calyx
spathaceously split to below the middle, thin, the narrowing tip more or less
reflexed; corolla lavender to magenta, tubular-infundibuliform, lepidote outside;
anthers glabrous, the thecae straight, divaricate; pollen grains ecolpate, the
exine finely reticulate; ovary rounded-conical to narrowly cylindrical, lepidote,
the ovules 2-seriate in each locule; disc lacking. Fruit a capsule, the valves
parallel to the septum, the surface densely lepidote, smooth but microscopically
crystalline-rough, the median nerve indistinct; the capsule compressed and
linear-oblong with thin brown-winged, bialate seeds or short, ovid, and slightly
compressed with thick, corky wingless seeds.
Three species of continental tropical America ranging from Costa Rica

to Brazil and Bolivia. Phryganocydia can be recognized in flower by its
spathaceous calyx.
a. Leaves simple or 2-foliolate, coriaceous, the blades ovate, long-acuminate; capsule

ovoid (more than 3.7 cm wide), slightly compressed, appearing yellowish, the seeds
thick, corky, wingless, less than 3.5 cm wide; mangrove swamps and their fringes,
Pacific coast -2. P. phellosperma
aa. Leaves 2-foliolate, chartaceous, the blades elliptic to elliptic-ovate, acute or short-
acuminate; capsule linear-oblong (less than 2.3 cm wide), strongly compressed, appear-
ing grayish, the seeds thin, winged, more than 4.7 cm wide; well-drained sites, central
and eastern Panama ---------- 1. P. corymbosa
1. Phryganocydia corymbosa (Vent.) Bur. ex K. Schum. in Engler & Prantl,

Nat. Pflanzenf. 4(3b): 224, fig. 89H. 1894.-FIG. 26C-D.
Spathodea corymbosa Vent., Choix tab. 40. 1807; Mem. Math. Phys. Inst. NatI. France 1: 1-20.
1807. TYPE: Trinidad, Riedle s.n. (P).
S. laurifolia H.B.K., Nov. Gen. Sp. P1. 3: 114. 1819. TYPE: Venezuela, Sucre, Humboldt &
Bonpland s.n. (P).
S. orinocensis H.B.K., Nov. Gen. Sp. P1. 3: 147. 1819. TYPE: Venezuela, Orinoco, Humboldt
Bignonia coito Vell., Fl. Flum. 244. 1825; 6: tab. 19. 1827. TYPE: Brazil (not seen).
Spathodea coito (Vell.) DC., Prodr. 9: 204. 1845.
Phryganocydia dipleuropus Mart. ex DC., Prodr. 9: 204. 1845, pro syn. TYPE: Brazil, Para,
Martius s.n. (BR, M).
Spathodea platypoda DC., Prodr. 9: 204. 1845. TYPE: Brazil, Bahia, Blanchet 1511 (G-DC, P).
Macfadyena corymbosa (Vent.) Griseb., Bonplandia 6: 10. 1858.
M. coito (Vell.) Miers, Proc. Roy. Hort. Soc. 3: 200. 1863.
M. laurifolia (H.B.K.) Miers, Proc. Roy. Hort. Soc. 3: 200. 1863.
M. lepidota Seem. ex Miers, Proc. Roy. Hort. Soc. 3: 200. 1863, nomen nudum. TYPE: West
Indies (sic), Shakespear s.n. (drawing by Miers at P).
M. amazonica Miers, Proc. Roy. Hort. Soc. 3: 200. 1863, nomen nudum. TYPE: Brazil,
Santarem, Spruce s.n. (P).
M. orinocensis (H.B.K.) Miers, Proc. Roy. Hort. Soc. 3: 200. 1863.
M. ovata Miers, Proc. Roy. Hort. Soc. 3: 200. 1863, nomen nudum. TYPE: Colombia,
Cuming 1118 (P).
M. platypoda (DC.) Miers, Proc. Roy. Hort. Soc. 3: 200. 1863.
Phryganocydia coito (Vell.) Bur. ex Baill., Hist. P1. 10: 35. 1888.
Macfadyena bangii Rusby, Bull. Torrey Bot. Club 27: 72. 1900. TYPE: Bolivia, junction of
Beni and Madre de Dios Rivers, Rusby 1139 (NY, US).
Phryganocydia orinocensis (H.B.K.) Schnee in Pittier et al., Cat. Fl. Venez. 2: 401. 1947.
Liana to 5 cm in diameter, the bark smooth and fibrous to noticeably rough-

lenticellate; stem with 8 phloem arms in cross section; branchlets subterete,
finely longitudinally striate, lenticels inconspicuous, lepidote to glabrous, inter-
petiolar glandular fields lacking; pseudostipules usually inconspicuous or lack-
ing, occasionally thickly foliaceous, but small and soon deciduous on young
twigs. Leaves 2-foliolate, sometimes with a tendril; leaflets elliptic to ovate-
elliptic, basally mostly acute, acuminate, or rounded, 4.1-22 cm long and 1.9-11
cm wide, chartaceous to subcoriaceous, secondary veins (3-)4-7(-8) on a
side, often 3-veined from the base, densely minute-lepidote beneath, less so
above, otherwise glabrous, olive to gray-brown when dry, the midvein often
reddish-brown beneath; tendril simple, 7-16 cm long; petiolules 0.5-3.0 cm
long, usually slightly winged, petiole 0.6-3.3 cm long, strongly lepidote. Inflo-
rescence a small axillary or terminal panicle, the branches lepidote, somewhat
enlarged and often flattened at articulations. Flowers somewhat sweet-smelling,
calyx spathaceously split from near the tip to 1 cm from the base, the long
narrowing tip curved away from the corolla, apically blunt lepidote, 2.1-4.1
cm long to the tip and 0.6-1.4 cm wide at widest point; corolla lavender with
a white throat and magenta spot at the base of the upper corolla lobes or
magenta with a white throat; tubular-infundibuliform, 4.6-9.5 cm long and
0.9-2.4 cm wide at the mouth, the tube 3.1-6.8 cm long, the lobes 1.3-3.2 cm
long; lepidote inside and out, glandular-pubescent at and below the level of
stamen insertion inside, stamens didynamous, the anther thecae divaricate, each
3.5-4.5 mm long, the longer filaments 1.8-2.1 cm long, shorter filaments 1.0-1.3
An.~~
FIGURE 26. Phryganocydia.-A-B. P. phellosperma (Hemsl.) Sandw. A. Habit (X

[After Stern et al. 5$9 (MO)] B. Fruit (X '/2). [After Dwyer 1093 (MO).]- C-D. P.
corymbosa (Vent.) Bur. ex K. Schum.-C. Ovary (X 21/2).-D. Ovary cross section (X 9).
cm long, the staminode 2-3 mm long, inserted 0.8-1.2 mm from base of corolla
tube, pistil 3.8-4.1 cm long, ovary linear, 3-4 mm long and 1 mm wide, lepidote,
the ovules 2-seriate in each locule; disc obsolete. Capsule linear-oblong, acute,
flattened, 13-53 cm long and 1.4-2.3 cm wide, densely lepidote, appearing
grayish, crystalline-rough under a lens, neither margins nor midrib pronounced;
seeds 1.5-2.0 cm long and 4.7-7.3 cm wide, the wings membranaceous but
brown, not demarcated from the seed body.
The distribution of this species is enigmatic. It is the commonest species

of Bignoniaceae in the Canal Zone and much of eastern Panama and is abundant
through the tropical moist forest of eastern Panama, occurring also to some
extent in tropical wet forest. However, it has never been collected west of
Cocle Province and is apparently absent from Costa Rica. It is thus the only
species of Bignoniaceae of the tropical moist forest life zone which occurs in
Panama but not Costa Rica; this is doubly surprising considering its extreme
abundance. To the south its range extends to Brazil. The species flowers
throughout the year. Fruit maturation takes about two months, and seed
release occurs, like flowering, throughout the year.
The fruit of this species is similar to that of Cydista aequinoctialis, but the
valves tend to be gray rather than black as in C. aequinoctialis; they also curl
upon falling while the valves of the latter merely bend up at the sides. Under
a lens the fruits of this species have an irregular, almost crystalline surface
texture, while those of C. aequinoctialis are smooth-textured. The seeds are
also distinguishable, but with difficulty-in this species they have a submar-
ginal hilum with the seed body relatively far from it, while seeds of C. aequin-
octialis have a nearly marginal hilum with the body of the seed crowded
against it. In sterile condition P. corymbosa can be distinguished best by its
tendency to flattened, almost alate, petiolules and terete, smooth, light gray
twigs. The leaves have a characteristic shape and waxy appearance that be-
comes readily recognizable after sufficient exposure.
CANAL ZONE: Barro Colorado Island, Aviles 83 (F), 90 (MO), 990 (F); Bailey & Bailey
314 (F), 608 (F, GH); Bangham 387 (A, F, US); Brown 20 (F); Croat 4975 (MO, SCZ),
5405 (MO), 5459, 5542, 5590, 6089, 6591 (all MO, SCZ), 7119 (MO), 7267 (MO, SCZ),
7870, 7988, 8168, 8272, 8437, 12684, 13228 (all MO); Foster 964 (PMA); Frost 94 (F);
Gentry 431 (MO, WIS), 711, 1989 (both MO); Hayden 19 (MO); Kenoyer 526 (US);
Salvoa 849 (A); Shattuck 466, 515, 784, pro parte (all F, MO), 990 (MO, US), 1042 (F,
MO); Standley 51126 (A, US); Starry 210 (F, MO); Wetmore & Woodworth 32, 73 (both
F), 853, 876, 878, 909 (all A); Woodson & Schery 976 (MO); Woodworth & Vestal s.n. (F),
376 (A, F), 394 (A, MO), 577 (A). Near Fort Kobbe, Allen 2020 (GH, MO). Rio Chilibre
between Juan Mina and Madden Dam, Bartlett & Lasser 16414 (MO). Vicinity of Albrook
tower, Blum 436 (MO, SCZ); Cowell 206 (NY). Gatuin, Cowell 313 (NY, US). Road S-li
NW of Escobal, Croat 12459 (MO, SCZ). Fort Sherman hill C-6, Duke 4403 (GH, K, MO).
Ancon Hill, Duke 4580 (GH, K, MO). Albrook site, Dwyer 7114 (GH, MO, US). Las Cruces
trail in Madden Forest, Gentry 1382 (MO). Pipeline road, Gentry 1416, 1584 (both MO).
Boy Scout road near Madden Lake, Gentry 2056 (MO). Between Summit and Gamboa,
Greenman & Greenman 5262 (MO). Paraiso Station, Panama Railroad, Hayes 158 (GH),
222 (BM, K). Summit, Hayes 591 (NY). Ancon Hill, Killip 3055 (F, US). Pipeline road,
Lewis & Sharp 32 (MO). Near Fort Randolph, Maxon & Harvey 6502 (US). Around Culebra,
Pittier 2150 (US). Cerro Gordo near Culebra, Standley 25952 (MO, US). Summit, Standley
26959 (US). Corozal, Standley 27350( (US). Las Cascades Plantation near Summit, Standley
29680 (US). East Paraiso, Standley 29836 (US). Vicinity of Summit, Standley 30093 (A,
US), 30104 (US). Albrook Air Force Base, Tyson s.n. (SCZ), 1412 (MO, SCZ). Pipeline
road, Tyson 1482 (MO, SCZ). Fort Sherman along road S-5, Tyson & Dwyer 1226 (MO).
Along road between Cocoli and Contractor's Hill, Tyson & Dwyer 6489 (SCZ). Without
locality, Johansen 40 (GH, US). COCLE: Bismarck above Penonome, Williams 550 (US).
COLON: Santa Rita Ridge, Gentry 1863, 1885 (both MO). E of Rio Piedras on road to
Portobelo, Gentry 2189 (MO). Between' Rio Piedras and Puerto Pilon, Lewis et al. 3213 (MO,
SCZ); Nee & Mori 3668 (MO). DARIEN: Trail between Pinogana and Yavisa, Allen 279
(MO). Vicinity of Boca de Cupe, Allen 888 (F, GH, MO, NY, US). Vicinity of Santa Fe
on Rio Sabana, Duke 4112 (MO). Between Rios Membrillo and Subcuti, Duke 8588 (MO).
Rio Balsa between Rio Areti and Manene, Duke 8781 (MO). Vicinity of Piflas, Duke 10635
(MO). Rio Balsa between Manene and Tusijuanda, Duke 13565, 13568 (both MO). Rio
Morti, Duke 14179 (F, MO, NY). Camp Lagarto, Duke 15201 (MO). Around La Palma,
Gentry 3923 (MO). Isla Lagarto in Golfo de San Miguel, Gentry 3998 (MO). Second large
point just before Punta Alegre NE of Garachine, Gentry 4062 (MO). S of Jaque, Gentry
4104 (MO). Ridge from top of Cerro Pavarando to top of Cerro Derumba, Gentry 4205, 4215
(both MO). S of La Palma, Gentry 4304 (MO). Rio Paya from Rio Tuira to village of
Paya, Gentry 4380 (MO). Rio Tuira between Rios Paya and Aspave, Gentry 4400 (MO).
Below Boca de Cupe on Rio Tuira, Gentry 4488 (MO). Below Boca de Yape on&Rio Tuira,
Gentry 4489 (MO). Midway between Yavisa and El Real on Rio Chucunaque, Gentry 4506
(MO). El Real, Gentry 4524, 4578 (both MO). Near Cerro Pirre on Rio Parasenico, Gentry
4687 (MO). Vicinity of La Palma, Pittier 6693, 6978 (both US). PANAMA: Rio Maestra,
Allen 48 (A, MO). Vicinity of Bejuco, Allen 976 (MO, US). Vicinity of Campana, Allen
3571 (BR, C, GH, K, MO, NY, P, U, US). Cerro Campana, Correa & Dressler 391 (PMA).
Between Capira and Chorrera, Dodge 10714 (F, MO). Along road between Panama and
Chepo, Dodge 10730 (MO). Along banks of Quebrada La Palma and cafion of Rio Chagres,
Dodge & Allen 17346 (MO). Rio Caffita, Duke 3812 (MO, SCZ). E of Chepo, Duke 4046
(MO), 4079 (K, MO). Rio Pita above confluence with Rio Maestra, Duke 4737 (MO).
Along Rio Terable and road to El Llano, Duke 5660 (CH, MO). San Jose Island, Duke
12555 (MO); Erlanson 7 (BM, NY, US); Harlow 4 (US), 30 (GH, US); Johnston 164 (BM,
GH, MO, US), 317 (BM, MO, US), 1263 (GH); Tyson & Loftin 5063 (SCZ). Near Caffitas,
Gentry 1406, 4966 (both MO). Rio Pasiga, Gentry 2202, 2259, 2299 (all MO). Rio Maestra,
Gentry 2211 (MO). E of Chilibre, Gentry 2508 (MO). Rio Piragua (Tabardi), Gentry 2529
(MO). Icanti, Gentry 2616 (MO). 10-15 km from Rio Bayano crossing on trail to Santa Fe,
Gentry 3833 (MO). W of Chepo, Gentry & Tyson 1639 (MO). Dam site on Rio Bayano,
Gentry & Tyson 1665 (MO). Punta Paitilla, Heriberto 211, 228 (both US). Chiman, Lewis
et al. 3342 (MO, SCZ). Trapeche Island, Miller 1887 (US). Near Chepo, Pittier 4653 (BM,
C, GH, NY, US). Near road to Cerro Azul, Rivas 10 (MO, PMA). Las Cumbres, Rivera 13
(GH; MO, PMA). Villa Rosario, Saldana 31 (PMA). Vicinity of Arenoso on lower Rio
Trinidad, Seibert 606 (A, K, MO, NY). Perlas Islands, without locality Stevenson 409 (F).
SAN BLAS: Isla Soskatupu, Duke 8954 (MO). Ailigandi, -Dwyer 2831 (MO). Above Puerto
Obaldia, Gentry 1489 (MO). Mainland opposite Playon Chico, Gentry 6416 (MO). Soskatupu,
Kirkbride 191 (MO, NY, SCZ). WITHOUT LOCALITY: Duchassaing s.n. (P).
2. Phryganocydia phellosperma (Hemsl.) Sandw., Kew. Bull. 1940: 302. 1941.

-FIG. 26A-B.
Macfadyena phellosperma Hemsl., Biol. Centr. Amer. Bot. 2: 492. 1882. TYPE: Panama,
Canal Zone, Hayes 81 (BM, K).
Liana to 2.5 cm in diameter, the bark smooth, gray, longitudinally lightly

striated; stem with 4 phloem arms in cross section; branchlets subterete or some-
what flattened, reddish-brown, lepidote, lacking conspicuous lenticels, inter-
petiolar glandular fields lacking; pseudostipules lacking. Leaves 2-foliolate,
occasionally simple, sometimes with a tendril; leaflets ovate, long-acuminate,
shallowly cordate to rounded, 5.1-13 cm long and 3.5-8 cm wide, coriaceous
to subcoriaceous, secondary veins 4-6 on a side, lepidote above and beneath,
otherwise glabrous, grayish olive when dry, midvein below reddish-brown, the
lateral veins inconspicuous; tendril simple, 8-15 cm long; petiolules 0.8-2.2 cm
long, petiole 0.8-3.8 cm long, + lepidote. Inflorescence 1-2 terminal flowers,

the pedicels lepidote. Flowers fragrant, sweet-smelling, the calyx spathaceously
split from near the tip to 2 mm from the base, the blunt tip bent slightly away
from the corolla, inconspicuously lepidote, 1.8-2.8 cm long and 0.5-1.1 cm
wide at widest point; corolla lavender with a white throat and magenta spot
at the base of the upper corolla lobes, the throat with 9 very fine magenta
nectar guides; tubular-infundibuliform, 4.2-7 cm long and 1.1-2.3 cm wide
at the mouth of tube, the tube 3-4.7 cm long, the lobes 1.2-2.5 cm long; incon-
spicuously lepidote outside, the lobes pubescent on the margins and sparsely
within, the tube mostly glabrous inside, glandular-pubescent at and below
3-4 mm long, the longer filaments 1.3-1.5 cm long and inserted 0.4-0.5 cm
from the base of the tube, the shorter filaments 0.8-1.1 cm long and inserted
0.2 cm from base of tube, the staminode to 4 mm long, inserted 1 mm from
base of tube or absent; pistil 2.0-2.2 cm long, the ovary rounded-conical, 1.5
mm long and 1.5 mm wide, 1.5 mm thick, lepidote, the ovules 2-seriate; disc
lacking. Capsule (including extralimital material) broadly ovoid to almost
suborbicular, slightly compressed, 3.5-6.5 cm long and 3.7-5.6 cm wide, densely
lepidote, crystalline-rough under a lens, drying yellowish; seeds thick, corky,
wingless, with a loose epidermis which pulls away from the seed body, 2.0-2.7
cm long and 2.2-3.5 cm wide, the hilum large, to 8 mm by 15 mm.
Ecologically restricted to mangrove swamp habitats, P. phellosperma occurs

only along the Pacific coast from Colombia to Costa Rica. It is locally abun-
dant but rarely collected. It flowers mostly in the wet season from late May
through July, although occasional flowers are produced throughout the year.
The seeds are water-dispersed.
It is closely related to P. corymbosa and quite probably derived from it.
The two similar congeners occasionally overlap ecologically at the edge of
the mangrove consociation, but the much -more coriaceous, long-acuminate
leaflets of P. phellosperma serve to distinguish it. The flowers of the two species
are nearly identical, although those of P. phellosperma are usually smaller and
have a more blunt-tipped calyx. The fruits are much shorter and thicker than
those of P. corymbosa. Thus Phryganocydia parallels Tabebuia and other gen-
era with species specialized to live in mangrove consociations; these specializa-
tions are reflected in corky, more or less wingless seeds which are water-dispersed
and short, thick fruits.
CANAL ZONE: Near Balboa, Alston 8830 (BM, K). Ancon, Bangham 609 (A). Miraflores
locks, Dwyer 1093 (MO). W end of Thatcher Ferry Bridge, Gentry 745 (MO, SCZ). Farfan
Beach road, Gentry & Dwyer 3551 (MO). Rio Grande swamp, Hayes 81 (BM, K). Near
Miraflores locks, Stern et al. 59, 80 (both GH, K, MO, US). Rio Grande, Woodson et al. 761
(A, MO, NY). DARIEN: Chepigana, Duke 279 (MO). Along Rio Tuira 5-15 mi. down-
stream from El Real, Duke 4947 (BM, GH, MO, US). Punto Sabanas across bay from La
Palma, Gentry 3959, 4089A (both MO). Isla Encanto in Golfo de San Miguel, Gentry 4008
(MO). Rio Sabanas below Santa Fe, Gentry 4082 (MO). Punta Triunfo up Rio Sabanas,
Gentry 4085 (MO). Mouth of Rio Jaque, Gentry 4190 (MO). PANAMA: San Jose Island,
Mouth of Rio Marino, Johnston 1296 (GH). Playa del Palma near San Carlos beach, Lewis
et al. 1502 (GH, MO, US). Taboga Island, Pittier 3611 (GHl, US).
27. PITHECOCTENIUM
Pithecoctenium Mart. ex Meisn., Gen. 1: 300; 2: 208. 1840. LECTOTYPE: P.

echinatum (Jacq.) Baill.
Liana; stem with 4 phloem arms in cross section; branchlets acutely hexa-
gonal with detachable ribs, without interpetiolar glandular fields; pseudo-
stipules spathulate, not persistent. Leaves 2-3-foliolate, the terminal leaflet
often replaced by a trifid tendril, this often divided to at least 15-fid, some
of the tendril-ends often with thickened discs which attach the vine to a
support. Inflorescence a terminal raceme or racemose panicle. Flowers with
the calyx cupular, truncate, thick, more or less minutely 5-denticulate, lepidote
and puberulous; corolla white, relatively thick, tubular-campanulate, densely
pubescent outside, anthers glabrous, the thecae straight, divaricate; pollen
grains esulcate, the exine alveolate, ellipsoid-cylindric, contracted above the
disc, densely appressed simple-pubescent, the ovules multiseriate in each locule;
disc annular-pulvinate. Fruit a thick, woody, somewhat compressed, elliptic
capsule, the valves parallel to the septum, densely echinate; seeds alate with
a thin body surrounded by a broad clearly demarcated hyaline-membranaceous
wing.
Three or four species in continental tropical America ranging from Mexico

to Argentina and Brazil.
1. Pithecoctenium echinatum (Jacq.) Baill., Hist. P1. 10: 8. 1888.-FIG. 27.
Bignonia echinata Jacq., Enum. P1. Carib. 25. 1760; Sel. Stirp. Amer. 183, tab. 176, fig. 52.
1763. TYPE: Colombia, Cartagena (not seen).
B. tiliaefolia H.B.K., Nov. Gen. Sp. P1. 3: 136. 1819. TYPE: Venezuela, Orinoco (?), Humboldt
& Bonpland (P).
B. squalus Vell., Fl. Flum. 230. 1825; 6: tab. 13. 1827. TYPE: Brazil (not seen).
B. phaseoloides Cham., Linnaea 7: 698. 1832, fide Bur. & K. Schum. TYPE: Brazil, Sellow
s.n. (not seen).
B. hexagona DC., Rev. Bign. (Bibl. Univer. Geneve) 21.- 1838, nomen nudum.
B. muricata Moc. ex DC., Rev. Bign. (Bibl. Univ. Geneve) 22. 1838, nomen nudum.
Pithecoctenium aubletii Splitg., Tijdschr. Natuurl. Gesch. Physiol. 9: 12. 1842. TYPE: Based
on Bignonia echinata Aubl., the fruit of which is P. echinatum but the leaves and flowers
Arrabidaea inaequalis (DC. ex Splitg.) K. Schum.
P. phaseoloides (Cham.) Schenk in Schimper, Bot. Mittheil. 4: 193. 1842.
P. cordifolium Mart., Flora 24(2): Beibl. 48. 1841. TYPE: Brazil, Cujaba, Martius 516
(MO, NY).
P. hexagonum DC., Prodr. 9: 195. 1845. TYPE: Mexico, Chalco, Andrieux 222 (G-DC, K).
P. lundii DC., Prodr. 9: 196. 1845. TYPE: Brazil, Sao Paulo, Lund 876 (G-DC).
P. muricatum Moc. ex DC., Prodr. 9: 194. 1845. TYPE: Mexico (not seen).
P. squalus (Vell.) DC., Prodr. 9: 194. 1845.
P. tribrachiatum Loes., Fedde Repert. 16: 209. 1919. SYNTYPES: Mexico, Veracruz, Schenck
860, 861 (not seen).
P. glaucum Rusby, Mem. New York Bot. Gard. 7: 354. 1927. TYPE: Bolivia, Rurrenabaque,
Cardenas 2063 (K, NY, US).
Liana to at least 10 cm in diameter, the bark loose, fibrous, vertically striate,

becoming a spirally twisted bush rope; stem with 4 phloem arms in cross sec-
tion; branchlets sharply hexagonal with the angles ribbed, lepidote with occa-
sional simple trichomes, the nodes without interpetiolar glandular fields, some-
- MC
FIGURE 27. Pithecoctenium echinatum (jacq.) K. Schum.-A. Habit (X 2/5). [After

Croat 5804 (MO).] B. Fruit ( X 2/5). [After Croat 7022 (MO).] C. Corolla split open
( X 3/a-).D. Fresh corolla in side view ( X 3/5).-E. Pistil ( X 3/a-).-F. Ovary and disc
( X 11/f-).-G. Ovary cross section ( X 63/5). [After Gentry 2701 (MO).]
times with a ridge connecting opposite petioles; pseudostipules elongate, ca.

10 mm long and 2 mm wide, caducous on mature twigs. Leaves 3-foliolate or
2-foliolate with a tendril or tendril scar; leaflets ovate to subrotund, acuminate,
cordate, 3.3-18 cm long and 2.0-14.7 cm wide, membranaceous, palmately
veined at the base, the secondary veins 4-6 on a side, strongly lepidote above
and beneath, variably pubescent above and beneath with simple trichomes
usually at the base of main nerves and sometimes scattered over the surface,
plate-shaped glands in the axils of basal nerves beneath, more or less olive
when dry; tendril trifid to 15-fid (perhaps more), 4-12 cm long to first
branching, the ultimate tips often thickened into discs which attach to bark
of supporting tree; petiolules and petiole tetragonal, the former 1.6-7.0 cm
long, the latter 2.9-7.2 cm long, lepidote and usually simple-pubescent at least
on the angles. Inflorescence a few- to 15-flowered terminal raceme or racemose
panicle with small, caducous bracts, the rachis and pedicels lepidote and simple-
pubescent. Flowers with musky odor, calyx cupular, coriaceous, truncate to
inconspicuously 5-denticulate with submarginal teeth, lepidote and densely
pubescent, 8-12 mm long and 9-11 mm wide, eglandular; corolla white with
yellow throat, relatively thick and fleshy, especially the basal 0.9-1.2 cm,
tubular-campanulate but usually bent at 900 angle in the middle, 3.6-6.1 cm
long and 1.0-1.8 cm wide at the mouth, the tube 1.2-1.7 cm to the bend,
2.2-3.5 cm long above the bend, the lobes 0.6-1.7 cm long, densely pubescent
with more or less moniliform trichomes outside, inside moniliform-pubescent
on the lobes, the tube glabrous except for dense moniliform trichomes at the
3-4 mm long, the longer filaments 1.7-2.1 cm long, shorter filaments 1.2-1.7 cm
long, the staminode 0.6-0.8 mm long, inserted 0.8-1.2 mm from the base of the
corolla tube; pistil 3.3-3.5 cm long, stigma heart-shaped, 4 mm by 4 mm, ovary
short-cylindric, swollen at the middle, 5-6 mm long and 4 mm wide at the
middle, 3 mm wide at the base and apex, densely simple-pubescent with multi-
cellular trichomes, the ovules + 15-seriate in each locule; disc annular-pulvinate,
2-3 mm long and 6 mm wide. Capsule rounded-oblong to elliptic, 12-31 cm
long and 5.2-7.5 cm wide, to 3.5 cm thick, the surface strongly echinate-
tuberculate, sometimes with a few simple trichomes at the base of tubercles;
seeds 2.3-4.1 cm long and 4.5-9.5 cm wide, the wings surrounding the seed
body, membranaceous, hyaline except the extreme base, clearly demarcated.
Both geographically and ecologically this is one of the most wide-ranging

species of Bignoniaceae, although never extremely abundant. It occurs from
Mexico to Argentina. It flowers during the wet season, mostly in May and
June but was collected in full bloom in several places at Almirante, Bocas del
Toro, in late November. The fruits take almost a year to mature, and seeds
are released in mid dry season from February through April. Robin Foster
(personal communication) considers P. echinatum to be the most effectively
wind-dispersed species occurring on Barro Colorado Island. The lightness and
aerodynamic stability of a floating seed of this species make its path of descent
a thing of beauty even in perfectly still air; with a breeze a seed remains air-
borne indefinitely. The dispersability of its seeds must be in large part respon-
sible for the great geographic distribution of the species.
This species is distinguished from all other Central American species except
Amphilophium by its sharply hexagonal twigs. Its elongate pseudostipules,
usually much-branched tendrils and lack of dendroid trichomes serve to dif-
ferentiate it from the two species of Amphilophium. The bent white corolla
and large echinate fruits are distinctive. "Peine de mono, monkey's hair-
brush."
BOCAS DEL TORO: Vicinity of Almirante, Gentry 2701, 2752, 2783 (all MO). CANAL
ZONE: Barro Colorado Island, Croat 5804 (MO, NY, SCZ), 5837 (MO), 7022 (MO, SCZ),
9418 (MO, SCZ), 10851, 14066 (both MO), 15115 (MO); Foster 840 (MO, PMA), 1173
(MO, PMA); Wetmore & Woodworth 36 (A, F). Las Cruces Trail in Madden Forest,
Gentry 1381 (MO). Road K-19, Gentry 1791 (MO). Boy Scout road near Madden Lake,
Gentry 2059, 2060 (both MO). Fort Sherman, Gentry & Tyson 4841 (MO)., CHIR
Finca Boquete, Oppenheimer & Hladik 209 (MO). COLON: Ridge W of Portobelo, Gentry
1762 (MO). DARIEN: Near Jaque, Gentry 4177 (MO). Slopes of Cerro Chucula, drainage
of Rio Pavarando, Gentry 4263 (MO). Below Paya on Rio Paya, Gentry 4392 (MO). El
Real, Gentry 4504 (MO). PANAMA: Rio Maestra, Gentry 2221, 2230A (both MO). Rio
Pasiga, Gentry 2252, 2270 (both MO). Rio Piragua (Tabardi), Gentry 2532 (MO). Icanti,
Gentry 2597 (MO). Near Rio Espave, Gentry 3789 (MO). Farfan Beach road, Gentry &
Dwyer 3556 (MO). SAN BLAS: Above Puerto Obaldia, Gentry 1476, 155-9 (both MO).
VERAGUAS: Above Santa Fe on slopes of Cerro Tute, Gentry 6256 (MO).
28. PLEONOTOMA
Pleonotoma Miers, Proc. Roy. Hort. Soc. 3: 184. 1863. LECTOTYPE: P. jasmini-
folia (H.B.K.) Miers.
Lianas; stems with 4 phloem arms in cross section, the extreme center
hollow; branchlets acutely tetragonal with detachable ribs, without interpetiolar
glandular fields; pseudostipules foliaceous or absent. Leaves biternate or tri-
ternate or ternately bipinnate or tripinnate, the terminal pinna often replaced
by a trifid tendril. Inflorescence an axillary or terminal raceme. Flowers with
the calyx cupular, truncate, usually minutely denticulate; corolla white to pale
yellow, narrowly campanulate-infundibuliform, the tube glabrous or variously
pubescent in upper part; anthers glabrous, the thecae straight, divaricate; pollen
grains 3-colpate, the exine microreticulate; ovary oblong, more or less lepidote
or pubescent, the ovules 2-seriate in each locule; disc cupular-pulvinate. Fruit
an elongate-linear to elongate-oblong compressed capsule, the valves parallel
to the septum, more or less smooth, drying blackish, the median nerve incon-
spicuous; seeds thin, bialate, the wings membranaceous, brown, poorly demar-
cated from the seed body.
Ten to twelve species in continental tropical America ranging from Costa

Rica to Argentina.
1. Pleonotoma variabilis (Jacq.) Miers, Proc. Roy. Hort. Soc. 3: 184. 1963.
-FIG. 28.
Bignonia variabilis Jacq., Hort. Schoenb. 2: 45, tab. 212. 1797. TYPE: Venezuela, Caracas,
Jacquin s.n. (W).
FIGURE 28. Pleonotoma variabilis (Jacq.) Miers. A. Habit (x '/2). [After Croat
5607 (MO).]-B. Fruit (X '/2). [After Gentry 3972 (MO).]
Liana to at least 3 cm in diameter, bark fibrous; stem with 4 phloem arm

in cross section, extreme center hollow; branchlets acutely tetragonal with the
angles ribbed, brown to green with ribs usually blackish, the nodes without
interpetiolar glandular fields, a linear ridge connecting opposite petioles; pesudo-
stipules narrow, 3-5 mm long and 1 mm wide, often deciduous. Leaves 3-
ternate or 2-ternate with a tendril or tendril scar; leaflets elliptic to elliptic-

ovate, acute to acuminate, widely cuneate to asymmetrically truncate, 2.1-16
cm long and 0.8-9.5 cm wide, secondary veins 5-7 on a side, with scattered
lepidote scales, slightly puberulous along the veins above, beneath usually
with simple trichomes in the axils of lateral nerves and sometimes along the
main veins, greenish when dry; tendril trifid, 2-14 cm long to branching, the
3 arms 0.1-2.1 cm long; petiole, petiolules, and subpetiolules all strongly angled,
lepidote to puberulous, the terminal subpetiolule 0.3-4.8 cm long, laterals
0.2-2.2 cm long, the terminal petiolule 2.5-6.5 cm long, lateral 1.2-6.5 cm long,
the petiole 2.1-6.7 cm long. Inflorescence a short, long-pedicelled raceme,
usually terminal, the pedicels and peduncle mostly glabrous, somewhat simple-
pubescent at the nodes. Flowers with the calyx cupular, more or less truncate,
5-9 mm long and 5-8 mm wide, mostly glabrous, more or less ciliate, sometimes
with scattered lepidote scales and with glands in the upper third; corolla white
or cream with light yellow tube, tubular-infundibuliform, 6.4-10.2 cm long
and 1.3-2.7 cm wide at the mouth, the tube 4.5-7.0 cm long, the lobes 1.4-2.2
cm long, tube glabrous outside, the lobes glandular-lepidote and puberulous
with plate-shaped glands outside at the base, tube glabrous within except for
long simple multicelled trichomes at and below the level of stamen insertion,
in bud long-pointed, conspicuously pubescent at the tip; stamens didynamous,
the anther thecae divaricate, ca. 5 mm long, the longer filaments 2.6-2.8 cm
long, shorter filaments 1.7-1.9 cm long, the staminode 4-5 mm long, inserted
1.9-2.1 cm from the base of the corolla tube; pistil 4.3-5.5 cm long, the ovary
linear-oblong, 4 mm long and 1.5 mm wide, lepidote, the ovules 2-seriate in
each locule; disc annular-pulvinate, 1.5-2 mm long and 3-4 mm wide. Capsule
linear to linear-oblong, acute, somewhat compressed, 15-30 cm long and 1.7-2.7
cm wide, 3-7 mm thick, lepidote, slightly and irregularly warty-tuberculate
with shallow projections, drying dark; seeds 1.1-1.5 cm long and 3.0-5.0 cm
wide, the wings membranaceous, brown, poorly demarcated from the seed body.
A vine of the tropical moist forest, premontane wet forest, and tropical wet
forest, P. variabilis ranges from Costa Rica to Venezuela and Trinidad. It
flowers during the dry season from February to May. Seed release is during
the dry season, and young plants are relatively common.
This is the only vine in the family in Central America to have bicompound
leaves. The leaves and strongly tetragonal stem make P. variabilis unmistakable,
even in vegetative condition.
BOCAS DEL TORO: Along railroad between Almirante and Changuinola, Gentry 2817 (MO).
CANAL ZONE: Barro Colorado Island, Croat 4006A, 5234, 5588, 6660, 7876, 7996A, 8133,
8247, 9147, 11083 (all MO); Gentry 432 (MO). Pipeline road, Gentry 1436 (MO), Gentry
& Dressler 1977 (MO). Fort Sherman, Gentry & Tyson 4840, 4849 (both MO). DARIEN:
Vicinity of La Palma, Gentry 3908, 4288 (both MO). Isla Lagarto in Golfo de San Miguel,
Gentry 3972 (MO). Isla Boca Grande in Golfo de San Miguel, Gentry 4032 (MO). S of
Jaque, Gentry 4097 (MO). Ridge from top of Cerro Pavarando to Cerro Venado and Cerro
Derumba, Gentry 4213 (MO). Below mouth of Rio Pucuro on Rio Tuira, Gentry 4326 (MO).
Between Rios Cube and Punusa, Gentry 4458 (MO). El Real, Gentry 4498, 4536 (both
MO). Near Cerro Pirre on Rio Parasenico, Gentry 4758 (MO). Cana and vicinity, Williams
796 (NY). LOS SANTOS: S of Macaracas, Lewis et al. 1615 (MO). PANAMA: Finca Indio
on slopes of Cerro Jefe, Gentry 2152 (MO). Rio Pasiga, Gentry 2197, 2257, 2272 (all MO).
Rio Maestra, Gentry 2213, 2237 (both MO). Rio Piragua (Tabardi), Gentry 2535 (MO).
Rio Bayano upriver from Rio Piragua, Gentry 2590 (MO). Icanti, Gentry 2607 (MO). E
of Cafiitas, Gentry 3695 (MO). Near Rio Espave, Gentry 3708 (MO). Ca. 16 km from Rio
Bayano crossing on trail to Santa Fe, Gentry 3812 (MO). San Jose Island, Johnston 1346
(GH, MO), 1400, 1412 (both GH). Along road between Empire and Chorrera, Killip 3403
(US). SAN BLAS: Above Puerto Obaldia, Gentry 1475, 1527 (both MO).
29. PODRANEA
Podranea Sprague in Thiselton-Dyer, Fl. Cap. 4(2): 449. 1904. TYPE: P.

ricolasiana (Tanfani) Sprague.
Vines or climbing shrubs; branchlets terete without interpetiolar glandular

fields; pseudostipules lacking. Leaves imparipinnately compound, without ten-
drils. Inflorescence a terminal panicle. Flowers with the calyx large, cam-
panulate, membranaceous, strongly 5-toothed, pale lavender when fresh; corolla
lavender with magenta markings, tubular-campanulate, glabrous outside; anthers
glabrous, the thecae straight, divaricate; ovary linear, tetragonal, glabrous, the
ovules ca. 6-seriate in each locule. Fruit a linear, leathery capsule dehiscing
perpendicular to the septum.
One or two species of east tropical and southern Africa.
1. Podranea ricolasiana (Tanfani) Sprague in Thiselton-Dyer, Fl. Cap. 4(2):

450. 1904.-FIG. 29.
Tecoma ricolasiana Tanfani, Bull. Soc. Tosc. Ortic. 1887: 17. 1887. TYPE: (not seen).
Pandorea ricolasiana (Tanfani) Baill., Hist. P1. 10: 40. 1888.
Vine or subscandent shrub. Leaves imparipinnately compound, usually 7-9
foliolate, without tendrils, the leaflets more or less ovate, obtuse to short-
acuminate, serrate, truncate to cuneate or somewhat attenuate, 2.5-3.8 cm long
and. 1.5-2.0 cm wide, petiolulate. Inflorescence a terminal panicle. Flowers
with the calyx large, cupular-campanulate, 1.5-2 cm long, strongly 5-dentate
with apiculate lobes, pale lavender when fresh; corolla pale lavender or pinkish
with a magenta patch at the base of the 2 adaxial lobes, the tube white with
deep magenta lines inside and some pinkish splotches near the base, cam-
panulate above a short cylindric base, 6-8 cm long, with scattered lepidote
scales on the lobes, glandular-pubescent at the level of stamen insertion inside
and pubescent with kinky trichomes in the sinuses between lobes, otherwise
glabrous; stamens didynamous, the thecae divaricate, 3 mm long, the longer
filaments 1.9-2.0 cm long, shorter filaments 1.4-1.5 cm long, the staminode
3 mm long, inserted ca. 10 mm from base of corolla tube; pistil 3.2-3.4 cm long,
the ovary linear, glabrous, 5 mm long and 1.5 mm wide, the ovules 6-seriate
in each locule; disc pulvinate, 0.5 mm long and 1.5-2.0 mm wide. Capsule
(Sprague) linear and leathery.
This species is native to South Africa but is widely cultivated in tropical

regions. It is a conspicuous ornamental, flowering through most of the year
but seldom setting fruit in cultivation.
CHIRIQUI: Bambito, 1 mi. SW of Cerro Punta, Tyson 5673 (MO, SCZ).
FIGURE 29. Podranea ricolasiana (Tanfani) Sprague, habit (X 7/10). [After Gentry
473, Costa Rica (MO).]
30. PYROSTEGIA
Pyrostegia K. B. Presl, Abh. Kbnigl. Bohm. Ges. Wiss., ser. 5. 3: 523. 1845. TYPE:
P. ignea (Vell.) Presl = P. venusta (Ker.) Miers.
Lianas; branchlets subangulate with 6-8 inconspicuous ribs, without inter-

petiolar glandular fields, pseudostipules small and inconspicuous. Leaves 2-
foliolate, often with a trifid tendril. Inflorescence a terminal or axillary pani
Flowers with the calyx cupular, truncate, 5-denticulate, more or less glandular
lepidote; corolla orange-red, narrowly tubular-infundibuliform; stamens ex-
serted, the anthers glabrous; pollen grains single, 4-colpate, the exine reticulate;
ovary linear-tetragonal, lepidote; disc annular-pulvinate. Fruit compressed,
linear, the valves parallel to the septum, smooth, the median nerve somewhat
raised; seeds thin, bialate, the wings brown with hyaline margins.
Three or four species of tropical South America ranging from Brazil and
Paraguay to southern Colombia.
1. Pyrostegia venusta (Ker.) Miers, Proc. Roy. Hort. Soc. 3: 188. 1863.-FIG. 30.
Bignonia venusta Ker., Bot. Reg. 3: 5. 249. 1818. TYPE: (not seen).
B. ignea Vell., Fl. Flum. 244. 1825; 6: Tab. 15. 1827. TYPE: (not seen).
Pyrostegia ignea (Vell.) K. B. Presl, Bot. Bemerk. 93. 1843.
P. acuminata Miers, Proc. Roy. Hort. Soc. 3: 188. 1863, nomen nudum.
P. intaminata Miers, Proc. Roy. Hort. Soc. 3: 188. 1863, nomen nudum.
P. pallida Miers, Proc. Roy. Hort. Soc. 3: 188. 1863, nomen nudum.
P. parvifolia Miers, Proc. Roy. Hort. Soc. 3: 188. 1863, nomen nudum.
P. reticulata Miers, Proc. Roy. Hort. Soc. 3: 188. 1863, nomen nudum.
Bignonia tecomaeflora Rusby, Mem. Torrey Bot. Club 6: 101. 1896. SYNTYPES: Bolivia, Bang
1596 (K, GH, M, MO), Bang 1510 (GH, MO, NY, US).
Liana; branchlets somewhat angled, puberulous, without interpetiolar glan

dular fields; pseudostipules inconspicuous or oblanceolate and subfoliaceous.
Leaves 2-foliolate, sometimes with a terminal tendril; leaflets ovate, acute,
basally rounded, 4-6 cm long and 2.5-4 cm wide, chartaceous, lepidote to more
or less puberulous; tendril shortly trifid; petiolules 1-1.5 cm long, petiole 1.5-
2.5 cm long. Inflorescence a terminal panicle. Flowers with the calyx cupular,
subtruncate to 5 denticulate, scattered lepidote, the margin densely ciliate;
corolla red-orange, tubular, the lobes narrow, valvate in bud, 5.5-6.5 cm long
and 0.9-1.1 cm wide at the mouth, the tube 4.0-5.0 cm long, the lobes 1.0-1.5
cm long, the tube glabrous outside, the lobes conspicuously puberulous on
margins and to some extent within, the tube pubescent inside at and below the
level of stamen insertion; stamens exserted, the anther thecae not or very
slightly divergent, ca. 4 mm long, the filaments 3.5-4.0 cm long, the staminode
3-4 mm long and inserted ca. 3 cm from base of tube, the filaments inserted
2.2-2.4 cm from base of tube; pistil 4.8-5.1 cm long, the ovary linear, lepidote,
4 mm long and 1 mm wide, the ovules 2-seriate in each locule but staggered
so apparently 1-seriate in most cross sections; disc cupular-pulvinate. Capsule
linear, acute at both ends, 25-30 cm long and 1.4-1.6 cm wide, the midrib
slightly raised; seeds thin, bialate, 1.2-1.4 cm long and 4-4.5 cm wide, the
wings brown with a hyaline tip.
FIGuRE 30. Pyrostegia venusta (Ker.) Miers, habit (X 3/5). [After Gentry 472, Costa
Rica (MO).]
No collections have been seen from Panama, and the above description is
taken from South American material. The species is widely cultivated in the
neotropics, especially at middle elevations. I have seen it in Costa Rica, and
it undoubtedly is cultivated in Chiriqui Province as well. It is native at least
to Brazil, Paraguay, and Bolivia, but its natural range is obscured by extensive
cultivation and sometimes naturalization. The conspicuous orange-red flowers
with narrow corollas and exserted anthers are distinctive.
31. SARITAEA
Saritaea Dugand, Caldasia 3: 262. 1945. TYPE: Arrabidaea magnifica Sprague

ex van Steenis or Bignonia magnifica W. Bull.
Lianas; branchlets terete without interpetiolar glands; pseudostipules con-

spicuous, foliaceous. Leaves 2-foliolate, often with a simple tendril. Inflores-
cence a few-flowered usually terminal corymbose panicle. Flowers with the
calyx cupular, truncate, lepidote; corolla magenta, tubular-campanulate, glab-
rous or minutely glandular outside; anthers glabrous, the thecae straight, divari-
cate; ovary linear-cylindric, lepidote, the ovules 2-seriate in each locule; disc
small, cupular-pulvinate. Capsule compressed, linear, the valves parallel to
the septum; seeds thin, bialate, the wings hyaline-membranaceous.
One species of Colombia and Ecuador.
1. Saritaea magnifica (Sprague ex van Steenis) Dugand, Caldasia 3: 263.

1945.-FIG. 31.
Arrabidaea magnifica Sprague ex van Steenis, Recueil Trav. Bot. Neerl. 24: 830. 1927.
TYPE: Colombia, Santa Marta, Smith 741 (BM, K, US).
Liana; twigs subterete, flattened at nodes, lepidote, longitudinally finely

striate, drying brown to gray, the nodes without interpetiolar glandular fields;
pseudostipules foliaceous, conspicuous, 0.6-4.2 cm long and 0.4-2.0 cm wide.
Leaves 2-foliolate, sometimes with a tendril or tendril scar; leaflets obovate,
obtuse, narrowly cuneate to decurrent, 5.2-11.5 cm long and 3.1-6.4 cm wide,
chartaceous, secondary veins 3-4 on a side, the lower pair almost straight and
making a sharper angle with midvein, lepidote above and beneath with con-
spicuous glandular fields in the axils of the basal nerve pair beneath, drying
olive green to olive brown above and beneath; tendril simple, 10-14 cm long;
petiolules 0.3-1.6 cm long, petiole 1.7-2.8 cm long, lepidote as the petiolules.
Inflorescence a few-flowered cymose panicle, terminal or axillary, its rachis
and pedicels lepidote. Flowers sweetly aromatic; calyx cupular, truncate, 7-8
mmn long and 6-7 mm wide, lepidote; corolla magenta, the throat white inside
with magenta nectar guides, tubular-campanulate, 7.9-9.1 cm long and 1.6-2.0
cm wide at the mouth, the tube 5.6-6.3 cm long, the lobes 2.2-3.1 cm long,
glabrous outside, pubescent inside at the level of stamen insertion with several-
celled, apiculate trichomes; stamens didynamous, the anther thecae divaricate,
5.5-6 mm long, the longer filaments 2.7-2.9 cm long, shorter filaments 2.0-2.1
cm long, the staminode 4-6 mm long, inserted 9-12 mm from the base of the
corolla tube; pistil 4.3-4.4 cm long, the ovary linear, 4 mm long and 1 mm
wide, glandular-lepidote, the ovules 2-seriate in each locule but staggered so
often appearing uniseriate in cross sections; disc cupular-pulvinate, 1 mm long
and 2 mm wide. Capsule linear, compressed; seeds thin, bialate, the wings
hyaline-membranaceous.
Native to Colombia but cultivated through much of the tropics, this species
is a plant native to the New World which first became known to science as a
cultivar in the Old World. For some time its origin remained a mystery, but
it has since been found growing wild in Colombia and Ecuador.
The large, glabrous corolla, lack of interpetiolar glandular fields, and
-J ~ ~ 1
N.~~~~~~'
FiGuRE 31. Saritaea magnifica (Spr. ex v. St.) Dugand, habit (x 2/3), [After Gentry
6049 (MO).]
conspicuous pseudostipules indicate that its relationship may not be so much

with Arrabidaea, near which it has been placed, as with Cydista and perhaps
Phryganocydia. It has the simple tendrils and corolla form of Cydista and
Phryganocydia, both of which tend to develop foliaceous pseudostipules. Its
fruit is somewhat intermediate between that of Arrabidaea and that of Cydista
and Phryganocydia. Saritaea has most of the characteristics one would expect
of a predecessor to Cydista and, except for the calyx, to Phryganocydia. I suspect
that it represents a link between these genera, which differ from it chiefly in
lacking a nectiferous disc, and the complex of genera centered around Arrabidaea.
The cuneate, few-veined leaflets and conspicuous pseudostipules make the
sterile plant easy to recognize.
A E
G H
J~~~~~~~~~~~~~
FIGURE 32. Schiegehia.A E. S. rncaraguenszs Standi-A. Ovary cross section ( X 8).

-B. Pistil and disc ( X 1).-C. Corolla (x 1 /2)-D. Anther ( X 31/2). [After Gentry 6545
(MO).]-E. Fruit (X 1/2). [After Lewis et at. 5248 (MO).] F J. S. fuscata A. Gentry.-
F. Habit (x X1/2) CG Ovary cross section ( X 6).-H. Flower (x X'/2). I. Pistil ( X1/2).-
J. Anther (x 5 ) [After Gentry 2875 (MO).]-K-0 S. dresslert A. Gentry.-K. Leaves
CANAL ZONE: Ancon Hill across from Governor's House, Croat 15026 (MO); Gentry
1474B (MO), 6049 (MO).
32. SCHEGELIA
Schlegelia Miq., Bot. Zeitung (Berlin) 2: 785. 1844. TYPE: S. lilacina Miq. = S.
violacea (Aubl.) Griseb.
Dermatocalyx Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjbenhavn 1855: 29. 1856.
TYPE: D. parviflorus Oerst. = Schlegelia parviflora (Oerst.) Monachino.
Vines or scandent shrubs, sometimes epiphytic; stems without anomalous

vasculature; branchlets more or less terete without interpetiolar glandular fields;
pseudostipules small, blunt-conical or subulate, more or less appressed against
the branchlet. Leaves simple, opposite, often strikingly coriaceous. Inflores-
cence an axillary raceme or a terminal or axillary (usually narrow), panicle,
often conspicuously bracteate. Flowers with the calyx cupular, truncate or
irregularly lobed; corolla white to pink, purple or red (or ?yellowish), tubular-
campanulate to narrowly tubular, often short, more or less glabrous outside,
conspicuously glandular lepidote on lobes inside; anthers glabrous, the thecae
relatively short and thick, divaricate; pollen grains 3-colpate, the exine finely
reticulate; ovary globose, bilocular with a single central placenta in each locule,
the ovules multiseriate on each placenta, sometimes apically unilocular, the ovules
then parietal on 2 bifid placentae fused lower in ovary. Fruit usually a spherical
berry with a thin crustaceous pericarp, the lower part sheathed by the persistent
calyx, more or less ellipsoid and hard-shelled in some species; seeds small, angu-
lar, wingless, surrounded by pulp.
About a dozen species (after the recent segregation of the genus Gibson-
iothamnus) of tropical America ranging from Guatemala and the West Indies
to Brazil.
Schlegelia is a confusing genus intermediate between the Bignoniaceae and
Scrophulariaceae. I have accepted its traditional placement in the Bignoniaceae
with some reservation. Williams (1970) recently described Gibsoniothamnus,
a new genus of Scrophulariaceae which he interprets correctly as an additional
link between that family and Schlegelia, and also proposed transfer of Schlegelia
itself to Scrophulariaceae. The discovery of S. dressleri, the most bignoniaceous-
looking species of Schlegelia yet described, provides an additional link between
this genus and Bignoniaceae. Schlegelia species fall into two groups which
may prove to be separate genera, one in the Bignoniaceae and one in the Scro-
phulariaceae. One group, closely allied with Bignoniaceae, has relatively large
flowers and, in the two species where fruit is known, a hard-shelled, Crescentia-
like fruit. The second group, more closely related to Scrophulariaceae, has
species with small flowers and baccate fruits with only a crustaceous pericarp.
(X 3/10). [After Dressler & Maas s.n. (MO).]-L. Inflorescence (X 1/2).-M. Corolla split
open ( X 1/2).-N. Ovary cross section near apex ( X 21/2). -O. Ovary cross section near
middle ( X 21/2). [After Dressler 3507 (MO).]
One other species may occur in Panama. Schlegelia darienensis Seem. has
been collected in the adjacent Choco of Colombia. It is recognized by its
terminal inflorescence with leaf-like bracts.
Useful reference:
Williams, L. 0. An overlooked genus of the Scrophulariaceae. Fieldiana,

Bot. 32: 211-214. 1970.
a. Corolla large (more than 3.9 cm long and 1.1 cm wide at the mouth of the tube),
tubular-campanulate; fruit ca. 4 cm in diameter (where known).
b. Inflorescence a densely contracted, many-flowered raceme subtended by a con-
spicuous fascicle of basal bracts; leaves coriaceous, somewhat bullate, more than
30 cm long --- -------- 1. S. dressleri
bb. Inflorescence a few-flowered axillary fascicle without a conspicuous fascicle of basal
bracts; leaves chartaceous to subcoriaceous, not bullate, less than 12 cm long
--- - - ----- ------ ----- 3. S. nricaraguensis
aa. Corolla small (less than 2.5 cm long and 0.5 cm wide at the mouth of the tube), tubular
or shortly campanulate; fruit less than 1.5 cm in diameter.
c. Corolla narrowly tubular with minute (1-2 mm long) non-reflexed lobes, more than
2 cm long; inflorescence a congested, densely branched, contracted, fastigiate pani-
cle, each branchlet short, thick, and conspicuously jointed -5-- - 5. S. sulfurea
cc. Corolla shortly tubular with larger (3-6 mmn long) reflexed lobes, less than 1.3 cm
long; inflorescence less-branched, not fastigiate, its branchlets not conspicuously
thickened nor jointed.
d. Inflorescence an open, unbranched raceme, sometimes a short, 3-flowered side
branch replacing the lowest flower on each side; leaves drying a dark grayish
brown --- ---- - 2. S. fuscata
dd. Inflorescence a more or less contracted panicle, sometimes almost fasciculate;
most leaves drying olive to olive brown ------ 4. S. parviflora
1. Schlegelia dressleri A. Gentry.6 TYPE: Panama, La Eneida near Cerro Jefe,

Dressler 3507 (MO, holotype; PMA, isotype) .-FIG. 32K-O.
Vine, the stem without phloem arms in cross section; branchlets terete,
rough-barked, lenticellate, without interpetiolar ridges or glandular fields.
Leaves opposite or ternate, simple, elliptic,'rounded at both ends, 30-45 cm
long and 11-20 cm wide, extremely coriaceous, bullate, secondary veins 6-7
on a side, the main veins impressed above, conspicuously raised beneath, densely
minute lepidote above and beneath, otherwise glabrous, with plate-shaped
glands near base of the midvein beneath, drying olive, the petiole 2.4-3 cm
long, lepidote, rough-barked. Inflorescence to 20-flowered, a contracted raceme
2-2.5 cm long subtended by a fascicle of subulate basal bracts, borne on older
stems, the rachis and pedicels more or less lepidote and puberulous, each
pedicel subtended by a light-drying subulate bract, 2-5 mm long, the pedicels
1.2-2.2 cm long. Flowers with the calyx cupular, truncate, 9-14 mm long and
6 Schlegelia dressleri A. Gentry, sp. nov. Frutex scandens, ramulis subteretibus cortic
suberoso, sine consociebus glandularum in nodis inter petioles; folia opposita vel ternata, ellip-
tica, amplissima, percoriacea, bullata; inflorescentia floribus in racemis bracteatis contractis,
fasciculo bractearum basalium subtenta; calyx cupulatus, truncatus, glaber; corolla albida,
tubulo-campanulata; stamina didynama, thecis divergentibus; ovarium globosum, placentis
duabus parietalibus apice, supra medium connatis, ovulis in utroque locule multiseriatis;
fructus ignotus.
9.0-13 mm wide, drying dark brown, glabrous with scattered plate-shaped

glands; corolla white, tubular-campanulate, 5.4-6.2 cm long and 1.4-1.8 cm
wide at the mouth, the tube 4.3-4.8 cm long, the lobes 0.8-1.1 cm long,
minutely lepidote outside near the base, otherwise glabrous, inside the lobes
glabrous apically, becoming pubescent near the base with glandular lepidote
trichomes merging into the glandular-puberulous trichomes of the upper third
of the tube, lower part of the tube glabrous except for simple, multicellular,
gland-tipped trichomes at the level of stamen insertion; stamens didynamous,
the anther thecae divergent, 3 mm long and 1.5-2.0 mm wide, the longer fila-
ments 2.6-2.7 cm long, shorter filaments 2.0-2.1 cm long, the staminode 5 mm
long, inserted 18-19 mm from the base of the tube; pistil 0.9 cm long, the
ovary globose, 6 mm long and 6-7 mm wide, inconspicuously puberulous, the
ovules multiseriate on 2 stalked parietal placentae in the upper 1 mm of ovary,
these fused below so placentation is mostly axile with a single parietal placenta
in each locule. Fruit unknown.
Apparently endemic to the Cordillera de San Blas of eastern Panama, S.

dressleri has been collected only east of Cerro Jefe, Panama Province. This
striking plant is the most distinctive species of the genus; its giant, leathery,
somewhat bullate leaves are unmistakable. Its corolla is also the largest in
Schlegelia. To date it is known only from two vines in a region fast being
cleared by campesino squatters. It was in flower in March (1973) and late
May (1.968).
PANAMA: La Eneida, region of Cerro Jefe, Dressler 3507 (MO, PMA). Campo Tres, 3
mi. NE of Altos de Pacora, Liesner 565 (MO).
2.- Schlegelia fuscata A. Gentry.7 TYPE: Panama, Cerro Jefe, Gentry 2875
(MO).-FIG. 32F-J.
Vine ("epiphyte," Tyson et al. 3195) to at least 2 cm in diameter, the bark

dark, indistinctly ridged; stem without phloem arms in cross section; branchlets
terete,. scattered lepidote, without interpetiolar ridges or glandular fields. Leaves
opposite, simple, elliptic, apically rounded, basally truncate to wide-cuneate,
7.6-12.6 cm long and 4.2-9.5 cm wide, extremely coriaceous, secondary veins
4-5 on a side, the main veins impressed above, raised beneath, lepidote above
and beneath, otherwise glabrous, the lepidote trichomes glandular and con-
gested at the base of blade beneath, this glandular area usually black from
infection by a member of the Melanconiales (fungus), when fresh glossy green
above and dull pale green beneath, drying brown above and beneath, the
petiole 1.3-2.5 cm long. Inflorescence a 5-12-flowered axillary raceme or a
subpanicle with the basal pedicels of the raceme forked and bearing 2 flowiers
7Schlegelia fuscata A. Gentry, sp. nov. Frutex scandens; folia opposita, elliptica, per-
coriacea, in sicco fusca; inflorescentia floribus 6-12, in racemis laxis vel paniculiformis
pedicellis basilaribus furcatis ferentibus duos vel tres flores, bracteis minimis inconspicuis;
calyx cupulatus, plus minusve vadose lobatus, glabrus vel leviter lepidotus, margine pubescenti;
corolla tubo albo, lobis roseis, tubulosa lobis refractis; stamina didynama, thecis non divergen-
tibus; ovarium globosum, ovulis multiseriatis, in placenta singulari ad centrum utriusque loculi;
fructus globosus, ad 1.2 cm diam., calyce persistenti.
the branches simple-pubescent, each pedicel pair and each flower subtended
by a pair of acute bracts with ciliate tips, the rachis 1.2-5 cm long, pedicels
4-8 mm long. Flowers with the calyx cupular, widest at the middle, with 4-5
round-tipped lobes, 6-7 mm long and 1-6 mm wide, glistening white with a
slight pink tint when fresh, black when dry, inconspicuously lepidote, simple-
pubescent at the tips; corolla greenish-white on the tube outside, the lobes
pink, paler at the tips and deeper at the base shading into the purplish-pink
of the throat, tubular with reflexed lobes, 1.0-1.1 cm long and 4 mm wide at
the mouth, the tube 7 mm long, the lobes 3-4 mm long, the posterior pair
sometimes partially fused, glabrous outside, the lobes glandular-lepidote to
glandular-puberulent inside, the tube glabrous inside except for long simple
multicellular glandular trichomes at the level of stamen insertion; stamens
didynamous, the anther thecae not divergent, 1.5-2 mm long, the filaments
all ca. 0.8 cm long but the posterior pair bent more sharply holding the posteri
anthers lower in the throat, the staminode 3 mm long, inserted 4-5 mm from
the base of the tube; pistil 5 mm long, the ovary spherical, 2-2.5 mm long and
2.5-3.0 mm wide, glabrous, the ovules 8-9-seriate on a single central placenta
in each locule; disc absent. Fruit a spherical fleshy berry to 1.2 cm in diameter,
pale purple when fresh, the calyx persistent and covering the lower half to
third of the fruit, glabrous but large-papillate; seeds angular, papillate-surfaced,
2-3 mm, long, less than 1 mm wide.
This species is only known from the Cerro Jefe and Santa Rita Ridge regions
of central Panama in the tropical wet forest and premontane rain forest at an
elevation of 800 to 1000 m, where it has been collected a number of times.
Schlegelia fuscata is closest to S. parviflora, differing most conspicuously in its
open, more or less racemose inflorescence which is unbranched or with a single
short, basal, 3-flowered side branch on each side and in its uniformly dark-
drying, extremely coriaceous leaves. The leaves are always conspicuously black-
ened below in the glandular area at the base of blade, presumably from a
fungal infection.
'This species may be no more than a local form of the heterogeneous S.
parviflora, but on the basis of present evidence it is different enough to be
considered specifically distinct.
COLON: Santa Rita Ridge, Dressler 3618 (PMA). PANAMA: Cerro Jefe summit, Dwyer
et al. 7276 (MO); Gentry 2875 (MO), 3692 (MO); Tyson et al. 3195, 4363, 5304 (all SCZ).
3. Schlegelia nicaraguensis Standl., Trop. Woods 16: 44. 1928. TYPE: Nica-
ragua, Bluefields, Englesing 99 (F, GH, K). -FIG. 32A-E.
Woody vine, stem without phloem arms in cross section; branchlets lepido
sparsely pubescent at least at the nodes, without interpetiolar ridges or glandu-
lar fields, pseudostipules subulate, to 3 mm long, pubescent. Leaves opposite,
simple, elliptic-obovate, apically rounded or apiculate, basally wide cuneate,
usually abruptly short-attenuate, 7.6-11 cm long and 3.8-7.4 cm wide, char-
taceous to rigid-chartaceous, the secondary veins 5-8 on a side, raised above,
the midvein impressed above, all main veins raised beneath, the midvein
conspicuously so, inconspicuously lepidote (often deciduously) above and be-

neath, sparsely puberulous along main veins beneath, otherwise glabrous, plate-
shaped glands scattered over the leaf surface beneath, congested at the base
of the blade, each gland surrounded by a raised epidermal papilla, drying brown
to gray brown, petiole 0.8-1.6 cm long, slightly lepidote, flat-topped, incon-
spicuously puberulous along the angles. Inflorescence a few-flowered, axillary
fascicle on older branchlets below the leaves, the pedicels puberulous. Flowers
with the calyx cupular, truncate to slightly lobed, 1 cm long and 1 cm wide,
when fresh violet with white spots, dark brown when dry, lepidote, puberulous
mostly near rim and base with immersed plate-shaped glands mostly toward
the apex; corolla cream or yellowish at base, the lobes magenta, tubular-
3.1-3.7 cm long, the lobes 0.5-0.8 cm long, glabrous outside and at the tips
inside, lepidote near the middle and base, the pubescence merging into the
simple multicellular trichomes of the upper third of the tube, lower part of
tube glabrous except for simple gland-tipped trichomes at the level of stamen
insertion; stamens didynamous, the anther thecae thick, slightly divergent, 3
mm long and 1.5-2.0 mm wide, longer filaments 2.0-2.2 cm long, shorter fila-
ments 1.5-1.7 cm long, staminode 3-5 mm long, inserted 5 mm from base of
corolla tube; pistil 0.5-0.7 cm long, the ovary truncately spherical, glabrous,
3 mm long and 3 mm wide, the ovules multiseriate on 2 bifid, stalked, parietal
placentae in upper 2/3 of ovary, these fused in the lower third and placentation
axile with a single central placenta in each locule; disc thin, sunken into the
base of the ovary. Fruit spherical, hard-shelled, glabrous, ca. 4 cm in diameter,
subtended by the persistent calyx, the seeds small, angular, 2-3 mm long and
1 mm wide, borne on an incompletely fused septum.
Previously known only from Nicaragua and Guatemala, the Panamanian

collections from the Cerro Jefe and Santa Rita regions of eastern Panama extend
the range of this species the length of Central America. It is evidently restricted
to the~ poorly known tropical wet forest. The species has been collected in
flower in December, January, and June in Panama, indicating an irregular
flowering season.
Schlegelia nicaraguensis is distinguishable from most species of the genus
by its large (ca. 4 cm long) corolla. It differs from S. dressleri most strikingly
in its much smaller leaves. This species is closely related to S. parasitica of the
West Indies and might prove to be a variety of that species.
BOCAS DEL TORO: Vicinity of Chiriqui Lagoon, von Wedel 1051 (GGH, MO). COLON:
Santa Rita Ridge, Croat 13201 (MO, SCZ), 14188 (MO); Gentry 6545 (MO); Lewis et al.
5248 (MO). PANAMA: Altos del Rio Pacora, Lewis et al. 2319 (MO, SCZ).
4. Schlegelia parviflora (Oerst.) Monachino, Phytologia 3: 103. 1949.
Dermatocalyx parviflorus Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kj0benhavn 1855:
28: 1856. TYPE: Costa Rica, Alajuela, Oersted s.n. (not seen; illustration in type
herbarium at C).
Citharexylum ramizii Glaziou, Bull. Soc. Bot. France 58, Mem. 3: 545. 1911, nomen nudum.
Tabebuia micrantha Kranzl., Fedde Repert. 17: 215. 1921. TYPE: Brazil, Sdo Paulo, Mosen
3662 (not seen; K, photo).
Schlegelia ramizii Sandw., Kew Bull. 1930: 211. 1930. TYPE: Brazil, Rio de Janeiro, Glaziou
15327 (C, K).
Aegiphila congesta Standl. ex Moldenke, Brittonia 1: 471. 1934, nomen nudum.
Schlegelia ramizii var. costaricensis Sandw., Kew Bull. 1940: 303. 1941. TYPE: Costa Rica,
San Jose, Skutch 4125 (A, K, MO).
S. ramizii var. macrandra Sandw., Kew Bull. 1940: 303. 1941. TYPE: Peru, Loreto, Klug
2340 (K).
S. ramizii var. trichandra Sandw., Kew Bull. 1940: 304. 1941. TYPE: British Honduras,
Peck 666 (K).
S. roseiflora Ducke, Trop. Woods 76: 29. 1943. TYPE: Brazil, Amazonas, Ducke 1142 (MO,
NY, US).
S. organensis Kuhlm., Rodriguesia 20: 7, pl. 1. 1946. TYPE: Brazil, Rio de Janeiro, Kuhlmann
s.n. (not seen).
S. macrophylla Ducke, Trop. Woods 90: 29. 1946. TYPE: Brazil, Amazonas, Ducke 1854 (K,
NY, US).
S. gentlei Lundell, Wrightia 4: 95. 1968. TYPE: Guatemala, Alta Verapaz, Contreras 4471
(US).
Woody vine; stem without phloem arms in cross section; branchlets terete,
glabrous, without interpetiolar ridges or glandular fields. Leaves opposite,
simple, obovate to elliptic-obovate, apically rounded to acute, basally cuneate,
11.2-19.5 cm long and 4.8-14.7 cm wide, coriaceous, secondary veins 6-8 on a
side, minutely lepidote at least beneath, otherwise glabrous with a glandular
field near the base of the blade below, drying olive to olive brown, the petiole
1.0-2.4 cm long. Inflorescence a few- to many-flowered axillary, more or less
contracted, sometimes almost fasciculate panicle, the branches puberulous with
small bracts, 1-2 mm long, subtending many of its dichotomies, these more
or less short-puberulous at least along the margins. Flowers with the calyx
cupular with 2-3 round-tipped lobes, 4-6 mm long and 3-5 mm wide, incon-
spicuously lepidote or subpuberulous at least at the base, usually drying tan;
corolla white or cream flushed with pink, tubular with reflexed lobes, 1.0-1.2
cm long and 0.4 cm wide at the mouth, the tube 0.6-0.8 cm long, the lobes
0.4-0.6 cm long, glabrous outside, the lobes glandular-lepidote to glandular-
puberulous inside, the tube scattered pubescent in the upper half inside, glab-
rous below the level of stamen insertion; stamens more or less equal, the
anther thecae thick, slightly divergent, 1 mm long and almost 1 mm wide, the
filaments 4 mm long, the bases swollen and pubescent, the staminode 2-3 mm
long, inserted 4 mm from the base of the corolla tube; pistil 3-4 mm long, the
ovary spherical, 1.5-2 mm long and 2 mm wide, glabrous, the ovules multi-
(ca. 8-)seriate on a central placenta in each locule; disc absent. Fruit a spher-
ical, fleshy berry, 0.9-1.1 cm in diameter, glabrous, the lower part covered by
the persistent calyx; seeds angular, ca. 2 mm long, less than 1 mm wide.
One of the most-collected species of the genus, S. parviflora occurs in the
tropical wet forest and premontane wet forest from Guatemala and British
Honduras to Brazil. In addition to the listed synonyms, a number of manu-
script names have been used for various collections of S. parviflora.
The species is characterized vegetatively by its stiff coriaceous leaves, much
smaller than those of S. dressleri. They differ from the round-tipped leaves of
S. fuscata in being usually more or less subacuminate and from the elliptic-
oblong leaves of S. sulfurea in being more or less obovate. In flower or fruit

S. parviflora differs from the likewise small-flowered S. fuscata by its fasciculate
or paniculate inflorescence as compared with the open, few-branched racemose
or subpaniculate inflorescence of the latter.
Schlegelia parviflora is extremely' variable as here interpreted, but under-
standing of its variation has so far proved difficult. It seems best to follow
Sandwith (in herb.) in using the name in a relatively wide sense until a thorough
revision of the genus is possible.
A confusing collection is von Hagen & von Hagen 2122 (NY) from Chiriqui,
Panama, which is intermediate between S. monachinoi Moldenke and S. parvi-
flora. It was determined by Moldenke as S. monachinoi of Colombia, a taxon
of uncertain status.
BOCAS DEL TORO: Rio Teribe between Quebrada Treglo and Puerto Palenque, Kirkbride
& Duke 538 (MO, SCZ). COLON: Santa Rita Ridge, Correa & Dressler 747 (MO, PMA,
SCZ). DARIEN: Camp Summit, Duke 15486 (MO). PANAMA: Along road from Cerro Azul
to Cerro Jefe, Tyson 5304 (SCZ). WITHOUT LOCALITY: Seemann 66 (BM).
5. Schlegelia sulfurea Diels, Notizbl. Bot. Gart. Berlin 14: 39. 1938. TYPE:
Ecuador, Schultze-Rhonhof 1927 (not seen).
S. fastigiata Schery, Ann. Missouri Bot. Gard. 29: 367. 1942. TYPE: Panama, Bocas del Toro,
von Wedel 773 (GH, MO, US).
Woody vine; stem without phloem arms in cross section; branchlets terete,
glabrous or inconspicuously short puberulous, without interpetiolar ridges or
glandular fields; pseudostipules lanceolate, acute, paired, appressed against the
branchlet. Leaves opposite, simple, elliptic-lanceolate to narrowly elliptic, acute
to 'short-acuminate, basally broadly cuneate to truncate, 10-33 cm long and
4-13.2 cm wide, chartaceous, to rigid-chartaceous, secondary veins 7-12 on a
side, veins slightly raised above, conspicuously so beneath, essentially glabrous
above and beneath with some plate-shaped glands beneath, these concentrated
at the base of the blade, drying dark olive beneath,. blackish above, the petiole
usually thick, often woody, slightly winged apically from the more or less
attenuate leaf blade, flat or shallowly grooved above, convex beneath, 0.5-1.5
cm long. Inflorescence a congested, extremely densely branched, contracted
panicle, the branchlets short and Conspicuously jointed, woody, usually shortly
puberulous. Flowers with the calyx cupular, truncate, 5-8 mm long and 5-7
mm wide, glabrous to slightly lepidote or puberulous, pink when fresh, drying
dark red; corolla creamy white with pink tip (Porter et al. 4432), narrow-tubular,
the lobes small and not reflexed, 2.0-2.5 cm long and 0.3-0.4 cm wide at the
mouth of tube, the tube 1.9-2.4 cm long, the lobes 1-2 mm long, the tube
glabrous inside and out, the lobes slightly glandular-lepidote outside, the anther
thecae slightly divergent, each 1-1.5 mm long and ca. 1 mm thick, the longer
filaments 0.9-1.0 cm long, shorter filaments 0.8-0.9 cm long, puberulous at the
base, the staminode 3 mm long, inserted 10-12 mm from the base of the tube;
pistil 4 cm long, the ovary spherical, lepidote, 2 mm long and 2 mm wide,
the ovules multi-seriate in each locule on a single central placenta; disc cupular-
pulvinate, somewhat sunken into the base of the ovary, 1 mm long and 2.5 mm
wide. Fruit a berry, spherical, 1.0-1.4 cm long and 1.3-1.5 cm wide, its lower
quarter covered by the expanded persistent calyx, glabrous.
This is a plant of the tropical wet forest. It ranges from Guatemala to

Ecuador at altitudes from sea level to almost 1000 m.
The type of S. sulfurea was destroyed at Berlin, but other specimens from
the type locality in Ecuador are clearly the same as S. fastigiata. I have seen
a duplicate of the Ruiz and Pavon collection which was mentioned in the orig-
inal description of S. sulfurea and it, too, is conspecific with the plant formerly
known as S. fastigiata.
The relatively long, narrow leaves and subulate-appressed pseudostipules are
good recognition characters, and the inflorescence is distinctive. Schlegelia
darienensis is known from just across the Colombian border and might be
expected to occur in Panama. It has somewhat similar fastigiate branches in
its inflorescence but this inflorescence differs in being terminal, elongate, and
having conspicuous leaf-like bracts.
BOCAS DEL TORO: Region of Almirante, Cooper 167 (F, NY, US), 169, 169A (both F).
Cerro Bonyic above Quebrada Huron, Kirkbride & Duke 600 (MO). Isla Colon, von Wedel
44 (CH, MO). Vicinity of Chiriqui Lagoon, von Wedel 154 (MO, US), 1397 (GH, MO,
US). Water Valley, von Wedel 773, 1447 (both GH, MO, US), 1468 (GH, MO), 1659
(GH, MO, US). Old Bank Island, von Wedel 2126 (GH, MO, NY, US). Fish Creek moun-
tains, von Wedel 2361 (MO). Fish Creek lowlands, von Wedel 2375 (MO). Fish Creek
Hills, von Wedel 2419 (GH, MO, US). COCLE: Summit of Cerro Pilon above El Valle de
Anton, Dwyer et al. 4503 (MO, SCZ); Dwyer & Lallathin 8691 (MO). 2 mi. N of Cerro
Pilon, Liesner 729 (MO). Foot of Cerro Pilon, Porter et al. 4432 (MO). DARIEN: Along
Sea Level Canal Route 17, Oliver et al. 3707 (MO); Duke 15491 (MO). PANAMA: N of El
Llano, Gentry 5086 (MO). SAN BLAS: Headwaters of Rio Cuadi (drill site 22), Duke et al.
3635 (MO). VERAGUAS: Along Rio Primero Brazo ca. 5 mi. NW of Santa Fe, Croat 23127
(MO); Liesner 1000 (MO).
33. SCOBINARIA
Scobinaria Seib., Carnegie Inst. Washington .Publ. 527: 408. 1940. TYPE: S.
verrucosa (Standl.) Seib. = S. japurensis (DC.) Sandw.
interpetiolar glandular fields, these usually sharply limited above by a more
or less V-shaped line connecting opposite petioles; pseudostipules small and
inconspicuous. Leaves 2-foliolate, often with a simple tendril. Inflorescence
a few-flowered axillary or terminal panicle. Flotvers with the calyx large and
thin, campanulate, bilabiate, variously lepidote at least at the base; corolla
lavender to magenta, tubular-campanulate, puberulous outside; anthers gla-
brous, the thecae straight, divergent; pollen grains 3-colpate; ovary linear-tetrag-
onal, finely lepidote-papillate, the ovules 2-seriate in each locule. Fruit a com-
pressed linear-oblong capsule, the valves parallel to the septum, verrucose
tuberculate, drying black, the median nerve not visible; the seeds thin, bialate,
the wings mostly brown, hyaline at the extreme margin, not demarcated from
the seed body.
Probably with only one species, the genus is often included with Arrabidaea.
It occurs in continental tropical America, ranging from southern Mexico to
Amazonia.
1. Scobinaria japurensis (DC.) Sandw., Kew Bull. 13: 440. 1959.-FIG. 33.
Adenocalymma japurense Mart. ex DC., Prodr. 9: 203. 1845, pro syn.

Phryganocydia japurensis Mart. ex DC., Prodr. 9: 214. 1845, pro syn.
Tabebuia japurensis DC., Prodr. 9: 214. 1845. TYPE: Brazil, Amazonas, Martius s.n. (M;
G-DC, fragment).
Arrabidaea japurensis (DC.) Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 65. 1896.
A. lenticellosa Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 64. 1896. TYPE: Brazil, Matto
Grosso, Riedel 1469 (K).
A. oligantha Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 64. 1896. TYPE: Brazil, Gardner
1258 (K, as Hostmann 1258 from Surinam).
Paragonia schumanniana Loes., Bot. Jahrb. Syst. 23: 130. 1897. TYPE: Nicaragua, Matagalpa,
Rothschuh 230 (not seen).
Arrabidaea nicotianaeflora Kranzl., Notizbl. Bot. Gart. Berlin 6: 369. 1915. TYPE: Brazil,
Rio Acre, Ule 9782 (K).
Adenocalymma verrucosum Standl., Publ. Field Columbian Mus., Bot. Ser. 4: 323. 1929.
TYPE: Honduras, Atlantida, Standley 54891 (A, F, US).
Arrabidaea belizensis Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 48. 1930. TYPE:
British Honduras, Middlesex, Schipp 284 (BM, F, K, MO, NY, US).
Martinella verrucosa (Standl.) Standl., Contr. Arnold Arbor. 5: 138. 1933.
Scobinaria verrucosa (Standl.) Seib., Carnegie Inst. Washington Publ. 522: 408. 1940.
S. amethystina Dugand, Caldasia 7(31): 24. 1955. TYPE: Colombia, Bolivar, Romero-
Castafieda 1735 (COL).
Liana to at least 10 cm in diameter, the bark gray and conspicuously orange-

lenticellate; stem with 4 phloem arms in cross section; branchlets variable, sub-
tetragonal to subterete, more or less longitudinally striate, glabrous to lepidote
or puberulous, brownish when fresh, the nodes normally with interpetiolar
glandular fields sharply limited above by a more or less V-shaped line con-
necting opposite petioles; pseudostipules lacking. Leaves 2-foliolate, sometimes
with a tendril; leaflets more or less ovate, acuminate to obtuse, rounded to
broadly cuneate, 5.2-17 cm long and 2.4-11.8 cm wide, chartaceous to mem-
branaceous, secondary veins (3-) 4-6 on a side, the basal pair usually not
parallel to the others, scattered lepidote especially on the veins, pubescent
beneath with long simple trichomes in leaf axils and sometimes along the base
of the main veins, above usually minutely puberulous at the base of the mid-
vein, usually drying greenish-olive with reddish veins and veinlets beneath;
tendril simple, 8-20 cm long; petiolules 1.0-4.2 cm long, petiole 1.3-5.1 cm long,
lepidote and puberulous at least above. Inflorescence a few-flowered axillary
panicle, the branches lepidote, each pedicel subtended by a bract and with 2
bracteoles, these minute, subulate, ciliate. Flowers with sweet slightly spicy
aroma, the calyx tubular-campanulate, strongly bilabiate, membranaceous, 1.8-
2.4 cm long and 0.8-1.2 cm wide, white when fresh, strongly lepidote or lepidote
only at the base; corolla lavender to magenta with a white throat, tubular-
3.2-5.7 cm long, lobes 0.7-1.8 cm long, pubescent outside with simple multi-
cellular trichomes, the lobes ciliate-fringed and pubescent, the tube mostly
glabrous inside, pubescent at the level of stamen insertion; stamens didynamous,
FIG
6267 (MO).] B. Fruit (X '/2). [After Gentry 4185 (MO)] C. Corolla split open (X '/2).
-D. Ovary and disc (X 1'/2). E. Ovary cross section (X 71/2). [After Gentry & Dwyer
5509 (MO).]
the anther thecae divaricate, 3 mm long, the longer filaments 1.7-2.0 cm long,
shorter filaments 1.1-1.4 cm long, the staminode 1-3 mm long, inserted 1.2-1.4
mm from base of corolla tube; pistil 2.5-2.8 cm long, the ovary linear-oblong,
more or less tetragonal, 3 mm long and 1-1.5 mm wide, densely lepidote,
minutely papillate, the ovules 2-seriate in each locule; disc annular pulvinate,
1 mm long and 2.5 mm wide. Capsule linear, acute to obtuse, flattened, 12-46
cm long and 1.3-2.5 cm wide, the surface black, conspicuously verrucose-
tuberculate, sometimes lepidote between tubercles, the tubercles sometimes
short and little more than papillae; seeds 1.2-1.8 cm long and 3.9-5.1 cm wide,
the wings membranaceous, brown, sometimes with the extreme margin hyaline,
not well demarcated from the seed body.
A common but infrequently collected vine of the tropical wet forest, this
species also occurs less frequently in premontane wet forest and tropical moist
forest. It ranges from British Honduras to Venezuela and Boliva. The species
flowers during the wet season from June to November. Seed release is during
the dry season.
Recognition of sterile vines in the field is by a combination of characters.
In general S. japurensis has interpetiolar glandular fields with a distinct and
often centrally notched interpetiolar ridge forming an upper margin. The bark
of mature plants is grayish with distinctive orange lenticels. The lower lateral
nerve pair of the leaflets tends to ascend arcuately in contrast to the other
lateral nerves. However, any combination of these characteristics seems possible,
especially in young plants, making this species difficult to identify when sterile.
The linear, verrucose-tuberculate fruit is unique and resembles a wood rasp
in texture. Some specimens have the tubercles much more accentuated than
others, but this is of no taxonomic significance. The flower with its flattened
tube and large, bilabiate, membranaceous calyx is also distinctive.
Sandwith (Kew Bull. 22:403-420. 1968) reunited Scobinaria with Arrabidaea,
but it is sufficiently distinct to be retained, even if it should prove monotypic.
The peculiar fruit and calyx, the small inflorescence, never 3-foliolate leaves,
much thicker stem with a different type of bark, and especially the large, rela-
tively thick-textured, dorsally compressed corolla all argue for its separation.
Most of the series Macrocalycinae of Arrabidaea as constituted by Bureau and
K. Schumann in Flora Brasiliensis is conspecific with S. japurensis. Although
I have not included A. egensis Bur. & Schum. in the synonymy of S. japurensis,
I suspect that it is probably a pubescence form; certainly it is a Scobinaria
rather than an Arrabidaea.
CANAL ZONE: Barro Colorado Island, Bailey & Bailey 609 (F); Croat 9040, 10(129 (both
MO), 13266 (MO, SCZ), 13983 (MO), 14102 (MO, SCZ), 14995, 15168 (both MO),
15564 (MO, SCZ); Foster 1071 (PMA), 1387 (MO, PMA); Gentry 709 (MO); Kenoyer
532, 535 (both US); Shattuck 438 (F, US), 903 (MO, US); Starry 151 (F, MO); Wetmore
& Abbe 56 (F); Woodworth & Vestal 324 (F). Las Cruces Trail in Madden Forest, Gentry
1385 (MO). CHIBIQUI: 12 mi. N of Puerto Armuelles, Croat 21965 (MO). 7.5 mi. W of
Puerto Armuelles, Croat 22486 (MO). COLON: Ridge W of Portobelo, Gentry 1765 (MO).
Rio Guanche, Gentry 6318 (MO). DARIEN: Rio Pifias, Duke 10562 (MO). Rio Balsas be-
tween Manene and Tusijuanda, Duke 13562 (MO). Puerto Sta. Dorothea, Dwyer 2236
(MO). S of Jaque, Gentry 4098 (MO). Rio Jaque upriver from Jaque, Gentry 4185 (MO).
FIGURE 34. Spathodea campanulata Beauv.-A. Habit (X 1/2). [After Croat 8890 and
Gentry 700B (both MO)] B. Fruit and seed (x 1/2). [After Croat s.n. (MO).]
S of La Palma, Gentry 4308 (MO). Rio Tuira between Rios Paya and Aspave6, Gentry 4427
(MO). Trail to Cerro Pirre from village of Pirre, Gentry 4779 (MO). PANAMA: Rio Pasiga,
Gentry 2193, 2312 (both MO). Between Rio Sulugandi and Rio Espave, Gentry 4990 (MO).
Junction of Rios Pacora and Corso to headwaters of Rio Corso, Oliver 2359 (MO, SCZ). SAN
BLAS: Near stream above Puerto Obaldia, Gentry 1536 (MO). Mainland opposite Playon
Chico, Gentry 6374 (MO). VERAGUAS: N of Santa Fe, Gentry 3016 (MO).
34. SPATHODEA
Spathodea Beauvois, Fl. Oware 1: 46. 1805. LECTOTYPE: S. campanulata

Beauvois.
Trees. Leaves imparipinnate, 9-10-foliolate. Inflorescence a terminal raceme,

the lower pedicels longer. Flowers with the calyx large, spathaceous, narrow-
ing to a recurved tip; corolla red-orange or crimson, usually with a yellow
fringe, large, broadly campanulate above a cylindrical base, glabrous outside;
anthers glabrous, the thecae divaricate; ovary narrowly oblong, the ovules multi-
seriate in each locule. Fruit a narrowly oblong-elliptic capsule, the valves keeled,
narrowed at the ends, dehiscing perpendicularly to the septum; seeds flat, winged,
the wings broad and hyaline-membranaceous.
A monotypic genus of tropical Africa.
1. Spathodea campanulata Beauvois, Fl. Oware 1: 47. tab. 27. 1805.-FIG. 34.
Tree to at least 25 m tall and 40 cm d.b.h. Leaves imparipinnate, usually

9-15-foliolate, the leaflets more or less elliptic, acute to acuminate, asymmetrically
subtruncate or widely cuneate basally, subsessile, slightly puberulous along main
veins. Inflorescence a terminal raceme, the lower pedicels longer and curving
upward to give a flat-topped effect. Flowers with the calyx spathaceous taper-
ing to a recurved point which bends away from the corolla, 4.5-5 cm long;
corolla red-orange with a yellow border, broadly campanulate above a cylindrical
base, large, 8.5-9.0 cm long and 4.5-5.0 cm wide at the mouth, the tube 6-6.5
cm long, the lobes 1.5-2 cm long, glabrous; stamens 4, subexserted, held against
the floor of the tube, the anther thecae divaricate, slender, 8 mm long; ovary
narrowly oblong, minutely papillate, the ovules m-ulti-seriate in each locule.
Capsule narrowly oblong-elliptic, dehiscing on one side, the valves boat-shaped,
narrowing to the ends, 17-25 cm long and 3.5-7 cm broad; seeds thin, winged,
the wings broad and hyaline-membranaceous, sharply demarcated.
A native of tropical Africa, S. campanulata is planted extensively as an

ornamental and shade tree throughout the tropics. Its bright-colored, showy
flowers are produced during most of the year and make it one of the most
attractive and best known species of Bignoniaceae. In the neotropics it often
sets fruit and at times is spontaneous though seldom naturalized. "African tulip
tree."
CANAL ZONE: Barro Colorado Island, Croat 5177, 8890 (both MO). Summit Garden,
Gentry 700B (MO). Curundu, Tyson 2042 (MO, SCZ).
35. STIZOPHYLLUM
Stizophyllum Miers, Proc. Roy. Hort. Soc. 3: 197. 1863. LECTOTYPE: S. denticu-
latum Miers = S. perforatum (Cham.) Miers.
Lianas; stems with 4 phloem arms in cross section, the extreme center mostly
hollow; branchlets terete without interpetiolar glandular fields; pseudostipules
inconspicuous or more or less spathulate and early deciduous. Leaves 3-foliolate,
conspicuously pellucid-punctate from the lepidote-peltate glands on the lower
surface, the terminal leaflet often replaced by a simple or trifid tendril. Inflo-
rescence a few-flowered axillary raceme. Flowers with the calyx campanulate,
sometimes more or less inflated, pubescent, irregularly shallowly bilabiate to
5-lobed; corolla white or cream, the lobes sometimes flushed with pink; anthers
glabrous or pubescent on the margin of the thecal openings, the thecae straight,
divaricate; pollen grains 3-colpate, the exine microreticulate; ovary linear-
tetragonal, lepidote, the ovules 2-seriate in each locule. Fruit a narrow, linear
capsule, the valves parallel to the septum, thin, convex, usually faintly striate
and more or less pubescent; seeds thin, bialate, the wings membianaceous,
brownish, weakly demarcated.
Probably three species in continental tropical America ranging from southern

Mexico to Brazil.
Miers' S. denticulatum matches in every respect except foliaceous pseudo-
stipules (which are ephemeral and inconstant in this genus) the genus described
with it and is the logical choice as lectotype. Although Bureau and Schumann
(1896) lump this species and all twelve of Miers' other species of the genus
under S. perforatum, I am not sure of the correct interpretation of this group.
Sandwith has suggested (Kew Bull. 1954: 608. 1955) that Stizophyllum might
constitute a single highly variable species, and Macbride followed this sugges-
tion in his treatment for Flora of Peru (Field Mus. Nat. Hist., Bot. Ser. 13(95):
56. 1961). I am convinced, however, that there are at least three species in
the genus, and perhaps some other variants deserve infraspecific recognition.
Sterile plants of this genus can be readily identified in the field by the
pellucid-punctate dots on their leaves (except when very young) and the con-
spicuously hollow stems. Fruits of the genus are among the narrowest in the
family and are difficult to discern from twigs at any distance. The white flower
color and slightly pubescent to glabrous anthers are also useful characters.
a. Branchlets and petiolules reddish hispid; tendrils trifid; calyx expanded-campanulate,

membranaceous; fruit villous-pubescent -1. S. inaequilaterum
aa. Branchlets and petiolules grayish or yellowish puberulous; tendrils simple; calyx short,
campanulate, chartaceous; fruit short-puberulous -2. S. riparium
1. Stizophyllum inaequilaterum Bur. & K. Schum. in Mart., Fl. Bras. 8(2):

221. 1896. TYPE: Peru: Mayna, Poeppig 1827 (W; B, destroyed) .-FIG.
35A-B.
Bignonia inaequilatera Poeppig ex Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 222. 1896, pro
syn., non Poeppig ex Bur., Addansonia 8: 289. 1868.
Cremastus rufo-villosus Herzog, Meded. Rijks-Herb. 29: 40. 1916. TYPE: Bolivia, Misiones de
Guarayus, Herzog 265, 1st journey (L; K, photo).
Liana to at least 2.5 cm in diameter, the bark smooth, somewhat scaly, stem
subtetragonal with hollow center and 4 phloem arms in cross section; branchlets
terete, hollow, longitudinally striate, strongly pubescent with long (to 1.5 mm)
FIGURE 35. Stizophyllum. A-B. S. inaequilaterum Bur. & K. Schum. A. Habit (X 1/2).
[After Gentry 7383 (MO).]-B. Fruit (x 1/2). [After Lewis et al. 5257 (MO).]-C-G. S.
riparium (H.B.K.) Sandw.-C. Anther (X 31/2 ) .-D. Ovary cross section (x 15) .-E. Corolla
(x '/2).-F. Calyx (x 1/2).-G. Pistil and disc (X 1/2). [After Gentry 2075 (MO).]
simple, usually kinky trichomes, the nodes without interpetiolar glandular fields;
pseudostipules usually inconspicuous or lacking, sometimes linear with a thick-
ened tip. Leaves 3-foliolate or 2-foliolate with tendril or tendril scar; leaflets
more or less oblong-elliptic, acuminate, asymmetrically truncate or subcordate,
5.6-13.1 cm long and 3.0-6.1 cm wide, membranaceous, sometimes inconspicu-
ously serrulate, more or less pinnately veined to the base, secondary nerves 4-7
on a side, regularly sunken-lepidote glandular below, hence pellucid punctate,
hirsute on nerves, especially beneath with long, simple trichomes, these arising
sporadically from even minute reticulations, drying dark olive to yellowish-olive,
the veins usually brownish beneath; tendril trifid, 8-16 cm long to branching,
the 3 arms 0.6-2 cm long; terminal petiolules 1.1-3.7 cm long, lateral petiolules
0.4-2.3 cm long, petiole 2.2-7.5 cm long, pubescent with long simple curly tri-
chomes. Inflorescence a long-bracteate, few-flowered terminal fascicle or a
contracted raceme terminal on a young lateral shoot, the branches hirsutely
reddish-pubescent, the bracts linear, pubescent, to 2 cm long. Flowers with the
calyx membranaceous, inflated-campanulate, 12-18 mm long and 9-10 mm wide,
the edge irregular, drying light green, slightly lepidote and sparsely hirsute,
usually with glands; corolla greenish-yellow with lavender lobes, 5-6.5 cm long
and 1-1.5 cm wide at the mouth, the tube 3.8-4.8 cm long, the lobes 0.8-1.0 cm
long, the tube glandular-lepidote and puberulous outside, the lobes pubescent,
the tube mostly glabrous inside; stamens didynamous, the anther thecae divari-
cate, sparsely ciliate with short trichomes along the margins, ca. 2 mm long,
the longer filaments 2-2.1 cm long, shorter filaments 1.5-1.6 cm long, inserted
1.3-1.4 cm from the base of the tube; pistil and ovary not seen in the 2 flowers
available for study. Capsule linear, narrow, subterete in cross section, 30-32.5
cm long and 4-5 mm wide, densely hirsute or villous.
A rarely collected vine of the tropical wet forest and premontane wet forest,
it is sometimes locally common but inconspicuous both in flower and fruit.
Flowering collections from Panama have been made in April and May.
This species is superficially similar to its congener, S. riparium, but field
study has shown the two species are readily distinguished even when sterile.
The coarser, more conspicuous, reddish-brown pubescence of twigs and leaves
of this species contrasts markedly with the dense, short, whitish-appearing
puberulence of. S. riparium. Stizophyllum inaequilaterum lacks the conspicuous
strap-shaped pseudostipules of young twigs of S. riparium, and the tendrils are
trifid, while those of S. riparium are simple; also the calyx and fruit of this
species are pubescent with much longer trichomes than those of S. riparium.
CANAL ZONE: Pipeline Road, Gentry 706, 1234, 1452, 3163 (all MO). Cultivated at
Summit Garden, Gentry 7383 (MO). COLON: Santa Rita Ridge, Gentry 466, 1882A (both
MO); Lewis et al. 5257 (MO). Hillside above Portobelo, Gentry 731C (MO). DARIEN:
Base of Cerro Pavarando, Gentry 4212 (MO). PANAMA: E of Cafnitas, Gentry 3694 (MO).
SAN BLAS: Mainland opposite Playon Chico, Gentry 6363 (MO).
2. Stizophyllum riparium (H.B.K.) Sandw., Lilloa 3: 462. 1938.-FIG. 35C-G.
Bignonia riparia H.B.K., Nov. Gen. Sp. P1. 3: 138. 1819. TYPE: Colombia, Rio Magdalena,
Humboldt & Bonpland s.n. (P).
B. brevipes Rusby, Bull. Torrey Bot. Club 27: 72. 1900. TYPE: Bolivia, Junction of Beni and
Madre de Dios Rivers, Rusby 1144 (NY).
Adenocalymma flos-ardeae Pittier, Contr. U. S. Natl. Herb. 18: 256. 1917. TYPE: Panama,
Colon, Pittier 3898 (BM, NY, US).
A. punctifolium Blake, Contr. U. S. Natl. Herb. 24: 22. 1922. TYPE: Guatemala, Izabal,
Pittier 8570 (GGH, NY, US).
Stizophyllum flos-ardeae (Pittier) Sandw., Recueil Trav. Bot. Neerl. 34: 212. 1937.
S. punctifolium (Blake) Sandw., Recueil Trav. Bot. Neerl. 34: 212. 1937.
Liana to at least 5 cm in diameter, the bark smooth, vertically striate; stem

subtetragonal with hollow center and 4 phloem arms in cross section; branchlets
terete, hollow, longitudinally striate, densely short-puberulent, the nodes with-
out interpetiolar glandular fields, often with lines connecting opposite petioles;
pseudostipules strap-shaped, caducous. Leaves 3-foliolate or 2-foliolate with
tendril or tendril scar; leaflets ovate or obovate to oblong-elliptic, acute to
acuminate, truncate to subcordate, 2.9-20 cm long and 1.9-12.5 cm wide, mem-
branaceous, often serrate when young, palmately nerved at the base, the
secondary veins (3-)4-6 on a side, regularly sunken-lepidote beneath, hence
pellucid-punctate at least when mature, puberulous on the nerves with short
simple trichomes especially beneath, drying olive to brownish with the veins
tannish beneath; tendril simple or rarely bifid with second arm minute, 7-15
cm long; terminal petiolules 3.2-7.2 cm long, lateral petiolules 0.7-4.0 cm long,
petiole 2.0-11.0 cm long, short-puberulous. Inflorescence a few-flowered axillary
raceme, the narrowly bracteate rachis and pedicels puberulous. Flowers with
inconspicuous sweet odor, calyx infundibuliform, irregularly 5-lobed to bilabiate,
5-12 mm long and 4-10 mm wide, densely puberulous, glandular on the lobes;
corolla uniformly white to cream, tubular, 3.2-5.0 cm long and 0.7-1.3 cm wide
9utside with moniliform, glandular-lepidote, and simple trichomes, pubescent
inside with moniliform trichomes on the lobes and with gland-tipped trichomes
at the level of stamen insertion; stamens didynamous, the anther thecae divari-
cate, sometimes slightly pubescent, 2-2.5 mm long, the longer filaments 1.5-1.8
cm long, shorter filaments 1.1-1.2 cm long, the staminode 3-5 mm long, inserted
10-12 mm from base of corolla tube; pistil 2.7-3.1 cm long, the ovary linear,
tetragonal, 3.5-4 mm long and 1 mm wide, glandular-lepidote, the ovules 2-
seriate in each locule; disc short-cupular, 1 mm long and 2.5-3 mm wide. Cap-
sule linear, tapering toward the ends, not flattened, 24-45 cm long and 0.5-0.7
cm wide, the surface longitudinally finely striate, short-puberulous; calyx usually
caducous; seeds 0.4-0.5 cm long and 1.3-2.4 cm wide, the wings membranaceous,
brown, weakly demarcated.
One of the commonest vines of the tropical moist forest, this species also
occurs sporadically in premontane wet forest and uncommonly in tropical wet
forest. Despite the great abundance of sterile young plants, it is seldom en-
countered in flower or fruit and is poorly represented in herbaria. It ranges
from Mexico to Peru and Bolivia. This species flowers mostly during the wet
season from April to November.
BOCAS DEL TORO: Hillside above Almirante, Gentry 2686 (MO). CANAL ZONE: Barro
Colorado Island, Croat 10220 (MO, SCZ); Gentry 5161 (MO); Kenoyer 534 (US); Shattuck
12 (A, F, MO); Standley 40923 (US); Wetmore & Woodworth 40 (A); Zetek s.n. (F, pro
parte). Along road to La Pita Signal Station, Croat 13957A (MO, SCZ). Fort Sherman, Duke
4383 (K). Boy Scout road near Madden Lake, Gentry 1375, 2052 (both MO). Las Cruces
Trail in Madden Forest, Gentry 1384 (MO). Pipeline Road, Gentry 2075, 2468 (both MO);
Gentry & Dressler 1964 (MO). Barbacoas Station, Panama Railroad, Hayes 736 (BM, K).
Rio Pedro Miguel near east Paraliso, Standley 29936 (US). CHIRIQUI: 12 mi. N of Puerto
Armuelles, Croat 21975 (MO). Burica Peninsula along Quebrada Tuco, Croat 22115 (MO).
W of San Bartolo Limite near Costa Rican border, Croat 22165 (MO). Along highway be-
tween Rios Tinta and Tabasara, Woodson et al. 386 (MO). COLON: Vicinity of Rio Piedras
along road to Portobelo, Blum et al. 2505 (SCZ). Ridge W of Portobelo, Gentry 1767 (MO).
Along Rio Fato, Pittier 3898 (BM, NY, US). DARIEN: Rio Tuira between Rio Punusa and
Rio Mangle, Duke 14628. S of Jaque, Gentry 4171 (MO). Santa Fe, Gentry 4309 (MO).
Village of Paya, Gentry 4394 (MO). Between Rios Cube and Punusa, Gentry 4468 (MO).
El Real, Gentry 4510 (MO). Near Cerro Pirre on Rio Parasenico, Gentry 4689 (MO).
PANAMA: Caflitas, Gentry 1407 (MO). Trail along Rio Pacora S from Pacora, Gentry 1413
(MO). Dam site on Rio Bayano, Gentry & Tyson 1678 (MO). Rio Pasiga, Gentry 2196,
2265, 2278 (all MO). Rio Maestro, Gentry 2228 (MO). Rio Piragua (Tabardii), Gentry
2530 (MO). Icanti, Gentry 2628 (MO). Near Capira, Gentry 3152 (MO). Ca. 16 km
from Rio Bayano crossing on trail to Santa Fe, Gentry 3811 (MO). Near Rio Espave, Gentry
3696 (MO). Near archeological site at edge of Madden Lake, Gentry 5009 (MO). Between
Pacora and Chepo, Gentry 5115 (MO). Pacora, Maurice 814 (MO, US). Juan Diaz, Standley
30480 (US). SAN BLAS: Mountains above Puerto Obaldia, Gentry 1481 (MO). VERAGUAS:
Santiago, Dwyer 1345 (MO).
36. TABEUIA
Tabebuia Gomes ex DC., Rev. Bign. (Bibl. Univ. Geneve) 14. 1838. TYPE:
T. uliginosa (Gomes) DC. = T. cassinoides (Lam.) DC.
Couralia Splitg., Tijdschr. Natuurl. Gesch. Physiol. 9: 14. 1842. TYPE: C. fluviatilis (Aubl.)
Splitg. = Tabebuia aquatilis (E. Meyer) Sprague & Sandw.
Handroanthus Mattos, Loefgrenia 50: 1. 1970. TYPE: H. albus (Cham.) Mattos = Tabebuia
alba (Chaam.) Sandw.
Shrubs to giant trees. Leaves simple, 1-foliolate, or digitately 3-7-foliolate.

Inflorescence terminal, an open or congested panicle or raceme, or reduced to
a single flower. Flowers with the calyx cupular, campanulate or tubular, truncate,
bilabiate or shallowly 5-lobed; corolla white, yellow, lavender, magenta, or red,
tubular-infundibuliform to tubular-campanulate (Panama), glabrous or puberu-
lous on the tube outside; anthers glabrous, the thecae straight, divaricate; pollen
grains single, 3-colpate, the exine finely reticulate; ovary linear oblong, often
somewhat lepidote, the ovules 2-multi-seriate in each locule. Fruit a subterete
elongate-linear or short oblong capsule, dehiscing more or less perpendicularly
to the septum, smooth to irregularly wavy-muricate, glabrous to lepidote or
variously pubescent; seeds thin and bialate with hyaline-membranaceous wings,
or thick, corky and suborbicular without wings
Fewer than 100 species in tropical America ranging from northern Mexico
and the West Indies to northern Argentina.
Useful references:
Buchinger, M. Las especies Argentinas del genero Tabebuia. Rev. Invest.

Forest. 2: 3-30. 1960.
Dugand, A. Bignoniaceae: El genero Tabebuia en Colombia. Mutisia 25:
1-31. 1956.
Gentry, A. H. A revision of Tabebuia (Bignoniaceae) in Central America.

Brittonia 22: 246-264. 1970.
a. Leaflets not stellate-pubescent; flowers (where known) rose, purple, or white; calyx
glandular-lepidote or mealy pubescent (if pubescent, corolla puberulous outside); fruit
glandular-lepidote or glabrous, the surface smooth.
b. Leaflets glandular-lepidote, not pubescent, linear-oblong to elliptic-oblong; corolla
(where known) glabrous outside; calyx and fruit strongly glandular-lepidote.
c. Leaves simple to 3-foliolate, blades linear-oblong; inflorescence (where known)
few-flowered; corolla (where known) white.
d. Tree; leaves simple; calyx (at least in fruit) spathaceously split and striate
with 5 raised lines; fruit linear, more than 12 cm long; seeds thin, bialate;
well-drained sites -8. T. striata
dd. Shrub or small tree; leaves simple to 3-foliolate; calyx 2-3-labiate, not striate;
fruit oblong, less than 12 cm long, attenuate at both ends; seeds thick, corky,
wingless; Pacific coast mangrove swamps -6. T. palustris
cc. Leaves mostly 5-foliolate, blades elliptic-ovate to elliptic-oblong; inflorescence
few- to many-flowered; corolla pale pink, rose, or lavender, rarely white.
e. Leaflets acuminate or sharply acute; inflorescence usually many-flowered; ma-
ture capsule more than 22 cm long; native and cultivated -- 7. T. rosea
ee. Leaflets obtuse; inflorescence few-flowered; mature capsule usually less than
15 cm long; rarely cultivated -5. T. pallida
bb. Leaflets pubescent at least in axils of lateral nerves beneath, ovate to ovate-lanceo-
late; corolla puberulous outside; calyx mealy pubescent; fruit glabrous
--------------------------------------------------------------------3. T. impetiginosa
aa. Leaflets stellate-pubescent at least in axils of lateral nerves beneath; flowers yellow;
corolla glabrous outside or with a few trichomes below sinuses; calyx with at least ves-
tigial yellowish to reddish-stellate tomentum; fruit stellate to woolly pubescent, or if
only inconspicuously lepidote then wavy-muricate.
f. Leaflets pubescent only in axils of lateral nerves beneath; the leaves often 7-foliolate;
calyx sparingly thick-stellate pubescent; fruit more or less lepidote, wavy-mucronate --
--------------------------------------------------------------------2. T. guayacan
ff. Leaflets with stellate trichomes scattered over undersurface or along veins; the
leaves rarely 7-foliolate; calyx stellate-pubescent, usually also with longer barbate or
simple trichomes; fruit with some stellate trichomes.
g. Leaves pubscent mostly along veins and petioles; corolla lobe venation of dried
specimens conspicuously reticulate to the margins, the color of lobes same as
that of tube; calyx without long simple trichomes, pubescent with short stellate
trichomes sometimes with some barbate trichomes, its lobes reflexed; fruit more
or less rugose, shortly stellate-pubescent; tropical wet forest (in Panama and
Costa Rica) -1. T. chrysantha
gg. Leaves densely pubescent over whole surface beneath; corolla of dried specimens
with venation becoming obscure in the lobes, the lobes appearing yellow and
distinctly different from darker-drying corolla tube; calyx golden woolly-pubes-
cent with many long (to 7 mm) simple trichomes in addition to the short-stel-
late tomentum, its lobes erect; fruit woolly-pubescent; mostly in tropical dry
forest and drier parts of tropical moist forest -4. T. ochracea
1. Tabebuia chrysantha (Jacq.) Nichols., Dict. Gard. 4: 1. 1887.
Bignonia chrysantha Jacq., Hort. Schoenb. 2: 45, tab. 211. 1797. TYPE: Venezuela, Caracas
(not seen).
Tecoma chrysantha (Jacq.) DC., Prodr. 9: 221. 1845.
Tabebuia rufescens J. R. Johnston, Proc. Amer. Acad. Arts 40: 696. 1905. TYPE: Venezuela,
Margarita Island, Johnston 79 (GHi).
Tecoma evenia Donn. Sm., Bot. Gaz. (Crawfordsville) 20: 8. 1895, pro parte. TYPE:
Guatemala, Santa Rosa, Heyde & Lux 3110 (MO; US, flowers only).
Tabebuia glomerata Urb., Fedde Repert. 14: 305. 1916. SYNTYPES: Trinidad, Tobago,
Jamaica (not seen).
Tecoma palmeri Krinzl., Fedde Repert. 17: 220. 1921. TYPE: Mexico, Guerrero, Palmer 419
(K, MO, US).
Tree to 25 m tall and 50 cm d.b.h., the bark pale to dark gray, scaly; wood
hard and heavy, dark olive brown with yellow (lapachol) deposits in the vessels;
twigs subtetragonal, varyingly stellate-pubescent when young, glabrescent, the
nodes without interpetiolar glands or pseudostipules. Leaves palmately 5(-7)-
foliolate; the leaflets broadly elliptic to oblong-obovate, abruptly acuminate,
obtuse to truncate or asymmetrically subcordate, the terminal leaflet 6-27 cm
long and 4-12.3 cm wide, the intermediate leaflets 5-25 cm long and 3.4-11
cm wide, the lateral leaflets 2.5-14 cm long and 1.4-8.3 cm wide, entire or serrate,
membranaceous to rigid-chartaceous, lepidote above and beneath, sometimes stel-
late-pubescent above, always to some extent beneath especially along main
veins, the trichomes scattered over the lamina; terminal petiolule 1.2-9 cm long,
intermediate petiolules 0.7-6.7 cm long, lateral petiolules 0.3-2.1 cm long, the
petiole 5-23 cm long, stellate-pubescent to glabrate. Inflorescence a contracted
terminal panicle, the branches stellate-pubescent, the bracts extremely reduced.
Flowers slightly aromatic, calyx campanulate, 5-lobed, the lobes often more or
less reflexed, 5-13 mm long and 4-12 mm wide, shortly stellate-pubescent;
corolla tubular-infundibuliform, 3-8 cm long, yellow with reddish penciling in
the throat and the venation (dried) reticulate to the margins of the lobes, the
dried tube and lobes indistinguishable in color, the tube 2.2-5 cm long and
0.6-2.2 cm wide at the mouth, the lobes 0.9-2.6 cm long, glabrous outside
and 4 lines of simple or usually once-branched, 2-3-celled trichomes extending
inside from the 4 lower sinuses to the level of stamen insertion, the level of
stamen insertion glandular-pubescent; stamens didynamous, inserted 4-8 mm
from the base of the corolla tube, the longer filaments 1.6-2.2 cm long, shorter
filaments 1.0-1.5 cm long, the anther thecae divaricate, 2-3 mm long, the
staminode 4-9 mm long; pistil 1.8-3.1 cm long, the ovary linear, 3.5-5 mm long
and 1.5-2 mm wide, glabrous to lepidote or sparsely stellate-pubescent, the
ovules 8-10-seriate in each locule; disc pulvinate, 0.5-1 mm long and 2-3 mm
wide. Capsule linear-cylindric, tapered at the ends, to 50 cm long and 2.0 cm
wide, sparingly short stellate pubescent; seeds 0.4-0.9 cm long and 1.4-3.3 cm
wide, the wings hyaline-membranaceous, well demarcated from the seed body.
A perplexing species, Tabebuia chrysantha includes at least two ecologically

distinct but morphologically similar forms. As now understood, one of these
occurs from Mexico to Honduras and in Trinidad and northern Venezuela mostly
in tropical dry forest. The other ranges from Nicaragua to Venezuela in tropical
and premontane wet forest.
This species flowers during the dry season with collections of flowers made
mostly in February and March. A single August collection (Gentry 5681) with
flowers was made during the abnormally dry wet season of 1972.
Despite an extensive search I was unable to locate type material of Jacquin's
Bignonia chrysantha. The illustration in Hortus Schoenbrunnensis matches either
the plant of the Venezuelan dry forest which has been known as T. rufescens
or the otherwise unnamed wet forest entity which I also include under T. chry-
/ N
_ ' _ A s~~~~~~~
FIGURE 36. Tabebuia.-A-B. T. guayacan (Seem.) Hemsl.-A. Inflorescence (X 2/5).

[After Croat 5388 (MO).]-B. Leaf (X 2/5). [After Gentry 4287 (MO).]-C. T. striata A.
Gentry, fruit (x 2/5). [After Gentry 1404 (MO).]
santha. Tabebuia chrysantha of Venezuela and Trinidad is almost identical

with the T. chrysantha of Mexico and northern Central America, but in the
intervening area of Costa Rica and Panama only the wet forest form of the
species occurs.
CANAL ZONE: Pipeline Road, Gentry 1795 (MO, SCZ), 2461 (MO). CHrRIQUi: Burica
Peninsula along Quebrada Guanabano, Croat 22514 (MO). Puerto Armuelles, Liesner 136
(MO). Area W of Puerto Armuelles, Stern & Chambers 130 (NY, US). COCLE: Base of
Cerro Pilon, Gentry & Dwyer 3661 (MO). Above El Valle, Gentry 5681 (MO). COLON:
Santa Rita Ridge, Gentry 456 (MO, WIS). DARIEN: Base of Cerro Pavarando, Gentry 4211
(MO). Base of Cerro Pirre, Gentry 4584 (MO). Summit of Cerro Pirre, Gentry 4611 (MO).
PANAMA: W of El Llano, Gentry 5074 (MO). VERAGUAS: Mountains above Santa Fe,
Gentry 3082 (MO).
2. Tabebuia guayacan (Seem.) Hemsl., Biol. Centr. Amer. Bot. 2: 495. 1882.
-FIG. 36A-B.
Tecoma guayacan Seem., Bot. Voy. Herald 180. 1854. TYPE: Panama, Cruces, Seemann 398
(BM, K).
Tree to 50 m tall and 2 m d.b.h., the bark split by long, shallow, vertical
furrows with the plates between furrows broken into scales, light tan or buff
to gray brown in color; wood extremely hard, the heavy heartwood dark olive
brown with yellow (lapachol) deposits in the vessels; twigs subtetragonal, gla-
brous, the nodes without interpetiolar glands or pseudostipules. Leaves palmately
5-7-foliolate; the leaflets lanceolate to ovate, acuminate, basally rounded to
obtuse, the terminal leaflet 9-30 cm long and 3.7-15.5 cm wide, the intermediate
leaflets 8-30 cm long and 2.8-15 cm wide, the lateral leaflets 6-21.5 cm long
and 2.1-10.7 cm wide, these often partially divided (if complete leaflets 7, the
basal pair smaller), entire (or serrulate in seedlings), membranaceous, minutely
lepidote at least beneath, stellate-pubescent with multicellular usually 3-branched
trichomes in the axils of lateral nerves beneath; terminal petiolule 2.9-7.4 cm
long, intermediate petiolule 2.4-6.0 cm long, lateral petiolule 1.1-3.4 cm long;
petiole 7-23 cm long, glabrous. Inflorescence a terminal panicle, the flowers in
2's or 3's, the branches slightly stellate-rufescent with reduced, caducous bracts.
Flowers with very faint odor, calyx campanulate, irregularly 2-5-lobed, 7-15 mm
long and 2-12 mm wide, sparsely stellate pubescent with short thick trichomes;
corolla yellow with reddish penciling in the throat; tubular-infundibuliform, 6-11
cm long and 1.3-2.2 cm wide at the mouth of tube, the tube 3.6-5.7 cm long,
the lobes 2.2-3.8 cm long, glabrous outside, inside with long mostly branched,
3-4-celled trichomes in the sinuses and from the 4 lower sinuses to the level
of stamen insertion, glandular-pubescent at the level of stamen insertion; stamens
didynamous, the anther thecae divaricate, 2-3 mm long, the longer filaments
15-21 mm long, shorter filaments 11-16 mm long, the staminode 3-10 mm long,
inserted 4.5-8 mm from the base of the corolla tube; pistil 2.4-3.3 cm long, the
ovary linear, 3-5 mm long and 1.5-2.5 mm wide, glabrous to somewhat lepidote,
the ovules 9-10-seriate in each locule; disc cupular, 0.5-1 mm long and 3-4 mm
wide. Capsule linear-cylindric, 29-61 cm long and 1.0-2.9 cm wide, essentially
glabrous or inconspicuously lepidote or thick-stellate pubescent; seeds 0.9-1.1
cm long and 3.54.0 cm wide, the wings hyaline-membranaceous, sharply de-

marcated from the seed body.
A tree of the tropical moist forest, Tabebuia guayacan occurs also, but
sparingly, in the premontane wet forest. It ranges from southern Mexico to
Colombia. This species flowers almost entirely during the dry season from
January to May with an occasional individual blooming asynchronously during
the wet season. Seed release occurs during the late dry season and early wet
season; the seeds are wind-dispersed.
Tabebuia guayacan can be recognized vegetatively by the stellate trichomes
beneath in the axils of the lateral nerves of an otherwise glabrous leaflet and
by its tendency to 7-foliolate leaves. It is closely related to T. serratifolia (Vahl)
Nichols., a species widespread in South America, which differs in its simple
trichomes in the axils of the lateral nerves beneath and in the longer trichomes
inside the corolla tube.
CANAL ZONE: Barro Colorado Island, Carpenter 66 (F); Croat 4120 (SCZ), 4121,
5388, 7940 (all MO, SCZ); Gentry 450 (MO, WIS), 712A (MO); Shattuck 821 (F, MO);
Standley 31419 (US). Near Gamboa, Allen 228 (A, GH, US). Rio Cristal on Pan Am High-
way, Playa Coronado, Croat 14256 (MO). Balboa, Gentry 423 (MO, WIS). Pipeline Road,
Gentry 1586, 2410, 4788 (all MO). Road to Gamboa airport, Gentry 2045, 4855 (both MO).
Albrook NCO Club, Gentry 1736 (MO). Road K-10 at edge of Canal Zone, Gentry 1958
(MO). Vicinity of Summit Garden, Gentry 4284, 5119 (both MO). Around Gamboa, Pittier
2694 (BM, GH, NY, US). Near village of Cruces, Seemann 398 (BM, K). Summit, Sharp
s.n. (SCZ). Hills N of Frijoles, Standley 27532 (US). Albrook NCO Club, Tyson 5403
(MO, SCZ). N of Summit, Woodson et al. 768 (A, MO, NY, US); Woodworth & Vestal 732
(F). Without locality, Johansen 72 (US). CHIRIQUI: Progreso, Cooper & Slater 311 (F, US).
COLON: Along Rio Boqueron above Peluca Hydrographic Station, Hunter & Allen 656 (G,
GH, MO, NY). DARIEN: Vicinity of Pinogana, Allen 4294 (CGH, MO). S of La Palma,
Gentry 4287 (MO). Near village of Pirre, Gentry 4781 (MO). PANAMA: Ciudad de Panama,
Gentry 1940 (MO, SCZ); Holdridge 6477 (PMA).
3. Tabebuia impetiginosa (Mart. ex DC.) Standl., Publ. Field Mus. Nat.

Hist., Bot. Ser. 11: 176. 1936.
Bignonia heptaphylla sensu auct., non Vell. Fl. Flum. 6: 251. 1825; pl. 48. 1827.
Tecoma heptaphylla (Vell.) Mart., Flora 24, Beibl. 2: 13. 1841, sensu auct. non Vell.
Tecoma impetiginosa Mart., Syst. Mat. Med. Veg. Bras. 54. 1843, nomen nudum.
T. ipe Mart., Syst. Mat. Med. Veg. Bras. 55. 1843, nomen nudum.
T. impetiginosa Mart. ex DC., Prodr. 9: 218. 1845. TYPE: Brazil, Piauhy, Martius 2446 (G-
DC, M).
T. eximia Miq., Linnaea 22: 803. 1849. LECTOTYPE: Brazil, Bahia, Blanchet 3963, leaves
flowers excluded (BR, C, G, K, MO, P, U).
Tabebuia avellanedae Lorentz ex Griseb., Symb. Fl. Argent. 258. 1879. TYPE: Argentina
(not seen).
T. palmeri Rose, Contr. U. S. Natl. Herb. 1: 109. 1891. TYPE: Mexico, Sonora, Palmer 320
(GH, US).
Tecoma impetiginosa var. lepidota Bur., Vidensk. Meddel. Dansk Naturhist. Foren. K
1893: 114. 1894. TYPE: Brazil, Rio de Janeiro, Glaziou 11241 (P).
T. ipe Mart. ex K. Schum. in Engler & Prantl, Nat. Pflanzenf. 4(3b): 238. 1894. TYPE:
Brazil (not seen).
T. adenophylla K. Schum. ex Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 412. 1897. TYPE:
Brazil, Goyaz, Glaziou 21841 (BR, C).
Gelseminum avellanedae (Lorentz ex Griseb.) Kuntze, Rev. Gen. P1. 3, 2: 245. 1898.
Tecoma ipe var. integra Sprague, Bull. Herb. Boiss., ser. 2. 5: 86. 1905. TYPE: Paraguay,
Hassler 3065 (K).
Tecoma avellanedae (Lorentz ex Griseb.) Speg. in Speg. & Girola, Cat. Descr. Maderas 379.
1910.
T. avellanedae var. alba Lillo, Segunda Contr. Conoc. Arbol. Argen. 13. 1917. TYPE: Argentina
(not seen).
T. integra (Sprague) Chodat, Bull. Soc. Bot. Geneve, ser. 2. 9: 242. 1917.
Tabebuia nicaraguensis Blake, Contr. Gray Herb. 52: 95. 1917. TYPE: Nicaragua, Hacienda
Campuscus, Baker 2258 (GH, holotype; MO, US, isotypes).
T. dugandii Standl., Trop. Woods 36: 17. 1933. TYPE: Colombia, Atlantico, Dugand 345
(F, US).
T. ipe (Mart. ex Schum.) Standl., Field Mus. Nat. Hist., Bot. Ser. 11: 176. 1936.
T. eximia (Miq.) Sandw., Lloydia 2: 213. 1939.
T. avellanedae var. paulensis Toledo, Arquiv. Bot. Est. Sdo Paulo, n. s. 3: 33. 1952. TYPE:
Brazil (not seen).
T. heptaphylla (Vell.) Toledo, Arquiv. Bot. Estado Sao Paulo, n. s. 3: 33. 1952, sensu
auct. non Vell.
T. schunkevigoi Simpson, Fieldiana, Bot. 36: 1. 1972. TYPE: Peru, Huanuco, Schunke 2596
(F, holotype; MO, isotype).
Tree to 20 m tall and 70 cm d.b.h., the bark relatively smooth, gray; wood
greenish brown, dense, the vessels containing yellow powder (lapachol); twigs
terete, glabrate, tips of shoots mealy pubescent, the nodes without interpetiolar
glands or pseudostipules. Leaves palmately 5-7-foliolate, often anisophyllous;
the leaflets ovate to elliptic, acuminate, cuneate to rounded or almost subcordate,
the terminal leaflet 5-19 cm long and 1.5-8 cm wide, the intermediate leaflet
4-16 cm long and 2.8-9 cm wide, the basal leaflet 2.2-11 cm long and 1.5-8.3
cm wide, entire or serrulate, membranaceous to chartaceous, somewhat lepidote
above and beneath, pubescent with simple or forked trichomes at least in the
axils of lateral veins beneath or pubescent over the whole undersurface, often
puberulous on the midrib above; terminal petiolules 1-4.2 cm long, intermediate
petiolules 0.9-4 cm long, lateral petiolules 0.2-2 cm long, the petiole 4-13 cm
long, lepidote and puberulous. Inflorescence a terminal, usually more or less
congested panicle, the flowers in 3's, the branches whitish to tannish mealy-
pubescent with short, thick-stellate trichomes. Flowers with the corolla pinkish-
purple to deep magenta outside and on the lobes, the throat opening yellow,
turning magenta, tubular-campanulate, 4-7.5 cm long and 1.2-5 cm wide at
the mouth, the tube 2.5-5 cm long, the lobes 0.9-2.0 cm long, puberulous outside,
inside with a few, scattered, simple trichomes in the tube and pubescent at
cm long, the staminode 3-7 mm long, inserted 5-6 mm from the base of the
corolla tube; pistil 2.7-3.4 cm long, the ovary linear, 3-4 mm long and 1 mm
wide, subtetragonal in cross section, glabrous to slightly lepidote, the ovules
ca. 4-seriate in each locule; disc cupuliform, 1-1.5 mm long and 2 mm wide.
Capsule elongate-cylindrical, attenuate at both ends, 12-56 cm long and 1.3-2.5
cm wide, glabrous; seeds 1-1.6 cm long and 3-4.8 cm wide, the wings hyaline-
membranaceous, sharply demarcated from the seed body.
A characteristic tree of the premontane moist forest, Tabebuia impetiginosa

also occurs to some extent in the tropical dry forest. It ranges from northern
Mexico to Argentina. This species flowers in the early dry season from De-
cember to February. The wind-dispersed seeds are released in late dry season.
The name Tabebuia impetiginosa has been used only in central Brazil where
it has been separated from the other species of a wide ranging complex because
of its pubescent leaflets. However, pubescent-leaved plants occur as occasional
variants throughout the range of the species in Central America, and have been
named as T. nicaraguensis. In Brazil the glabrate form has been given the
names T. impetiginosa var. lepidota and T. adenophylla.
Geographical separation and characters of the number of leaflets and leaf
indumentum have been used to maintain these various populations as separate
species. Disjunctions in the range of this composite species are like those of
a number of other organisms which are restricted to the dry forest habitat with
the species occupying all of the suitable habitats, and the various populations
separated only by the present expansion of the intervening wet forest areas.
The Central American, Colombian, and western Argentinian populations
have 5-foliolate leaves with entire leaflets which are pubescent only in the axils
of the lateral nerves below; the eastern Argentinian plants have mostly 7-foliolate
leaves with dentate leaflets which are pubescent only in the axils of the lateral
nerves beneath; and most Brazilian plants have entire leaflets which are pu-
bescent over the whole lower surface.
Recent collections from the dry forest area around the Golfo de San Miguel,
Darien Province, come from the middle of the supposed area of disjunction
between the Central American and Colombian populations, further supporting
the morphological evidence for union of T. palmeri and T. dugandii under T.
impetiginosa. The Darien specimens are noteworthy in having the mostly
7-foliolate leaves of the Argentinian and some Brazilian forms.
The serrate-leaved form of eastern Argentina and adjacent Brazil was con-
sidered distinct by Buchinger (1960) as T. ipe (Mart. ex K. Schum.) Standl.,
but I suspect it merits varietal rather than specific recognition.
Most specimens identified as Tabebuia heptaphylla (Vell.) Toldeo belong
to this species, and that name has been used (as Tecoma heptaphylla) in Brazil.
The basionym Bignonia heptaphylla Vell. antedates the basionym of Tabebuia
impetiginosa. Vellozo's drawing of a 7-foliolate, serrate-leafleted plant could
refer to this species or several others. The description of the corolla as "violacea"
points toward this species, but the strongly 5-dentate calyx does not. The fruit
as figured and described is wrinkled-striate with a persistent calyx, conditions
which do not match T. impetiginosa. In the absence of a type specimen, B.
heptaphylla has been subject to diverse interpretations, and because of these
several problems, I consider Tabebuia heptaphylla to be a nomen confusum
and have adopted the name T. impetiginosa.
DARIEN: Isla Cartagena, Golfo de San Miguel, Gentry 3935 (MO). Shore near Majagualito
across from La Palma, Gentry 3989 (MO). Isla Encanto in Golfo de San Miguel, Gentry 4023
(MO). PANAMA: Vicinity of Bejuco, Allen 1630 (F, GH, NY, US), 4203 (MO).
4. Tabebuia ochracea (Chain.) Standl., Field Mus. Nat. Hist., Bot. Ser.
11: 176. 1936.
Tecoma ochracea Cham., Linnaea 7: 653. 1832. TYPE: Brazil, Sellow s.n. (K, W).
T. hypodictyon DC., Prodr. 9: 217. 1845. TYPE: Brazil, Villa Barra, Blanchet, 1840 (G-DC).
T. hassleri Spr., Trans. Proc. Bot. Soc. Edinburgh 48: 435. 1904. SYNTYPES: Paraguay, San
Estanislao, Hassler 4159 (CGH, K), 4164 (K).
Tabebuia hypodictyon (DC.) Standl., Field Mus. Nat. Hist., Bot. Ser. 11: 176. 1936.
4a. Tabebuia ochracea subsp. neochrysantha (A. Gentry) A. Gentry, comb.

& stat. nov.
T. neochrysantha A. Gentry, Brittonia 22: 260. 1970. TYPE: Costa Rica, Guanacaste, Gentry
355 (BM, F, MO, UC, US, WIS).
?Bignonia heterotricha DC., Rev. Bign. (Bibl. Univ. Geneve) 22. 1838, nomen nudum.
?Tecoma heterotricha DC., Prodr. 9: 219. 1845. TYPE: Venezuela, Caracas, Vargas s.n. (G-DC).
?Tabebuia heterotricha (DC.) Hemsl., Biol. Centr. Amer. Bot. 2: 495. 1882.
T. chrysantha (Jacq.) Nichols., Dict. Gard. 4: 1. 1897, sensu Sandw., non Jacq.
Tree to 25 m tall and 50 cm d.b.h., the bark with dark shallow vertical
furrows separating lighter, flat-surfaced ridges; wood hard, heavy, the heart-
wood dark olive brown with yellow (lapachol) deposits in the vessels; twigs
subtetragonal, stellate-pubescent when young, glabrescent, the nodes without
glands or pseudostipules. Leaves palmately 5-foliolate; the leaflets oblong-
obovate or oblong-elliptic, abruptly acuminate to apiculate, basally obtuse to
truncate, the terminal leaflet 5-22 cm long and 1.8-14.4 cm wide, the inter-
mediate leaflets 4.2-19.8 cm long and 1.4-13.2 cm wide, the lateral leaflets
2.8-11 cm long and 1.6-8.0 cm wide, entire to serrate, membranaceous, lepidote
above and beneath, stellate pubescent but glabrescent above, densely and per-
sistently stellate-pubescent beneath; terminal petiolule 0.8-5.8 cm long, inter-
mediate petiolules 0.7-5.1 cm long, lateral petiolules 0.2-1.8 cm long, the petiole
6-18 cm long, stellate-pubescent. Inflorescence a contracted terminal panicle,
the pedicels and peduncle obsolete, the branches stellate-pubescent. Flowers
with a sweet fragrance, calyx campanulate, 5-lobed, 8-15 mm long and 5-1.0
mm wide, woolly-pubescent with shorter, stellate tomentum and thick covering
of long (to 7 mm), simple trichomes, these usually branched at the extreme
base; corolla yellow with reddish penciling in the throat, the venation drying
obscure in the lobes, the tube and corolla different in color, the tube usually
brownish, the lobes yellowish; tubular-infundibuliform, 4-8.3 cm long and
0.8-2.1 cm wide at the mouth, the tube 3-5.8 cm long, the lobes 1.0-2.4 cm
long, glabrous outside except for some trichomes at the base of the lobes,
villous on the anterior surface within, also with a few scattered simple trichomes
adaxially in the throat, glandular-pubescent at the level of stamen insertion;
stamens didynamous, the anther thecae divaricate, 1.5-2.5 mm long, the longer
filaments 1.5-2.0 cm long, shorter filaments 0.9-1.5 cm long, the staminode
2-11 mm long, inserted 6-7 mm from the base of the corolla tube; pistil 2.0-2.7
cm long, the ovary linear, 3-5 mm long and 1.5-2 mm wide, lepidote to some-
what puberulous with simple and stellate trichomes, the ovules more or less
10-seriate in each locule; disc pulvinate, 1 mm long and 2-3 mm wide. Capsule
linear-cylindric, tapered at the ends, 13-35 cm long and 1-1.5 cm wide, abun-
dantly golden woolly pubescent with a short stellate tomentum and longer
indumentum of simple trichomes (these usually with stellate base); seeds
0.4-0.8 cm long and 1.8-2.9 cm wide, the wings hyaline-membranaceous, well

demarcated from the seed body.
A tree of the tropical dry forest, premontane moist forest, and drier parts
of the tropical moist forest, Tabebuta ochracea subsp. neochrysantha occurs
from Honduras and extreme eastern El Salvador to Venezuela and Trinidad.
Other forms of the species occur south to Paraguay and Brazil. This species
flowers in the dry season from January to April. The wind-dispersed seeds are
released in mid to late dry season.
In the author's 1970 study the Central American population was considered
to be distinct from related populations in South America, mainly on the basis
of comparison of Central American material with the relevant types by curators
of European herbaria. However, consultation of type material indicates that
the Central American T. neochrysantha is conspecific with T. ochracea Cham.,
an earlier name.
There are overlapping quantitative differences between T. neochrysantha
and T. ochracea which are geographically correlated. The northern form has
usually longer calyx trichomes, a usually more conspicuous differentiation be-
tween corolla lobes and tube in dried specimens, somewhat shorter trichomes
in the corolla tube, and an often softer leaf indumentum with usually less
pronounced reticulation beneath. Sandwith in unpublished notes at Kew (K)
separated this form as T. chrysantha from T. ochracea on the basis of its longer
trichomes within the corolla tube on the anterior side, but this character shows
too much variability for specific segregation. The evidence supports formal
recognition for the northern plant, subspecies neochrysantha.
While the Brazilian type specimen of Tecoma ochracea (Sellow s.n., dupli-
cates at K, G-DC, W) is a reasonable match for Central American material
of T. neochrysantha, I have as yet been unable to separate the South American
taxa of this complex, some very different from Central American material, but
it is probable that the Brazilian material will prove divisible into several species.
However, the longer calyx trichomes and densely stellate-pubescent leaves, the
most conspicuous characters of T. neochrysantha, are much more variable than
previously supposed. Most Costa Rican populations have shorter calyx trichomes
than those from Panama or Honduras, while juvenile leaves are normally much
less thickly stellate pubescent than mature leaves of the same plant. Tabebuia
heterotricha, although considered distinct in previous studies, is apparently an
atypical form of this subspecies with less pubescent leaves similar to the common
juvenile form.
CANAL ZONE: Ancon Hill, Allen 4520 (G, MO). Balboa, Gentry 424. Madden Lake
near Natural Bridge, Gentry 701 (MO). Junction of Chiva Chiva Trail and Gaillard High-
way, Gentry 4786, 4861A, 4861B, 5117 (all MO). N of Paraiso, Gentry 4958, 4959 (both MO).
Ancon, Keenan 323 (US). Sabanas, Paul 307 (US). COCLE: S of El Valle de Anton, Allen
4477 (G, MO). Penonome and vicinity, Williams 194 (NY). DARIEN: Punto Sabanas opposite
La Palma, Gentry 3944 (MO). Punta Patinio NNE of Garachine, Gentry 4043 (MO).
HERRERA: E of Las Minas, Gentry 3146 (MO). PANAMA: Vicinity of Chorrera, Allen 1698
(F, MO, NY, US), 4265 (C, GlH, MO). Rio Pasiga, Gentry 2248 (MO). Farfan Beach road,
Gentry & Dwyer 3543 (MO). W of Cafiitas, Gentry 4964 (MO). Alhajuela, Pittier 3501
(GH, NY, US). VERAGUAS: S of Santa Fe, Gentry 2937 (MO). Near Rio Santa Maria S of
San Francisco, Gentry 3118 (MO). WITHOUT LOCALITY: Warscewicz 52 (G).
5. Tabebuia pallida (Lindl.) Miers, Proc. Roy. Hort. Soc. 3: 199. 1863.8
Bignonia pallida Lindl., Bot. Reg. 12: tab. 965. 1826. TYPE: St. Vincent, cultivated, Caley
s.n. (not seen).
Shrub or small tree. Leaves 1-5-foliolate (in Panama mostly 5-foliolate)

the leaflets elliptic to elliptic-oblong, obtuse, basally rounded, chartaceous,
lepidote. Inflorescence a few-flowered terminal panicle, often reduced to one
or 2 flowers. Flowers with the calyx cupular, bilabiate, lepidote; corolla pinkish-
lavender to almost white, the throat opening yellow; tubular-infundibuliform;
stamens didynamous, the thecae divaricate; ovary lepidote, the ovules 2-seriate
in each locule. Capsule linear-cylindric, attenuate at both ends, 7-15 cm long
(rarely to 22 cm, fide Sandwith, 1954), lepidote, the calyx persistent; seeds
bialate, 7-9 mm long and 2-3 cm wide, the wings hyaline-membranaceous,
sharply demarcated from the seed body.
A common and highly variable West Indian species, T. pallida is cultivated

as an ornamental in various parts of the tropics. It is known in Panama only
from a single tree at Summit Gardens. Tabebuia pallida is close to T. rosea
but is separated by its usually smaller, obtuse leaflets.
CANAL ZONE: Summit Gardens, cultivated, Nee & Mori 3579 (MO, WIS).
6. Tabebuia palustris Hemsl., Biol. Centr. Amer. Bot. 2: 495. 1882. TYPE:
Panama, Canal Zone, Hayes 80 (BM, K).
Shrub or small tree, often twisted, to 4 m tall and 5 cm d.b.h., the bark
smooth, pale gray; wood whitish, soft; twigs terete, lepidote, the nodes without
interpetiolar glandular fields or pseudostipules. Leaves simple or 3-foliolate;
the leaflets narrowly elliptic or oblong, acute, cuneate to subsessile, the terminal
leaflet 10-19 cm long and 2.1-5.6 cm wide, lateral leaflets when present 5-19
cm long and 1.1-5.1 cm wide, subcoriaceous, densely lepidote beneath, less
so above, gray-green when dry, the terminal petiolule 1-3.5 cm long, lateral
petiolules when present 0.2-1.5 cm long, the petiole 2.3-10 cm long, lepidote.
Inflorescence a 2-5-flowered terminal cyme on a reduced branch usually in
a dichotomy of the branches, the pedicels and peduncle lepidote. Flowers
with the calyx irregularly bilabiate or 3-labiate, cupular, 10-20 mm long and
4-10 mm wide, lepidote; corolla white with yellow throat ridges, tubular-
infundibuliform, 5-7 cm long and 0.9-1.5 cm wide at the mouth, the tube
2.8-5.8 cm long, the lobes 0.8-1.8 cm long, glabrous outside, inside pubescent
on throat ridges and at the level of stamen insertion, sparsely so on lobes and
in sinuses; stamens didynamous, the anther thecae divaricate, 3 mm long, the
longer filaments 1.8-2.6 cm, shorter filaments 1.4-1.9 cm long, the staminode
5-7 mm long, inserted 4-6 mm from base of corolla, pistil 2.9-3.1 cm long,
the ovary linear-conical, 4.5-5 mm long and 1 mm wide, densely glandular-
8The synonymy of this species is much more complex than indicated here, with m
local varieties given specific epithets on different West Indian islands. Only these names have
been used for Panamanian material.
lepidote, the trichomes extending up the base of style, the ovules 2-seriate in
each locule; disc pulvinate, 2 mm long and 3 mm wide. Capsule oblong-cylindric,
attenuate at both ends, 8-11 cm long and 1.6-2.6 cm wide, densely lepidote,
the calyx persistent; seeds suborbicular, thick, corky, brown, 1.4-1.8 cm long
and 1.8-2.2 cm wide, wings essentially lacking.
Restricted to coastal mangrove swamps and especially common along the

mouths of rivers near the fresh water/salt water interchange, this seldom-
collected plant ranges along the Pacific coast from central Costa Rica to Colom-
bia. It blooms more or less throughout the year.
This species is distinct from other Central American species of Bignoniaceae
in its shrubby habit, mangrove habitat and thick, corky seeds. Its closest rela-
tive is probably T. aquatilis (E. Meyer) Sprague & Sandw. of the Amazon and
Orinoco valleys, which has similar fruit and seeds but 5-foliolate leaves with
the veins drying reddish-yellow beneath. Tabebuia insignis of South America
is also related but has winged seeds.
CANAL ZONE: Along creek at Miraflores Lock, Gentry 422 (MO, WIS), 2104 (MO).
Rio Grande swamp, Hayes 80 (BM, K). Miraflores Locks area, Tyson 1269 (SCZ). Rio
Grande, Woodson et al. 760 (A, MICH, MO, NY, US). COCLE: Penonome and vicinity, Wil-
liams 292 (NY, US). DARIEN: Chepigana, Duke & Bristan 282 (MO). 0-4 mi. up Rio
Sabanas from Santa Fe, Duke 4148, 4172 (both MO). Rio Sabanas just below Santa Fe,
Gentry 4080 (MO). Rio Sabanas at junction with Rio Lara, Gentry 4084 (MO). Rio Jaque
near mouth, Gentry 4191 (MO). E of Santa Fe, Tyson et al. 4679 (MO, SCZ). PANAMA':
San Jose Island, Erlanson 224, 330 (both GH, US); Harlow 59 (GH); Johnston 2 (GH, US),
798 (GH).
7. Tabebuia rosea (Bertol.) DC., Prodr. 9: 215. 1845.
Tecoma rosea Bertol., Fl. Guatimal. 25. 1840. TYPE: Guatemala, Esquintla, Velasquez s.n.
(FI, fide Sandw.).
T. mexicana Mart. ex DC., Prodr. 9: 218. 1845. TYPE: Mexico, Karwinski s.n. (M).
Sparattosperma rosea (Bertol.) Miers, Proc. Roy. Hort. Soc. 3: 99. 1863.
Tabebuia mexicana (Mart. ex DC.) Hemsl., Biol. Centr. Amer. Bot. 2: 495. 1882.
T. pentaphylla (L.) Hemsl., Biol. Centr. Amer. Bot. 2: 495. 1882, non B. pentaphylla U.
Couralia rosea (Bertol.) Donn. Sm., Bot. Gaz. (Crawfordsville) 20: 9. 1895.
Tecoma evenia Donn. Sm., Bot. Gaz. (Crawfordsville) 20: 9. 1895. TYPE: Guatemala, Santa
Rosa, Heyde & Lux 3110 (MO, US, leaves only).
T. punctatissima Kranzl., Fedde Repert. 17: 221. 1921. TYPE: Colombia, Cundinamarca,
Karsten s.n. (W).
Tabebuia punctatissima (Kranzl.) Standl., Trop. Woods 36: 18. 1933.
Tree to 25 or 30 m tall and 1 m d.b.h., the bark narrowly vertically fissured

with corky ridges, dark gray to blackish; wood superficially similar to oak, of
medium density, light grayish brown with distinct pattern from stripings of
brown paratracheal parenchyma, when fresh with faint sweet odor suggesting
watermelon; twigs subtetragonal, lepidote, with thick pith, the nodes without
interpetiolar glands or pseudostipules. Leaves palmately 5-foliolate, often aniso-
phyllous; the leaflets elliptic to elliptic-oblong, acute to acuminate, basally
rounded to cuneate, the terminal leaflet 8-35 cm long and 3-18 cm wide, the
intermediate pair 6-34 cm long and 3-17 cm wide, the lateral pair 3.5-25 cm
long and 1.4-12 cm wide, entire, subcoriaceous to chartaceous, lepidote above
and beneath, gray-green when dry; the terminal petiolule 2-11 cm long, inter-
mediate petiolules 1.5-9.5 cm long, lateral petiolules 0.2-2.4 cm long, the

petiole 5-32 cm long, lepidote. Inflorescence a terminal panicle with a pair
of subulate bracts subtending each dichotomy, the branches densely lepidote.
Flowers slightly sweet-smelling, calyx cupular, bilabiate, 11-21 cm long and
6-12 cm wide, densely lepidote; corolla white to pinkish-lavender or magenta,
the throat opening yellow, turning white, tubular-infundibuliform, 5-10 cm
long and 1.5-3.2 cm wide at the mouth of tube, the tube 3-5.8 cm long, the
lobes 2.2-2.6 cm long, glabrous outside, ciliate, sparsely pubescent within with
small mostly unicellular trichomes on throat ridges and gland-tipped trichomes
at the point of stamen insertion; stamens didynamous, the thecae divaricate,
cm long, the staminode 2-6 mm long, inserted 5-10 mm from base of corolla;
pistil 1.9-3.2 cm long, the ovary linear, 5-8 mm long and 1 mm wide, densely
lepidote, the ovules 2-seriate in each locule; disc tapered-cupular, 2-3 mm long
and 3-3.5 mm wide. Capsule linear-cylindric, attenuate at both ends, 22-38
cm long and 0.9-1.5 cm wide, lepidote, the calyx usually persistent; seeds 0.7-1.0
cm long and 2.84.4 cm wide, the wings hyaline-membranaceous, sharply de-
marcated from the seed body.
A widespread tree in tropical wet forest, tropical moist forest, and tropical
dry forest, it is also extensively cultivated as an ornamental and its wood is
much used for furniture. Altitudinally it reaches 1500 meters. It ranges from
southern Mexico to Venezuela. This species flowers mostly during the dry
season, occasionally early in the wet season. The wind-dispersed seeds are
released mostly in the early wet season in late April and May.
The commercial importance of Tabebuia rosea is such that it is usually
harvested by timber cruisers before clearing or general lumbering of an area
begins. In Panama it is often the fourth tree species after Swietenia, Cedrela,
and Bombacopsis to be removed and is usually the last species to be selectively
cut.
For a discussion of the nomenclatural problems of this species see Sandwith
(Kew Bull. 1953: 453. 1954), who showed the propriety of using the name
T. rosea. It has also been widely known as T. pentaphylla.
Tabebuia rosea is recognized by its 5-foliolate, lepidote but otherwise com-
pletely glabrous leaves.
BOCAS DEL TORO: Punta Robalo to Robalo River, Seibert 1564 (MO, US). CANAL ZONE:
Barro Colorado Island, Brown 18 (F); Croat 4662 (MO, SCZ), 4674, 4884 (both MO),
5436 (MO, SCZ), 5726 (MO), 7771 (MO, SCZ), 7928 (MO), 8184 (MO, SCZ),
8299 (MO), 8394 (MO, SCZ); Gentry 444 (WIS); Shattuck 565 (MO), 650 (F, MO),
Standley 40928 (US); Wetmore et al. 85 (A, F, GH, MO); Wilson 102 (F, MO). Monte
Lirio, Christopherson 125 (NY, US). Boy Scout road near Madden Lake, Gentry 2057 (MO).
Madden Dam, Gentry 5067 (MO). Around Culebra, Pittier 2767 (GH, NY). Las Cascades
Plantation near Summit, Standley 25800 (US). Mount Hope Cemetery, Standley 28825 (US).
Curundu in back of Club house, Tyson 3657 (SCZ). CHIRIQUI: Progreso, Cooper & Slater
268 (F, US), 311 (NY). COCLE1: Near El Valle de Anton, Gentry & Dwyer 3597, 3685
(both MO). Nata, Seemann 1126 (K). Penonome and vicinity, Williams 429 (NY, US).
COLON: Road to Portobelo, Gentry 1772 (MO). Tres Brazos sawmill at Icacal between Salud
and Boca de Rio Indio, Howell 55 (MO). DARIEN: Vicinity of Pinogana, Allen 4287 (G,
MO, NY, US). S of Jaqu-e, Gentry 4118 (MO). Summit of Cerro Chucula, Gentry 4243
(MO). S of La Palma, Gentry 4291 (MO). El Real, Gentry 4318, 4579 (both MO). Rio
Paya from Rio Tuira to village of Paya, Gentry 4390 (MO). Quebrada Sierpe between Rios
Cube and Punusa, Gentry 4469 (MO). Near Cerro Pirre on Rio Parasenico, Gentry 4694
(MO). HERRERA: S of Ocu', Gentry 3121 (MO). PANAMA': Universidad de Panamai, Diaz s.n.
(MO, PMA). Vicinity of El Llano, Duke 5819 (MO). Perlas Islands, Isla Espiritu Santo, Duke
10456 (MO). Cafiitas, Gentry 1404 (MO). Between El Llano and Chepo, Gentry 1408 (MO).
Cerro Campana above Su-Lin Motel, Gentry 1827 (MO). Rio Pasiga, Gentry 2194, 2343 (both
MO). Rio Piragua (Tabardi), Gentry 2521 (MO). Near Rio Espave, Gentry 3712 (MO).
Near last river before Rio Parti on trail from Bayano crossing to Santa F6, Gentry 3799 (MO
W of El Llano, Gentry 4962 (MO). Dam site on Rio Bayano, Gentry & Tyson 1648 (MO).
Swamp between El Jagua Hunting Club and El Congor Hill, Hunter & Allen 476 (F, G, K
MO). Perlas Islands, San Jose Island, Johnston 519 (GH). Chepo, Kluge 20 (F, US). E
of Rio Tocumen, Standley 26666 (US). Rio Tapia, Standley 28094 (MO, US). SAN BLAS:
Vicinity of Puerto Obaldia, Gentry 1528A (MO). VERAGUAS: 1-2 mi. above Santa Fe, Gentry
3039 (MO). Without locality, Seemann s.n. (K). WITHOUT LOCALITY: Hayes 982 (NY).
Isthmus of Darien, Barclay s.n. (BM, K).
8. Tabebuia striata A. Gentry.9 TYPE: Panama, San Blas, Puerto Obaldia,

Gentry 1484 (MO).-FIG. 36C.
Tree to 15 m tall and 10 cm d.b.h., the bark pale gray, smooth, with a
faint sweetish odor when cut, the branches strongly ascending; twigs terete
with a large pith, lepidote. Leaves simple, opposite, elliptic to narrowly obovate,
acute to obtuse, basally cuneate, 21-38 cm long and 6.5-13.3 cm wide, regularly
lepidote above and beneath with 2 distinct sizes of scales, somewhat lepidote-
glandular at the base of the blade, drying olive with the veins tan, midvein
conspicuously raised beneath; petiole 1.2-6 cm long. Inflorescence unknown.
Flowers with the calyx (in fruit) spathaceous, the apex often somewhat split,
2.5-3 cm long and 0.9-1.9 cm wide, densely lepidote, drying gray, conspicu-
ously 5-striate with slightly raised lines; corolla, stamens and pistil unknown.
Capsule subterete, dehiscing perpendicularly to the septum, 12-25 cm long
and 2-2.8 cm wide, the valves coriaceous, densely lepidote, drying gray, the
calyx persistent; seeds thin, bialate, 1.1-1.5 cm long and 1.9-2.5 cm wide, the
wings short, thin, basally membranaceous at the extreme tip.
A tree of the tropical wet forest, this species is known only from the type
locality in extreme eastern Panama almost on the Colombian border. It is
locally not uncommon, and seedlings were found at several places on the
mountain slopes above Puerto Obaldia.
Although flowers of this species are not known, it is distinct from all othe
species of the genus. Its simple leaves and lack of pubescence other than lepido
scales relate it to T. insignis (Miq.) Sandw. and T. stenocalyx Sprague & Stapf.
However its leaves are thinner and much longer than in either of these specie
Its spathaceously split calyx in fruit differs from that of T. insignis. The fru
of T. stenocalyx is similar to T. striata but narrower, 1.5-2 cm wide (fide
Sandwith, 1954). Its calyx also splits spathaceously in fruit but is not so
conspicuously striate. Although T. striata is close to T. stenocalyx, I do not
9Tabebuia striata A. Gentry, sp. nov. Arbor ad 15 metros, cortice laevi; folia simplicia
obelliptica, cuneata ad basem, lepidota; inflorescentia ignota; calyx (in fructo) spathaceus,
dense lepidotus, striatus quinque lineis elevatis; corolla, stamina et pistillum ignota; capsula
subteres perlepidota; semina tenuia, bialata.
think the two are the same, especially in view of the latter's distribution (Trini-
dad, the Guianas, and the Departments of Bolivar and Delta Amacuro in
Venezuela). Romeroa verticillata Dugand of Colombia also seems related to
T. striata, especially on the basis of leaves. It differs in a much wider fruit,
usually whorled leaves, orbicular seeds with opaque wings, and in lacking a
striate calyx.
SAN BLAS: Puerto Obaldia, Gentry 1484, 1488 (both MO).
37. TANAECIUM
Tanaecium Sw., Nov. Gen. Sp. P1. 2: 91. 1788. TYPE: T. jaroba Sw.
Osmohydrophora Barb. Rodr., Vellosia, ed. 2. 1: 48. 1891. TYPE: 0. nocturna Barb. Rodr.
= Tanaecium nocturnum (Barb. Rodr.) Bur. & K. Schum.
Lianas; stems with 8 or 16 phloem arms in cross section; branchlets terete,

with or without interpetiolar glandular fields; pseudostipules inconspicuous or
lacking. Leaves 2-3-foliolate, the terminal leaflet often replaced by a simple
tendril; vegetative parts often with a conspicuous almond (cyanide) odor.
Inflorescence an axillary or terminal raceme or racemose panicle. Flowers
fragrant, calyx cupular, more or less truncate, sometimes minutely 5-denticulate,
often with plate-shaped glands in the upper part; corolla white, elongate-tubular
to hypocrateriform, pubescent or glabrous without; stamens usually more or
less exserted, anthers glabrous, the thecae straight or arcuate, somewhat diver-
gent; pollen grains single, 3-colpate or irregularly many-porate, the exine finely
reticulate; ovary oblong, lepidote, the ovules multiseriate in each locule; disc
cupular-pulvinate. Fruit an oblong-cylindric capsule, the valves parallel to the
septum, thick, woody, not at all compressed, smooth, the median nerve invisible
or furrowed, wingless or bialate with brownish irregularly-edged, poorly
demarcated membranaceous wings.
Six species of tropical America ranging from Costa Rica and Jamaica to
Brazil.
a. Leaves mostly 3-foliolate, the leaflets elliptic, pinnately veined; corolla tube densely
puberulous outside; stamens more or less included; seeds thick, woody, wingless; Carib-
bean coast -1. T. jaroba
aa. Leaves 2-foliolate, the leaflets ovate to subrotund, conspicuously 3-veined from base;
corolla tube glabrate outside; stamens mostly exserted; seeds bialate with brownish
membranaceous wings; eastern Panama -2. T. nocturnum
1. Tanaecium jaroba Sw., Nov. Gen. Sp. P1. 92. 1788; Fl. Ind. Occ. 2: 1050,
tab. 20, fig 1. 1800. TYPE: Jamaica (not seen) .
T. albiflora DC., Prodr. 9: 245. 1845. TYPE: Based on T. jaroba Sw.

T. exsertum Griseb., Fl. Brit. W. Ind. Is. 450. TYPE: Jamaica, Mench 1070 (K).
Liana; branchlets terete, the nodes without pseudostipules, interpetiolar

glandular fields mostly lacking or inconspicuous, if present relatively small.
Leaves 3-foliolate or 2-foliolate with a tendril or tendril scar; leaflets elliptic,
acute or rounded, truncate or rounded at the base, 8-10 cm long and 4-6 cm
wide, pinnately veined with 5-7 lateral veins on a side, drying grayish-green,
the veins beneath beige, glabrous except for a few long trichomes at the extreme
base of the midvein above or puberulous with simple trichomes along the
veins beneath; tendril simple; petiole 5-6 cm long, petiolules 1-3 cm long,
grooved on the upper surface, pubescent with scattered long trichomes along
the groove, otherwise lepidote. Inflorescence a terminal raceme on a short
lateral branch; pedicels 1.3-2.4 cm long. Flowers with the calyx cupular, sub-
truncate or minutely 5-denticulate, more or less ciliate, glandular-spotted but
mostly glabrous with some plate-shaped glands on the apical part; corolla
white, elongate-tubular, expanded apically, 18-20 cm long and 1.2-1.7 cm wide
at the mouth, the tube 14-17 cm long, the lobes 2-2.8 cm long; densely glandu-
lar-puberulous outside, less densely glandular-lepidote and glandular-puberulous
on the lobes, more or less ciliate, tube mostly glabrous inside; stamens didyna-
mous, the anthers half-exserted, the thecae 5-6 mm long, the longer filaments
ca. 2.8 cm long, shorter filaments ca. 1.5 cm long, the staminode 5 mm long,
inserted 11-17 cm above the base of the corolla tube; pistil 16-17 cm long,
the ovary oblong, tetragonal, 1.5-2 mm long and ca. 1 mm wide, ovules ca.
6-seriate in each locule; disc cupular-pulvinate, 1.5-2 mm long and 2 mm wide.
Capsule elliptic-oblong, 14-22 cm long and 6-11 cm wide, 6-8 cm thick (Elias
1306 (US), Dugand & Jaramillo 3500A (US), both from Colombia), brown
with paler lenticellate flecks, glabrous or with a few lepidote scales, the valves
woody, convex; seeds angular, woody, wingless but with a sharp edge on 3
sides 2.5-3.3 cm long and 3.0-3.2 cm wide, the hilum terminal and at right
angles to the seed body.
No Panamanian collections of this species have been seen, but it is probably

a member of the flora of Panama. I have collected it in Costa Rica only a few
miles from the Panamanian border, and it is also known from Colombia.
Vegetatively it resembles Arrabidaea in having 3-foliolate leaves, a simple
tendril, an interpetiolar glandular field, and a stem cross section with 4 phloemn
arms. Its strongly raised venation on the leaf undersides, the small size of its
interpetiolar glandular fields, and the swampy habitat separate it from species
of Arrabidaea.
A plant of lowland swampy areas along the Caribbean coast, T. jaroba occurs
in regions of tropical moist forest but is probably restricted more by local edaphic
than by climatic factors. It ranges from Costa Rica to Venezuela and also to
Jamaica. It is only doubtfully distinct from the West Indian and Venezuelan
T. crucigerum Seem.
Presumably moth pollinated, it was collected in flower in Costa Rica in
late May. Its thick, wingless seeds are water-dispersed, correlating with its
swampy habitat and more or less circum-Caribbean distribution.
2. Tanaecium nocturnum (Barb. Rodr.) Bur. & K. Schum. in Mart. Fl. Bras.
8(2): 185. 1896.-FIG 37.
Osmohydrophora nocturna Barb. Rodr., Vellosia, ed. 2. 1: 49. 1891; 3: tab. 8-9. 1891. TYPE:
Brazil, Amazonas, Barbosa Rodriguez s.n. (not seen).
FIGURE 37. Tannaecium nocturnum (Barb. Rodr.) Bur. & K. Schum.-A. Habit (X 1/2).
[After Tyson et al. 4808, Bristan 1061, and Gentry 4543 (all MO).]-B. Fruit (X 1/2). [After
Gentry & Clewell 6926 (MO).]-C. Ovary and disc (X 1'/2).-D. Anther (X 21/2).-E.
Ovary cross section ( X 81/2). [After fresh material of Gentry 5233, Colombia.]
Liana to at least 5 cm in diameter, the bark smooth, pale gray; stem with
8(-16) arms in cross section; vegetative parts with a conspicuous almond odor;
branchlets subterete, finely longitudinally striate, elenticellate, lepidote, the
nodes without interpetiolar glandular fields or ridges; pseudostipules lacking.
Leaves 2-foliolate, sometimes with a tendril; leaflets ovate to subrotund, acumi-
nate to acute, rounded to slightly cordate, 8.8-16.0 cm long and 5-12.5 cm
wide, membranaceous, palmately veined at the base, the basal pair of lateral
nerves conspicuous, strongly arcuate, with axillary glandular fields beneath,
other secondary veins inconspicuous, mostly glabrous above and basally lepi-
dote, beneath glabrous to slightly lepidote; tendril simple, 7-18 cm long;
petiole and petiolules thinly lepidote, petiolules 2.5-7.7 cm long, apically glandu-
lar; petiole 2.7-7.5 cm long. Inflorescence a terminal racemose panicle, the
branches lepidote. Flowers fragrant, calyx tubular, more or less truncate, 16-18
mm long and 8-9 mm wide, scattered lepidote with glands; corolla white,
elongate-tubular, somewhat curved, 8.5-13.0 cm long and 0.7-1.6 cm wide at
the mouth, the tube 6.6-9.5 cm long, the lobes 1.2-2.4 cm long, glabrous to
sparsely glandular-lepidote outside, the lobes densely glandular-lepidote inside,
the tube glabrous; stamens exserted, anther thecae somewhat divergent, arcuate,
6-10 mm long, the filaments about equal, 4-5 cm long, the staminode 2-3 mm
long, inserted 4.7-7.2 cm from base of the corolla tube; pistil 7.8-9 cm long;
ovary elongate-oblong, lepidote, 6-7 mm long and 2 mm wide, the ovules 4-6-
seriate in each locule; disc pulvinate, 1 mm long and 3-4 mm wide. Capsule
ovoid-cylindrical, 12.7-20 cm long and 4-4.8 cm in diameter, 3.5-4 cm thick,
the surface impressed-lepidote, drying light brown; seeds 1.2-2.3 cm long and
4.4-7.5 cm wide, the wings membranaceous but brownish, irregular-edged, not
A vine of the tropical moist forest and tropical wet forest, it is known in
Central America only from Darien Province. To the south it ranges to Ama-
zonian Brazil. It was previously reported only from the drainage of the Ama-
zon and northeastern Colombia. It flowers during the early wet season from
May through July. Seeds are released during the dry season and are mostly
wind-dispersed; however the thickness of the central portion of the seed sug-
gests an intermediate stage in the evolution of water-dispersed seeds.
Tanaecium nocturnum is recognized by its pronounced, arcuate basal pair
of lateral veins, the almond odor, or by its smooth bark, which is reminiscent
of that of Carpinus caroliniana. The long, tubular, white flower of Tanaecium
is almost unique among the Bignonieae and strikingly convergent with the
flowers of Tabebuia stenocalyx and Ekmanianthe of the Tecomeae. The oblong
sausage-shaped fruit is also distinctive in the Bignonieae though clearly related
to the fruit of Ceratophytum. The superficial similarity of the fruit of Tanaecium
to that of Crescentia of the Crescentieae led to much confusion among early
taxonomists as to its tribal affiliation.
Vines of this genus contain enough cyanide (Dugand, Caldasia 1(5): 29-35.
1942) to be poisonous to livestock. The cut stems or crushed leaves of T. noc-
turnum give off a strong almond smell, and the plant is considered to have
aphrodisiacal properties by the Choco Indians. They maintain that a gift of
leaves of this plant to a girl will cause her to fall in love with the giver, or, in
a more complicated ritual, that when a handkerchief is scented with the juice
of this vine and presented to a girl, she will reciprocate the suitor's attentions
within three days.
DARIEN: Between Quebrada Venado and Peje Swamp on headwaters of Rio Tuqueza,
Bristan 1061 (MO, SCZ). Rio Pirre, Bristan 1478, 1501 (both MO). Rio Tuira between Rios
Paya and Aspave, Gentry 4399 (MO). El Real, Gentry 4543 (MO). Near Cerro Pirre on
Rio Parasenico, Gentry 4684 (MO). Rio Pirre near village of Pirre, Gentry & Clewell 6926
(MO). E of Santa Fe, Tyson et al. 4808 (MO, SCZ). Cana and vicinity, Williams 982 (NY).
38. TECOMA
Tecoma Juss., Gen. 139. 1789. LECTOTYPE: T. stans (L.) Juss. ex H.B.K.
Stenolobium D. Don, Edinburgh Phil. Jour. 9: 264. 1823. TYPE: S. castanifolium D. Don
= T. castanifolia (D. Don) Melch.
Kokoschkinia Turcz., Bull. Soc. Imp. Naturalistes Moscou 2: 33, tab. 2. 1849. TYPE: T.
gaudichaudi DC. (fide Index Kewensis).
Shrubs or small trees. Leaves simple, 3-foliolate or imparipinnately com-

pound, the leaflets serrate. Inflorescence a terminal raceme or racemose panicle.
Flowers with the calyx cupular, with 5 shallow, often apiculate, deltoid lobes,
corolla yellow or orangish, tubular-campanulate to narrowly tubular-infundib-
uliform, glabrous outside; stamens exserted or included, the anthers variously
divergent, glabrous or pilose; pollen grains single, oblong, 3-colpate, the exine
microreticulate; ovary narrowly cylindric, lepidote, the ovules 2-seriate in each
locule; disc cupular-pulvinate. Fruit a linear capsule, somewhat compressed
parallel to the septum but dehiscing perpendicularly to it, the valves smooth,
more or less glabrous; seeds thin, bialate, the wings hyaline-membranaceous,
About a dozen species ranging from extreme southern Arizona and southern
Florida through tropical America to northern Argentina, especially in the Andes.
1. Tecoma stans (L.) Juss. ex H.B.K., Nov. Gen. Sp. P1. 3: 144. 1819.-FIG. 38.
Bignonia stans L., Sp. P1., ed. 2. 2: 871. 1763. TYPE: (not seen).
Tecoma mollis H.B.K., Nov. Gen. Sp. P1. 3: 144. 1819. TYPE: Mexico, Guanaxuato, Humboldt
Tecoma incisa Sweet, Hort. Brit., ed. 1. 284. 1827, nomen nudum. TYPE: Trinidad (not seen).
Bignonia frutescens Mill. ex DC., Prodr. 9: 224. 1845, pro syn.
B. incisa Hort. ex DC., Prodr. 9: 224. 1845, pro syn.
Tecoma stans var. apiifolia Hort. ex DC., Prodr. 9: 224. 1845. TYPE: Guadeloupe, Bertero
s.n. (G-DC).
T. stans var. velutina DC., Prodr. 9: 224. 1845. SYNTYPES: Cultivated in Hort. Madrid, from
Mexico (G-DC); cultivated in Jard. Sarme (G-DC).
Stenolobium molle (H.B.K.) Seem., Jour. Bot. 1: 91. 1863.
S. stans (L.) Seem., Jour. Bot. 1: 88. 1863.
S. stans var. pinnata Seem., Jour. Bot. 1: 89. 1863. TYPE: Panama, Seemann 558 (BM).
S. stans var. apiifolium (DC.) Seem., Jour. Bot. 1: 89. 1863.
Gelseminum molle (H.B.K.) Kuntze, Rev. Gen. 2: 480. 1891.
G. stans (L.) Kuntze, Rev. Gen. 2: 479. 1891.
Stenolobium incisum Rose & Standl., Contr. U. S. Natl. Herb. 26: 174. 1913. TYPE: Mexico,
Chihuahua, Pringle 960 (US).
Tecoma stans var. angustatum Rehder, Mitt. Deutsch. Dendrol. Ges. 1915: 227. 1915. TYPE:
United States, Texas, El Paso, Jones 4187 (A).
Stenolobium quinquejugum Loes., Fedde Repert. 16: 211. 1919. TYPE: Mexico, El Calabaral,
Langlasse 478 (K).
S. tronadora Loes., Fedde Repert. 16: 210. 1919. LECTOTYPE: Mexico, Durango, Palmer
131 (MO).
BE
B It ll E~~~~/
FIGURE 38. Tecoma stans (L.) Juss. ex H.B.K.-A. Habit (x 1/2).-B. Corolla split
open ( x 1/2). C. Calyx and pistil ( x 1/2).-D. Ovary and disc ( x I1/2).-E. Anther (x 5).
-F Ovary cross section (x 121/2). [After Gentry 2895 (MO).]
Shrub or small tree to 10 m tall and 25 cm d.b.h., the bark dark and ridge
twigs lepidote and irregularly subpuberulent, more or less terete, drying dark
brown when young, light brown when mature. Leaves (in Panama) impari-
pinnately compound, 3-9-foliolate, the first pair of leaves on a new branch
often simple or 1-foliolate, the leaflets opposite, serrate, lanceolate, apically acute,
basally cuneate, the terminal leaflet often attenuate, 2.4-15 cm long and 0.8-6
cm wide, progressively larger distally, somewhat lepidote above and beneath,
simple puberulous on the midvein above and beneath, often puberulous beneath
at the base of the secondary nerves and sometimes on the leaf surface, espe-
cially in the vein axils; petiolules of the lateral leaflets lacking or to 2 mm long,
the terminal leaflet 4-20 mm long; petiole 1-9 cm long, rachis lacking (in
3-foliolate leaves), to 8 cm long, petiole and rachis slightly lepidote, puberulous
at the leaflet bases. Inflorescence a terminal or subterminal raceme of to 20
flowers, only a few flowers opening at a time; pedicels and inflorescence rachis
lepidote. Flowers fragrant with a sweet odor similar to vanilla; calyx elongate-
cupular, evenly 5-dentate, the teeth ca. 1 mm long and usually apiculate, 3-7
mm long and 3-4 mm wide, somewhat lepidote throughout, ciliate with con-
spicuous sunken submarginal glands; corolla yellow with 7 reddish lines in the
throat, 2 additional fainter red lines on top of throat ridges and a short faint
line at the base of the 2 upper lobes, tubular-campanulate above a narrowed
base 0.9-1.0 cm long, 3.5-5.8 cm long and 1.2-2.4 cm wide at the mouth, the
tube 3.0-4.3 cm long, the lobes 1.0-1.6 cm long, glabrous outside with a glandu-
lar epidermis, inside glabrous except for gland-tipped trichomes at the level
of stamen insertion and twisted trichomes in sinuses and on throat ridges;
stamens didynamous, the anther thecae divaricate, 3.5 mm long, pubescent
with twisted, simple, multicellular trichomes, the longer filaments 2.1-2.4 cm
long, shorter filaments 1.5-1.8 cm long, the staminode 4-5 mm long, inserted
8-9 mm from the base of the tube; pistil 3.1-3.4 cm long, ovary narrowly
cylindric, 3 mm long and 1 mm wide, glandular-lepidote, the ovules 2-seriate
in each locule; disc cupular-pulvinate, 1 mm long and 1 mm wide. Capsule
linear, tapering at the ends, subterete when fresh, 7-21 cm long and 5-7 mm
wide, surface lenticellate, more or less glabrous, sometimes slightly and incon-
spicuously lepidote; seeds 3-5 mm long and 2.4-2.7 cm wide, wings hyaline-
membranaceous, sharply demarcated from the seed body.
A characteristic plant of the premontane moist forest and also occurring

sparsely and locally in drier parts of the tropical moist forest, T. stans is culti-
vated frequently throughout the tropics and subtropics. Extremely wide-
ranging, it occurs naturally from southern Florida and southern Arizona through
the West Indies and Central America to northern Argentina.
This species flowers mostly during the early dry season from December to
February, also producing a few sporadic flowers throughout the year. Fruit
is produced mostly in the dry season.
Numerous varieties of this phenotypically plastic species have been named.
In Panama and Costa Rica the populations are relatively homogeneous, and
it is best to treat the plant as a single species without designation of varieties.
CANAL ZONE: Fort Kobbe, Allen 2019 (GH, MO, US). Along K-2 N of Cocoli, Croat
9171 (MO, SCZ). Culebra Island, Duke 4633 (MO, US), 4634 (MO). Madden Dam, Dwyer
3060 (MO). Summit of Cerro Luisa, Gentry 1402 (MO). Below Sosa Hill, Gentry 2895
(MO). Farfan beach road, Gentry & Dwyer 3547 (MO). SE of junction of K-2 with K-15
on K-2, Graham 315 (GH). Between Farfan Beach and Palo Seco, Hunter & Allen 442 (MO,
U, US); Lewis et al. 55 (GH, MO). Bella Vista, Salvoza 1009 (A). Balboa, Standley 30906
(US). Farfan Beach area, Tyson 1813 (MO, SCZ). Fort Amador on causeway and islands,
Tyson 2021 (MO, SCZ). Ancon below MARU research labs, Tyson 2798 (MO, SCZ, US).
La Boca, Williams 618 (US). Fort Grant, Woodson et al. 1552 (A, MO). CHIRIQUi: Boquete,
Sanchez 9 (MO, PMA). COCLEJ: Santa Clara Beach, D'Arcy & Croat 4096 (MO, SCZ);
Woodson et al. 1704 (A, MO). COLON: Santa Rita Trail, Cowell 107 (US). 2 km de la
carretera Transistmica, a mas o menos 1500 m del Rio Chagres, Rosas 31 (MO). DARIEN:
Vicinity of Vera Cruz, Duke 6062 (MO, SCZ). Isla Rosero in Golfo de San Miguel, Gentry
3983 (MO). Second large point just before Punta Alegre NE of Garachine, Gentry 4057 (MO).
LOS SANTOS: Playa de La Concepcion, Burch et al. 1259 (GH, MO, PMA, SCZ). Pedasi,
Dwyer 2488 (MO). Road from Macaracas to Chitre, Tyson et al. 3152 (MO, SCZ). PANAMA:
Ruins of Franciscan Convent, Celestine, 31 (US). Perlas Islands, San Jose Island, Correa 99
(SCZ); Duke 12527 (MO); Erlanson 75 (GH, US), 126 (GH); Johnston 547 (GH, MO);
Tyson & Loftin 5054 (CRU, MO, SCZ). Perlas Islands, Isla del Rey, Duke 9554 (MO).
Perlas Islands, Isla Espiritu Santo, Duke 10443 (MO). Facultad de Ciencias de la Universidad
de Panama', Escobar 27 (PMA). Vera Cruz, Lewis et al. 3007 (MO, SCZ). Vicinity of Panamai,
MacBride 2622 (US). Cerca de la Costa, Mejia 10 (MO, PMA). Bella Vista, Standley
25402 (A, US). In Chorrera, Tyson 5935 (SCZ). WITHOUT LOCALITY: Cuming 1096 (BM);
Seemann 558 (BM).
39. TECOMARIA
Tecomaria Spach, Hist. Nat. Veg. Phan. 9: 137. 1840. TYPE: T. capensis
(Thunb.) Spach.
Shrubs or subshrubs. Leaves imparipinnately compound, usually 9-11-

foliolate, the leaflets serrate. Inflorescence a raceme or racemose panicle.
Flowers with the calyx cupular, 5-toothed; corolla orange to scarlet, tubular,
curved; stamens exserted, the thecae more or less parallel, only slightly diver-
gent, attached to the filament ca. 1 mm from the fused end of the thecae. Fruit
a linear capsule, dehiscing perpendicularly to the septum; the seeds flat, mem-
branaceous-winged.
Two species of southern and eastern Africa, widely cultivated in the tropics
and subtropics.
1. Tecomaria capensis (Thunb.) Spach, Hist. Nat. Veg. Phan. 9: 137. 1840.
Bignonia capensis Thunb., Prodr. 105. 1794-1800. TYPE: (not seen).

Tecoma capensis (Thunb.) Lindl., Bot. Reg. tab. 1117. 1827.
Tecomaria krebsii Ki. in Peters, Reise Mossamb. Bot. 193. 1862-1864. TYPE: (not seen).
T. petersii Ki. in Peters, Reise Mossamb. Bot. 192. 1862-1864. TYPE: (not seen).
Shrub or subshrub, the branches flexuous. Leaves opposite, pinnately com-

pound, usually 7-11-foliolate; the leaflets elliptic to suborbicular, apically and
basally rounded or abruptly cuneate, sessile, ca. 1.5 cm long and 1.0 cm wide,
serrate, membranaceous, puberulous at least along main veins, the axils usually
with tufts of branched trichomes. Inflorescence a raceme or racemose panicle.
Flowers with the calyx cupular, 5-toothed, 5-7 mm long and 4-5 mm wide,
more or less puberulous, cilate with scattered plate-shaped glands in the upper
half; corolla orange to orange-scarlet, tubular, curved, 3.5-5 cm long and 0.6-
0.7 cm wide at the mouth, mostly glabrous, the lobes ciliate; stamens more
or less equal in length, anthers exserted, the thecae ca. 3 mm long, slightly
divergent for 1/2-%/. of their length, the slender filament attached ca. 1 mm
from the fused end of the thecae; pistil 5.5-6.5 cm long, the ovary oblong,
glabrous; disc cupular-pulvinate. Fruit rarely set in Central America, the
capsule linear, 5-11 cm long and 7-8 mm wide; seeds in two rows, thin, bialate
with hyaline membranaceous wings.
Native to South Africa, T. capensis is commonly cultivated in the subtropics

and at higher altitudes in the tropics. Reported by Standley as cultivated
frequently in Central America including Costa Rica, collections have not been
seen from Panama, but it is certainly to be expected. This plant is recognized
by its pinnately compound leaves and narrow tubular orange-red flowers with
strongly exserted anthers.
40. TOURRETTIA
Tourrettia Fougeroux, MWm. Acad. Paris 1784: 205. 1787; corr. Schreber Gen.
2: 406. 1791, orth. et nom. cons. TYPE: T. lappacea (L'Her.) Willd.
Dombeya L'Her., Stirp. Nov. 33. 1786, nom. rej. TYPE: D. lappacea L'Her. = T. lappacea
(L'Her.) Willd.
Medica Cothenius, Disp. Veg. 7. 1790, fide Index Kewensis.
Herbaceous vines; branchlets angled, nodes without interpetiolar glandular

fields; pseudostipules lacking. Leaves opposite, 2-3-ternate, often with a trifid,
dichotomously branching tendril, the leaflets serrate and often more or less
subdivided. Inflorescence a terminal spike-like raceme, the upper flowers sterile
and caducous. Flowers with the calyx red, bilobed, caducous; corolla tubular,
bilabiate, shorter than the calyx, pubescent at least toward the tip; stamens 4,
staminode absent, the anthers glabrous; pollen grains single, 3-colpate, the
exine finely granular; ovary ovoid, densely short-echinate; disc annular. Fruit
an uncompressed, ovoid, septicidally dehiscing capsule, usually not splitting to
the base, densely echinate with uncate spines; seeds small, thin, with a narrow,
hyaline-membranaceous wing.
One species of upland areas of continental tropical America ranging from

Guatemala to Argentina.
1. Tourrettia lappacea (L'Her.) Willd. in L., Sp. P1., ed. 4. 3: 263. 1801.-
FIG. 39.
Dombeya lappacea L'Her., Stirp. Nov. 33, pl. 17. 1785. TYPE: Peru, Pavon s.n. (BM).
Tourretia volubilis J. F. Gmel., Syst. Nat. 2: 940. 1791. TYPE: (not seen).
Herbaceous vine, the branchlets tetragonal. Leaves opposite, biternate with

a subterminal, finely branched, spirally coiling tendril, the leaflets membra-
naceous, glabrous, conspicuously and unevenly serrate, 3-8 cm long and 1.5-5
cm wide, the lateral pair of a ternation often divided to base. Inflorescence
FIGURE 39. Tourrettia lappacea (L'Her.) Willd.-A. Habit (X 3/5). [After White 23
and D'Arcy 6637 (both MO).]-B. Fruit (X 3/5). [After Woodson et al. 1116 (MO).]
A~~~~~~~~~~~~~~~~~~~~~~~~~~~
spicate but with short pedicels, narrowly bracteate, the bracts scarlet, the
terminal flowers mostly sterile with bright red calyces. Flowers with the calyx
of fertile flowers 1.3-1.5 cm long, somewhat hirsute, deeply 2-parted more than
half its length, the upper lobe long-acuminate, the lower lobe acute, the limbs
red; corolla purplish to greenish, tubular, 1.6-1.8 cm long and apically 3-5
cm wide, bilabiate, the tube 1-1.1 cm long, the lip 6-8 cm long, all lobes fused
forming a protective hood over the anthers, this open below, pubescent outside,
mostly glabrous inside, pubescent at the base of the stamens; stamens 4, the
anther thecae divaricate, glabrous, 1-2 mm long, shorter filaments 8-9 mm long,
longer filaments 9-11 mm long, staminode absent, inserted 7-8 mm from the
base of the tube; pistil 1.4-1.5 cm long, the ovary ovoid, densely short-tuber-
culate with thick, single-celled trichomes, 2.5-3 mm long and 1.5 mm wide,
4-celled, the ovules uniseriate in each locule; disc cupular-membranaceous,
1 mm long and 1.5 mm wide; capsule ovoid, somewhat woody, ca. 3.5 cm by
1.7 cm, strongly echinate with long spines to 1 cm long with recurved tips
and shorter straight spines, 2-valved, apically septicidally dehiscent, often
not splitting to base; seeds thin, flat, 6-7 mm long and 4-5 mm wide, the
dark brown body encircled by a narrow brown almost membranaceous wing.
This species is a plant of upper elevations from 600 m to 1800 m. It occurs
in the wet or moist premontane and lower montane regions from Guatemala
to Peru and in the Andes east to just across the Venezuelan border. The fruits
are burr-like and are probably epizoochorous with the seeds shaking a few at a
time out of the partially opened capsule.
The relationships of this monotypic genus are not clear. It is perhaps closer
to the Pedaliaceae than to the Bignoniaceae, but its compound leaves, tendrils,
and winged seeds have led to usual placement in the latter family.
CHIRIQUI: Trail from Paso Ancho to Monte Lirio in upper valley of Rio Chiriqui Viejo,
Allen 1477 (GH, MO). E side of Cerro Pando, D'Arcy 6637 (C, K, MO, P, UCWI). Cerro
Punta, Graham 275 (GH). Vicinity of Boquete, Lewis et al. 372 (GH, MO, US); Maurice
725 (US); Pittier 7001 (US). Rio Chiriqui Viejo valley near El Volca'n, White 23 (GGH, MO).
Finca Lerida to Boquete, Woodson et al. 1116 (A, MO, NY); Woodson & Schery 237 (GH,
MO, US). .
41. TYNNANTHUS
Tynnanthus Miers, Proc. Roy. Hort. Soc. 3: 193. 1863. LECTOTYPE: T. f

latus (Vell.) Miers.
Schizopsis Bur., Mon. Bign. 44. 1864. TYPE: S. labiata (Chain.) Bur. = Tynnanthus labiatus
(Chain.) Miers.
Lianas, perhaps small trees, stems with 4 phloem arms in cross section;
branchlets terete to subtetragonal without interpetiolar glandular fields, often
with a ridge between opposite petioles; pseudostipules lacking or foliaceous
but early deciduous. Leaves 2-3-foliolate, the terminal leaflet often replaced
by a simple or trifid tendril. Inflorescence an open or contracted axillary or
terminal panicle. Flowers with the calyx small, cupular, subtruncate, usually
5-denticulate or 5-setate; corolla white, often tiny, more or less infundibuliform.
bilabiately split about half its length, pubescent outside; anthers glabrous, sub-
exserted, the thecae divaricate, bent at the middle; pollen grains 3-colpate, the
exine smooth; ovary conical, densely pubescent, the ovules 2-4-seriate in each
locule. Fruit a more or less compressed linear capsule, the valves parallel to
the septum, flat with a slightly or conspicuously raised margin; seeds flat,
bialate with hyaline-membranaceous wings.
About 12 species of tropical America ranging from southern Mexico and the
West Indies to Brazil and Bolivia.
1. Tynnanthus croatianus A. Gentry, Ann. Missouri Bot. Gard. 58: 93. 1971.
TYPE: Panama, Canal Zone, Croat 11927 (MO, SCZ).-FIG. 40.
Liana, the stems with 4 phloem arms in cross section, the pith conspicuously
4-angled; branchlets tetragonal, becoming subterete, finely striate, becoming
noticeably lenticellate with age, young twigs more or less puberulous, the
nodes without interpetiolar glands or pseudostipules, vegetative parts with odor
of cloves. Leaves 3-foliolate, or 2-foliolate with a tendril or tendril scars; leaflets
membranaceous, papillose on both surfaces with scattered lepidote scales espe-
cially above, minutely puberulous below mostly on and near nerves and above
more or less glabrescently so over whole surface, drying light green to yellowish
with distinct yellow to reddish-brown reticulation beneath, darker green to
brownish above, the terminal leaflet when present broadly ovate, obtusely
acuminate, truncate to broadly cuneate, 5.5-12 cm long and 3.7-8 cm wide,
the lateral leaflets broadly ovate, obtusely acuminate, more or less asymmetrically
truncate or subcordate on one side at the base, 3.9-10 cm long and 2.7-6 cm
wide; terminal petiolule 1.5-3.3 cm long, laterals 0.6-2.5 cm long, the petiole
1.4-7.5 cm long, drying dark reddish; tendril strongly trifid; 11-15 cm long to
branching, the 3 arms 1.0-2.2 cm long. Inflorescence a small, usually axillary
panicle, 3-8 cm long and 2-6 cm wide, the branches strongly puberulous.
Flowers sweetly aromatic with odor similar to lilacs, calyx cupular, puberulous,
eglandular, truncate or minutely sub-denticulate, 3-4 mm long and 3-4 mm
wide; corolla white with 2 yellow lines in the throat; bilabiate, 1.2-2 cm long,
split about half its length, the 2 upper lobes almost fused, the 3 lower lobes
4-9 mm long, puberulous outside, inside pubescent on lower 3 lobes, margins
of upper 2 lobes, at the base of the lower side of the tube and sparsely
elsewhere, glabrous at the point of stamen insertion; stamens didynamous, the
anther thecae 1.5 mm long, divaricate, twisted near the base and reflexed
forward, the connective extended 0.3 mm beyond point of anther attachment,
the longer filaments 1.0-1.2 cm long, shorter filaments 0.9-1.0 cm long, the
staminode 0.5 cm long, inserted 2 mm from the base of corolla tube; pistil
16-17 mm long, the stigma small, bilamellate, 1 mm long, the style puberulous,
the ovary conical 1.5-2 mm long and 1 mm wide at the base, densely puber-
ulous, the ovules 4-seriate in each locule at the bottom of the ovary and 2-
seriate at the top; disc reduced, shortly cupuliform, densely pubescent on the
margin. Capsule linear, acuminate, flattened, 25-35 cm long and 0.8-1.2 cm
~~~
FiGuRE 40. Tynnanthus croatianus A. Gentry. A

(MO).] B. Fruit (X 1/2). [After Croat 12681 (MO)] C. Corolla split open (x 11/4).-
D. Ovary cross section (X 10). E. Anther (X 61/2). F-H. Front, back, and side views of
fresh corolla (X 1/2). I. Ovary (X 21/2). [After Gentry 5502 (MO).]
wide, 0.3-0.5 cm thick, the lateral margins somewhat raised, the corners sharply
angled, midvein not raised, sparsely to densely pubescent; seeds 0.6-0.8 cm
long and 1.7-2.9 cm wide, thin, bialate, the wings hyaline-membranaceous,
sharply demarcated from the brown seed body.
A plant of the tropical moist forest, T. croatianus is widely but sporadically

distributed through central and eastern Panama. It has been collected almost
to the Colombian border and probably occurs in that country as well. It flowers
during the wet season mostly in July and August. The seeds are wind-dispersed
and released during the dry season.
The trifid tendrils, subtetragonal young branchlets, 4-angled, almost cross-
shaped pith, and odor of cloves when cut are good recognition characters.
The fresh leaves are a glossy yellowish green in color quite similar to those of
Arrabidaea florida, but the trifid tendril is a sure distinguishing character.
The closest relative of this species is T. labiatus (Cham.) Miers, known only
from the Sellow type collection and a few other collections from eastern
Brazil. Although similar in general facies (calyx size and shape, corolla size,
leaf venation), T. labiatus differs in its lepidote rather than puberulous inflo-
rescence branches, glabrous leaves, flower color (described as "rosea," Blanchet
3267, Fl. Bras. 8(2): 193.), and in having the style glabrous except at extreme
base. The great range disjunction between these two species supports their
taxonomic separation.
CANAL ZONE: Barro Colorado Island, Croat 11927, 13975 (both MO, SCZ); Foster 1178
(MO); Gentry 708 (MO); Shattuck 1108 (F, MO, US); Starry 149 (F, MO). Boy Scout
Camp Road, Croat 12904 (MO); Gentry 5502 (MO). DARIEN: Rio Balsas between Manene
and Tusijuanda, Duke 13544, 13579 (both MO). El Real, Gentry 4538 (MO). PANAMA:
Alhajuela, Dwyer 1144 (MO, SCZ). Rio Pasiga, Gentry 2203, 2271, 2365 (all MO). 0-4
km from Rio Bayano crossing on road to Santa Fe, Gentry 3875 (MO). Near archeological
site at edge of Madden Lake, Gentry & Tyson 5024 (MO).
42. XYLOPHRAGMA
Xylophragma Sprague in Hook., Ic. P1. tab. 2770. 1903. TYPE: X. pratense
(Bur. & K. Schum.) Sprague.
conspicuous interpetiolar glandular fields; pseudostipules triangular to ovate,
short, acute. Leaves 3-foliolate, the terminal leaflet often replaced by a simple
tendril. Inflorescence a narrow-bracteate, contracted, racemose panicle, axillary
on older branchlets. Flowers with the calyx cupular, truncate, conspicuously
5-denticulate, usually stellate or dendroid pubescent; corolla lavender, more
or less tubular-campanulate, dendroid-puberulous without; anthers glabrous, the
thecae straight, divaricate; pollen grains 3-colpate, the exine almost smooth;
ovary flattened ovoid, densely lepidote, the ovules 6-8-seriate in each locule;
disc cupular-pulvinate. Fruit a compressed, woody, ellipsoid capsule, the valves
parallel to the septum, more or less smooth or with slightly raised glandular
areas, the median nerve not evident, the seeds thin, the wings hyaline-mem-
branaceous, sharply demarcated from the seed body.
Four species of continental tropical America ranging from Mexico to Bolivia,

Paraguay, and Brazil.
1. Xylophragma seemannianum (Kuntze) Sandw., Kew Bull. 1953: 469.

1954.-FIG. 41.
Bignonia dentata DC. sensu Seem., Bot. Voy. Herald 179. 1854, non DC.
Saldanhaea seemanniana Kuntze, Rev. Gen. P1. 2: 480. 1891. TYPE: (not seen).
Distictis rovirosana Donn. Sm., Bot. Gaz. (Crawfordsville) 20: 7. 1895. TYPE: Mexico,
Tabasco, Rovirosa 488 (K, US).
Tecoma floccosa Ki. ex Bur. & K. Schumn. in Mart., Fl. Bras. 8(2): 318. 1897. TYPE: British
Guiana, Schomburgk 334 (B, not extant; K, photo).
Adenocalymma cocleense Pittier, Contr. U. S. NatI. Herb. 18: 255. 1917. TYPE: Panama,
Cocle, Williams 522 (NY, US).
Tabebuia floccosa (KI. ex Bur. & K. Schum.) Sprague & Sandw., Kew Bull. 1932: 27. 1932.
Liana to at least 7 cm in diameter, often shrubby when young, the bark

relatively smooth with conspicuous raised whitish lenticels, stem with 4 phloem
arms in cross section; branchlets subtetragonal to subterete, more or less pubes-
cent with dendroid trichomes, glabrescent, the nodes with iTterpetiolar glandular
fields; pseudostipules inconspicuous or resembling minute bromeliads. Leaves
3-foliolate or 2-foliolate with a tendril or tendril scar (some leaves 5-foliolate
in Gentry 1112, Costa Rica (MO)); leaflets rhombic-ovate to rhombic-obovate,
acuminate to acute, truncate to broadly cuneate, 3.5-28 cm long and 1.5-14
cm wide, membranaceous to chartaceous, palmately 3-5-nerved from the base,
secondary veins 5-7 on a side, variably lepidote and pubescent above and
beneath, beneath with mostly dendroid trichomes, above often with simple tri-
chomes over most of surface and dendroid trichomes on main veins, drying
variably olive or brownish; tendril simple, 7-17 cm long; petiolules and petiole
dendroid-pubescent, the terminal petiolule 2.1-7 cm long, the lateral petiolules
1.0-2.6 cm long, the petiole 6.4-12 cm long. Inflorescence a contracted, lateral
raceme or panicle, each flower subtended by 2 narrow bracts, the branches
dendroid-pubescent. Flowers fragrant with a sweet odor with musky over-
tones; calyx cupular, 5-9 mm long and 3-5 mm wide, with 5 slender teeth, 1
mm long, pubescent with stellate or dendroid trichomes, especially dense in
S lines terminating in the marginal teeth, eglandular; corolla lavender outside
and on the lobes, the throat white inside, tubular-campanulate, 3.5-6.2 cm
long and 1.2-2.0 cm wide at the mouth, the tube 2.2-3.6 cm long, its lower
6-9 mm narrowed to 2 mm wide, the lobes 1.2-2.2 cm long, thickly dendroid
pubescent especially in bud outside and on its lobes inside, the tube within
mostly glabrous with simple trichomes at and just below the level of stamen
insertion; stamens didynamous, the anther thecae divaricate, 2.3-3.5 mm lon
the longer filaments 1.6-2.1 cm long, shorter filaments 0.9-1.5 cm long, the
staminode 1.5-3 mm long, inserted 7-10 mm from the base of the corolla tube;
pistil 2.1-2.8 cm long, ovary tapered-cylindrical and somewhat flattened, 1.5-2.5
mm long and 1.5-2 mm wide, densely lepidote, the ovules 6-8-seriate in each
locule; disc cupular-pulvinate, 0.5-1 mm long and 1.5-2 mm wide. Capsule
rounded-oblong, flattened, 5.5-16 cm long and 3.4-5 cm wide, 1-1.5 cm thick,
drying dark, inconspicuously lepidote, often with raised glandular areas; seeds
; \ ~~~~~~~~B
FIGuRE 41. Xylophragma seemannianum (Kuntze) Sandw.-A. Habit (x 1/2). [After

Porter et al. 5150 (MO).]-B. Leaf (X 1/2). [After Gentry 3776 (MO).]-C. Fruit (x 1/2).
[After Gentry 4965 (MO)] D. Corolla split open (x '/2). E. Ovary and disc (X 3).-
F. Ovary cross section ( X 9). [After Gentry 3890 (MO).]
1.6-2.9 cm long and 3.0-5.2 cm wide, the wings hyaline-membranaceous, sharply

demarcated from the seed body.
A common vine of tropical dry forest, premontane moist forest, and drier
parts of the tropical moist forest, Xylophragma seemannianum becomes pro-
gressively less common in wetter parts of the tropical moist forest. It ranges
from Mexico to Guyana. This species flowers during late dry season from
March to May. The fruits take about a year to mature, releasing seeds during
the next dry season.
Although first described as a species of Saldanhaea, this plant is quite dis-
tinct from that genus, which I have united with Cuspidaria. The fruit is
different from the relatively narrow, elongate fruit of Cuspidaria and lacks any
hint of raised margins or submarginal area. This species also lacks the distinctive
bent anther thecae and pollen tetrads of Cuspidaria (including Saldanhaea).
Xylophragma seemannianum is distinguished from other species with which
it might be confused by the dendroid trichomes of its leaves and petioles. In
the field this pubescence assumes a macroscopically characteristic mealy form.
The combination of interpetiolar glandular fields and relatively large leaves
also serves to mark this species.
The generic name, meaning woody partition, refers to the woody capsule
valves and comes from the Greek. It is neuter and the specific epithet seeman-
nianum thus takes the neuter ending used here rather than the feminine gender
given it by Sandwith (Kew Bull. 1953:469. 1954).
CANAL ZONE: Barro Colorado Island, Croat 5029 (SCZ), Croat 13962 (MO, SCZ); Foster
705, 798 (both PMA). Victoria Fill near Miraflores Locks, Allen 1755 (F, GH, MO, US).
Road C25B, E of Madden Forest, Croat 15265 (MO). Pipeline Road, Croat 9327 (MO,
SCZ); Gentry 1582, 4857 (both MO). Curundu, Gentry 1601 (MO). Boy Scout Camp road
near Madden Lake, Gentry 2051 (MO). Farfan beach road, Gentry 4903 (MO). N of
Paraliso, Gentry 4960 (MO). C-29 beyond Summit Gardens, Harvey 5286 (F). Along Las
Cruces Trail in Madden Forest, Lewis et al. 5304 (MO, NY, SCZ). Gamboa district, Maggs
II-44 (F, K). Gorgona, Seemann 603 (BM, K). CHIRIQUI: Remedios, Lindsay 468 (F).
COCLE: Penonome and vicinity, Williams 522 (NY, US). DARIEN: Cerro Piriaque, Duke
8115 (MO). Rio Tuira between Rios Punusa and Mangle, Duke 14583 (MO). Santa Fe,
Gentry 4310 (MO). Rio Tuira between Rios Pucuro and Paya, Gentry 4329 (MO). Near
village of Paya, Gentry 4396 (MO). El Real, Gentry 4547 (MO). Trail from Cerro Pirre
to El Real, Gentry 4783 (MO). Cerro Piriaque, Tyson et al. 3824 (MO, SCZ). HERRERA:
E of Las Minas, Gentry 3140 (MO). PANAMA: Perlas Islands, Isla de La Bayonetta,
Dwyer 1756 (MO). Near Capira, Allen 730 (CGH, MO, US). Dam site on Rio Bayano,
Gentry & Tyson 1663A (MO). Rio Maestro, Gentry 2225 (MO). Rio Pasiga, Gentry
2261, 2262, 2264, 2276 (all MO). Rio Piragua (Tabardi ), Gentry 2575 (MO). Icanti,
Gentry 2608 (MO). Near Rio Espave, Gentry 3739 (MO). Trail from Rio Espave toward
Rio Agua Clara, Gentry 3776 (MO). 10-15 km from Rio Bayano crossing on trail to Santa
Fe, Gentry 3822, 3886 (both MO). Rio Espave near junction with Rio Bayano, Gentry 3890
(MO). W of Cafiitas, Gentry 4965 (MO). E of Cafiitas, Gentry 4975 (MO). Between Chepo
and El Llano, Gentry & Tyson 1638, 1688 (both MO). Rio Cabra, Maurice 764 (US).
Between Rio Pacora and Chepo, Porter et al. 5150 (MO, SCZ). E of Rio Tocumen, Standley
26589 (GH, US). Vicinity of Capira, Woodson et al. 1310 (MO). VERAGUAS: S of Santa
Fe, Gentry 2935 (MO).
INDEX OF LATIN NAMES
Numbers in boldface type refer to descriptions; numbers in roman type refer to synonyms;
numbers with dagger (f) refer to names incidentally mentioned.
Adenocalymma 786 comosum 786 f

alboviolaceum 889 densiflora 896
apurense 787, 789T, 790T, 813k fissum 885
arthropetiolatum 789, 790i floribundum 833
brachybotrys 890 flos-ardeae 939
calderonii 787, 789k foveolatum 890
ciliolatum 888 friesiana 811
cocleense 968 grenadense 787
helicocalyx 885 Arrabidaea 803, 8101, 8151, 8181, 9211,

heterophyllum 823 9331, 9551
hintonii 787, 789k -ser. Macrocalycinae 9331
hosmeca 888 acuminata 811
inundatum 787, 789k actifolia 807
japurense 931 amazonica 809
kerere 890 barquisimetensis 811
laevigatum 888 belizensis 931
macrocarpum 888 blanchetii 818
magdalenense 790k broadwayi 817, 8181
micradenium 885 calodictyos 806, 807f
moritzii 885 candicans 804, 809f
obovatum 888 -var. latifolia 804
ocositense 826 carabobensis 819
pachypus 888 cardenasii 812
paniculatum 809 carichanensis 8031
perezii 885 chica 8071, 807
pohlianum 888 chironeura 809
portoricensis 808 conjugata 809
punctifolium 939 corallina 810, 8151
seleri 885 corymbifera 868
standleyanum 823 costaricensis 815f
stridula 890 cuprea 808
symmetricum 89,0 dischasia 896
verruciferum 885 dichotoma 8101
verrucosum 931 divaricata 812
Aegiphila duckei 877
congesta 928 egensis 933 f
Amphilophium 791, 912k floribunda 8331
macrophyllum 791, 793f, 795k florida 812, 8161, 9671
molle 791, 793f grosourdyana 8031
mutisii 791 guaricensis 811, 814
oxylophium 794 guatemalensis 837
paniculatum 791, 793f, 794f, 795k inaequalis 9101
-var. molle 791, 793f isthmica 814, 837
pannosum 794, 795f japurensis 931
vauthieri 791 larensis 808
xerophilum 791 lenticellosa 811, 931
Amphitecna 842 leucopogon 803f
macrophylla 842 f littoralis 814, 8151
obovata 846 lundellii 819
sessilifolia 848 macrocarpa 833
Anemopaegma 796 magnifica 9191, 920
belizeanum 798 mollicoma 814
carrerense 803k mollissima 8121, 814, 8671
chamberlaynii 803k nicotianaeflora 931
chrysanthum 796, 802i obliqua 810
chrysoleucum 798, 799f, 800k -var. hirsuta 811
flavum 800k obovata 810
grandiflorum 799k oligantha 931
grandifolia 796k orbignyana 806
karstenii 803k ovalifolia 811
lehmanii 802i oxycarpa 815
leptosiphon 877 pachycalyx 806
macrocarpa 798 panamensis 812, 8131
mirandum 796k patellifera 8151, 816
orbiculatum 796f, 800 pentstemonoides 816, 8181
pachypus 888 pleei 819
paraense 799k praecox 811
platypus 888k purinosa 806
puberulum 801, 802k pseudochica 837
punctulatum 798 pubescens 8071, 8151, 818
santa-ritense 802, 803k rego 803f
surinamense 799 f rhodantha 811
symmetricum 800k -var. oxyphylla 811
tobagense 823 rhodothyrsus 806
tonduzianum 837 rosea 807
vargasianum 839 rotundata 810
rubrinervis 833 heterotricha 948

samydoides 803f, 818k hexagona 910
sanctae-marthae 814 hibiscifolia 810
sceptrum 803 f hondensis 820
schomburgkii 806 hostmannii 836
seleriana 814 hymenaea 888
sieberi 819 ignea 918
spraguei 811 inaequilatera 936
subfastigiata 833 incarnata 836, 890
subincana 806 incisa 958
virescens 809, 810k inflata 874
-var. farinosa 809 kerere 890
zulianensis 806 laevigata 888
Avicennia 873k languinosa 874
Batocydia latifolia 820
exoleta 874 laurifolia 896
unguis 874 lenta 895f, 895f, 896
Bayonia 884 lepidota 840
fissa 885 littoralis 814
helicocalyx 884T, 885 longisiliqua 877
Bignonia magnifica 919k
acapulcensis 814 magnoliaefolia 850
acuminata 811 martinii 877
acutistipula 874 martiusiana 896
adenophylla 856 mollissima 814
aequinoctialis 836 muricata 910
aesculifolia 853 nitida 812
alliacea 850 nitidissima 836
apurensis 787 noterophila 826
balbisiana 810 obliqua 810
benensis 890 obovata 877
binata 826 orbiculatum 800
boliviana 811 pachyptera 871
brevipes 939 pallida 950
californica 874 paniculata 791
callistegioides 826 pannosa 794, 795f
candicans 804 patellifera 816
capensis 961 pentaphylla 951k
chica 807 phaseoloides 910
chrysantha 941, 942k picta 836
chrysoleuca 798 populifolia 880
ciliata 809 prieurei 882
coito 905 procera 862
colombiana 810 pseudounguis 874
colorata 804 pterocarpa 833f
conjugata 809, 810k pubescens 794, 818
copaia 862 purpurea 826
corallina 810 pyramidata 895
corymbifera 868 quadrivalvis 880
crucifera 820 riparia 938
cuprea 807 rotundata 810
cupulata 816 rupestris 896
dasyonyx 874 rusbyi 850
dentata 968 sagreana 839
dichotoma 810 sarmentosa 836
diversifolia 839 -var. hirtella 836
echinata 910 sinclairii 896
ehretioides 896 spectabilis 836
erubescens 808 squalus 910
exoleta 874 stans 958
farinosa 809 striata 896
floribunda 833k tecomaeflora 918
fockeana 877 tetragonolobum 823
frutescens 958 tiliaefolia 910
glabrata 810 triantha 874
gracilis 874 triphylla 807
heptaphylla 945, 947k tweediana 874
heterophylla 890 umbellata 826
umbrosa 868 -var. puberula 832

uncata 871 cuneifolia 832
uncinata 871 cuspidata 846
unguiculata 874 edulis 899, 900
unguis 874 fasciculata 832
-var. gracilis 874 latifolia 846
unguis-cati 874 lethifera 846
-var. exoleta 874 musaecarpa 900
-var. guatemalensis 874 obovata 846
-var. serrata 874 palustris 846
variabilis 836, 913 pinnata 864
venusta 918 plectantha 832
verrucifera 885, 887k spathulata 832
villosa 810, 814, 836 ternata 829
Bignoniaceae 781, 799f, 855f, 866T, 912f, toxicaria 846
923f trifolia 829
Bignonieae 781f, 790f, 801f, 957k Crescentieae 781f, 902f, 957k
Blepharitheca 833 Cupulissa 796
floribunda 833f, 833 grandifolia 796k
Bombacopsis 952f Cuspidaria 833, 970k
Callichlamys 819, 822f, 884k corymbifera 868
garnieri 820, 822f floribunda 833
latifolia 819f, 820 hibiscifolia 810
peckoltii 820 mollis 817
riparia 819f, 820 pterocarpa 833 f
rubiginosa 820 Cybistax
rufinervis 820 macrocarpa 853
splendida 820 Cydista 835, 921i
Carpinus aequinoctialis 799f, 828f, 835f, 836, 896T,
caroliniana 957 f 907f
Cedrela 952 f -var. hirtella 835f, 838f
Ceratophytum 823, 825f, 894f, 957k amoena 836
brachycarpum 823 diversifolia 839, 884k
capricorne 823f, 823 heterophylla 840
tetragonolobum 823, 898k incarnata 836
tobagense 823 pubescens 837, 838k
Chasmia 803 sarmentosa 836, 838f
Chodanthus seemannii 836
puberulus 801 spectabilis 836
Citharexylum vargasiana 838
ramizii 927 Delostoma
Clytostoma 826, 838f latifolia 820
binatum 815f, 826 Dendrosicus 842
callistegioides 826k isthmicus 843, 846f, 849k
elegans 828 kennedyi 844, 849k
floridum 826 latifolius 842f, 844f, 846, 848k
isthmicum 826 macrophyllus 842f, 846f, 849k
noterophilum 826 saxatilis 842f, 846
ocositense 828 sessilifolius 844f, 847f, 848
purpureum 826 silvicola 849k
Couralia 940 spathicalyx 848f, 849
fluviatilis 940k Dermatocalyx 923
rosea 951 parviflorus 923f, 927
Craterotecoma 866 Distictella 850, 852f
parviflora 866f broadwayana 850
Cremastus 803 kochii 850
balbisianus 810 magnoliifolia 850
glutinosa 803f mansoana 850k
rufo-villosus 936 racemosa 850, 852k
Crescentia 782f, 829, 923f, 957k Distictis 850
aculeata 900 kochii 850
acuminata 831 racemosa 850
alata 829, 847f, 902f rovirosana 968
angustifolia 832 Dolichandra
arborea 831 fenzliana 871
coriacea 846 Dombeya 962
cucurbitina 846 lappacea 962f, 962
cujete 829f, 831 Doxantha 870, 882k
acutistipula 874 colombiana 869

adunca 874 corymbifera 868, 870k
dasyonyx 874 dicheiolocalyx 869
exoleta 874 erionema 866f
lanuginosa 874 glabra 866
mexicana 874 phaseolifolia 868
praesignis 874 puberula 869
serrulata 874 schomburgkii 820
tenuicula 874 schumanniana 869
unguiculata 874 truncata 868
unguis 874 umbrosa 868
unguis-cati 870f, 874 valenzuelae 868, 869f
-var. dasyonyx 874 Macfadyena 870, 881f, 882f
-var. exoleta 874 amazonica 905
Ekmanianthe 957k bangii 905
Enallagma 842 coito 905
cucurbitina 842T, 846 corymbosa 905
latifolia 844f, 846 fenzliana 871
obovata 846 fockeana 877
sessilifolia 848 guatemalensis 871
Endoloma 791 hispida 871, 873k
purpurea 791f, 791 laurifolia 905
Etorloba 856 lepidota 905
Gelseminum mollis 871, 873k
avellanedae 945 obovata 877
molle 958 orinocensis 905
stans 958 ovata 905
Gibsoniothamnus 923k phellosperma 908
Godmania 853 platypoda 905
aesculifolia 853 pubescens 871
macrocarpa 853f, 853 uncata 870f, 871, 876k
uleana 853 uncinata 870f, 871
Handroanthus 940 unguis-cati 870f, 871f, 873f, 873, 882k
albus 940k Martinella 876
Haplophragma 855 iquitosensis 877
adenophyllum 855f, 856 martinii 876f, 877
Heterophragma obovata 876f, 877
adenophyllum 856 verrucosa 931
Iacaranda 856 Medica 962
Jacaranda 856, 863k Melanconiales 925k
acutifolia 864k Melloa 879, 882f
caerulea 856k populifolia 879f, 880
caucana 858, 863k quadrivalvis 871f, 879f, 880
-subsp. sandwithiana 858 Microbignonia 870
chelonia 863 auristellae 870k
copaia 862 Mussatia 882, 884k
filicifolia 858 hyacinthina 882
glabra 856k prieurei 882 f
gualanday 858 Neotuerckheimia
hesperia 860f, 861f gonoclada 848
mimosifolia 863 Onohualcoa 884
orbiculata 800 fissa 885
ovalifolia 863, 864k helicocalyx 885, 887k
procera 862 seleri 884f, 885
spectabilis 862 verrucifera 884f, 885
superba 862 Osmohydrophora 954
trianae 858 nocturna 954f, 955
Kigelia 864 Pachyptera 887, 894k
pinnata 864 alliacea 815f, 895f
Kokoschkinia 958 dasyantha 896
gaudichaudi 958k foveolata 887f, 890
Kordelestris 856 hymenaea 887f, 888, 895k
Laguncularia 873k kerere 887T, 890
Levya 835 -var. erythraea 890, 892f
nicaraguensis 835f, 836 -var. incarnata 892f
Lochmocydia 833 parvifolia 893
Lundia 866 perrottetii 896
chica 807 puberula 871
standleyi 894 Platolaria 796

striata 896 flavescens 796f, 800
umbelliformis 896 Pleonotoma 913
Pandorea diversifolium 839
ricolasiana 916 jasminifolia 913k
Panterpa 803 variabilis 913
leucopogon 803 f Podranea 916
mollissima 814 ricolasiana 916
Paragonia 825f, 895, 898f Pseudocalymma 887
pyramidata 895 alliaceum
schumanniana 931 -var. macrocalyx 894
Paramansoa 803 -var. microcalyx 889
grosourdyana 803k hymenaeum 889
Parmentiera 899, 902f laevigatum 887f, 889
aculeata 899 langlasseanum 889
alata 829 macrocarpum 889
cereifera 900, 902f, 904k pachypus 889
edulis 899f, 900 pohlianum 889
foliolosa 900 standleyi 894
lanceolata 900 Pteromischus 829
macrophylla 902 alatus 829f, 829
trunciflora 904f Pteropodium 856
Pedaliaceae 964 glabrum 856k
Peltospermum Pyrostegia 918
orbiculatum 800 acuminata 918
Pentelesia 803 ignea 918f, 918
discolor 803f intaminata 918
Periarrabidaea pallida 918
duckei 877 parvifolia 918
Petastoma 803 reticulata 918
breviflorum 817, 818k venusta 918
broadwayi 817 Rafinesquia 856
domatiatum 817, 818k caerulea 856 f
langlasseanum 889 Rhizophora 873k
laurifolium 826 Romeroa
leiophyllum 896 verticillata 954k
macrocalyx 896 Saldanha 833k
multiglandulosum 826 Saldanhaea 833, 970k
ocositense 826 floribunda 833
patelliferum 819 lateriflora 833 f
pentstemonoides 819 seemanniana 968
pubescens 819 subfastigiata 833
samydoides 803k Saritaea 919, 921k
tonduzianum 889 magnifica 920
Phoenicocissus 866 Schizolobium
Phryganocydia 904, 909f, 921k parahybum 863k
coito 905 Schizopsis 964
corymbosa 904f, 905, 907f, 909k labiata 964k
dipleuropus 905 Schlegelia 923, 925k
japurensis 931 darienensis 924f, 930O
orinocensis 905 dressleri 923f, 924, 927f, 928k
phellosperma 908 fastigiata 929, 930O
pisoniana 880 fuscata 925, 928f, 929f
Phrygiobureaua 904 gentlei 928
Piriadacus lilacina 923 f
hibiscifolius 811 macrophylla 928
Pithecoctenium 909 monachinoi 929k
aubletii 910 nicaraguensis 926
cordifolium 910 organensis 928
echinatum 794T, 910 parasitica 927k
glaucum 910 parviflora 923f, 926f, 927
hexagonum 910 ramizii 928
lundii 910 -var. costaricensis 928
muricatum 910 -var. macrandra 928
panamense 800 -var. trichandra 928
phaseoloides 910 roseiflora 928
reticulare 896 sulfurea 929
squalus 910 violacea 923k
tribrachiatum 910 Scrophulariaceae 923k
Scobinaria 930, 933k mansoana 820, 820f

amethystina 931 mexicana 951
japurensis 930f, 931 micrantha 928
verrucosa 930f, 931 neochrysantha 948, 9491f
Setilobus neurophylla 816
vicentinus 885 nicaraguensis 946, 947f
Sparattosperma ochracea 947
rosea 951 -subsp. neochrysantha 948
Spathodea 880f, 935 pallida 950
adenophylla 856 palmeri 945, 947f
campanulata 935 palustris 950
coito 905 pentaphylla 951, 952k
corymbosa 905 pisoniana 880
fraxinifolia 880 punctatissima 951
hispida 871 pyramidata 896
laurifolia 905 rosea 950f, 951
mollis 871 rufescens 941, 942k
obovata 877 rufinervis 820
orinocensis 905 schunkevigoi 946
pisoniana 880, 880f serratifolia 945k
platypoda 905 sessilifolia 848
uncata 871 speciosa 820
uncinata 871 stenocalyx 953f, 957k
Spirotecoma 855 f striata 953
Stenolobium 958 uliginosa 940f
castanifolium 958k Tanaecium 954, 957f
incisum 958 albiflora 9154
molle 958 crucigerum 955 f
quinquejugum 958 exsertum 954
stans 958 jaroba 954
-var. apiifolium 958 nocturnum 954T, 955
-var. pinnata 958 pinnatum 864
tronadora 958 zetekii 890
Stizophyllum 935 Tecoma 958
denticulatum 935f, 936k adenophylla 945
flos-ardeae 939 aesculifolia 853
inaequilaterum 936 avellanedae 946
perforatum 935f, 936k -var. alba 946
punctifolium 939 capensis 961
riparium 938 castanifolia 958k
Swietenia 952i chrysantha 941
Tabebuia 909f, 940 digitata 853
adenophylla 947k evenia 941, 951
aesculifolia 853 eximia 945
alba 940f floccosa 968
aquatilis 940f, 951k fuscata 853
avellanedae 945 guayacan 944
-var. paulensis 946 hassleri 948
calderonii 787 heptaphylla 945, 947k
cassinoides 940k heterotricha 948
chapadensis 810 hypodictyon 948
chrysantha 941, 948, 949k impetiginosa 945
cordata 877 -var. lepidota 945
dugandii 946, 947k incisa 958
eximia 946 integra 946
floccosa 968 ipe 945
fuscata 853 -var. integra 945
glomerata 941 mexicana 951
guayacan 944 mollis 958
heptaphylla 946, 947f ochracea 947, 949k
heterotricha 948, 949k palmeri 941
hypodictyon 948 pisoniana 880
impetiginosa 815f, 945 punctatissima 951
-var. lepidota 947k ricolasiana 916
insignis 951f, 953k rosea 951
ipe 946, 947k stans 958
-var. integra 945 -var. angustatum 958
japurensis 931 -var. apiifolia 958
latifolia 820, 877 -var. velutina 958
Tecomaria 961 Tynnanthus 964

capensis 961 croatianus 813i, 965
krebsii 961 fasciculatus 964k
petersii 961 hyacinthinus 882
Tecomeae 781f, 957k labiatus 964f, 967k
Temnocydia Vasconcellia 803
aequinoctialis 836 acutifolia 807
carajura 807 florida 812
elliptica 896 Xylophragma 967
lenta 896 pratense 967f
spectabilis 836 seemannianum 812i, 835i, 968
Tourrettia 962 Zenkeria 899
lappacea 962 Zeyheria
volubilis 962 surinamensis 896

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Flora of Panama. Part IX. Family 172.

FAMILY 172. BIGNONIACEAE2

1Assisted by National Science Foundation Grant GB-5674 (Thomas B. Croat, principal

ANN. MISSOURi BOT. GARD. 60: 781-977. 1973.

ous and provide useful taxonomic characters. Presence or absence of concen-

q. Leaves 2-3-ternate; branchlets acutely tetragonal and the tendril trifid

MM. Corolla puberulous or lepidote outside at least on lobes;

Adenocalymma Mart. ex Meisn., Gen. 1: 300; 2: 208. 1840. LECTOTYPE: A.

1. Adenocalymma apurense (H.B.K.) Sandw., Lilloa 3: 461. 1938.-FIG. 1A-D.

Liana to 10 cm in diameter, the bark smooth, gray, often 4-furrowed; twigs

FIGURE 1. Adenocalymma.-A-D. A. apurense (H.B.K.) Sandw.-A. Habit (X %).

2. Adenocalymma arthropetiolatum A. Gentry.4 TYPE: Panama, Canal Zone,

Liana to 3 cm in diameter, the bark smooth, greenish, with inconspicuous

on the surface, with q. few scattered plate-shaped glands; tendril simple, to 14 cm

a. Pseudostipules persistent and conspicuous; capsule compressed, rugose-tuberculate; vege-

Liana to at least 10 cm in diameter, the bark loose, fibrous, vertically striate;

FIGURE 2. Amphilophium paniculatum (L.) H.B.K.-A. Habit ( X%//). [After Gentry

angles, lepidote to strongly dendroid pubescent, especially on the ribs, interpetio-

Ecologically this is the most wide-ranging species of Central American Big-

Liana to at least 2.5 cm in diameter, branchlets sharply hexagonal with the

About 30 species; continental tropical America from Brazil and Argentina to

1. Anemopaegma chrysanthum Dugand, Caldasia 4: 307. 1947. TYPE: Colom-

FIGURE 3. Anemopaegma.-A-B. A. orbiculatum (Jacq.) DC.-A. Habit (x '/2).

2. Anemopaegma chrysoleucum (H.B.K.) Sandw., Lilloa 3: 459. 1938.-FIG.

Liana to at least 2.5 cm in diameter, usually less than 1 cm in diameter, the

(rarely 3-foliolate), sometimes cirrhose; leaflets narrowly elliptic, acute, cuneate

with. On the other hand, A. flavum Moro

3. Anemopaegma orbiculatum (Jacq.) DC., Prodr. 9: 190. 1845.-FIG. 3A-B.

4. Anemopaegma puberulum (Seib.) Miranda, Anal. Inst. Biol. Mexico 24:

Liana, branchlets subterete, puberulous or glabrate, the nodes without int

This species was supposedly restricted to Veracruz to Chiapas in southern

PANAMA: El Llano-Carti highway, 6-10 km N of El Llano, Dressler 4329 (MO, PMA)

5. Anemopaegma santa-ritense A. Gentry, Ann. Missouri Bot. Gard. 58: 93.

mm wide, 0.5 mm thick; disc conspicuous, conical-pulvinate, 1.5 mm long, 2.5 mm

1. Arrabidaea candicans (L. C. Rich.) DC., Prodr. 9: 185. 1845.-FIG. 4D-H.

FIGURE, 4. Arrabidaea.-A-C. A. florida DC.-A. Habit (X 3/5).-B

Liana to 7 cm in diameter, the bark yellowish to pale gray, smooth, 4-furrowed

Liana to 6 cm in diameter, the bark pale gray, smooth, 4-furrowed in age;

This is another of the species of Bignoniaceae with taxonomically confusing

3. Arrabidaea conjugata (Vell.) Mart., Flora 24(2): Beibl. 46. 1841.

4. Arrabidaea corallina (Jacq.) Sandw., Kew Bull. 1953: 460. 1954.

1.3-1.8 cm long and 3.6-5.2 cm wide, the wings hyaline-membranaceous at the

5. Arrabidaea florida DC., Prodr. 9: 184. 1845. TYPE: Brazil, Amazonas,

VascQncellia florida Mart. ex DC., Prodr. 9: 184. 1845, pro syn.

Liana to 5 cm in diameter, the bark gray, inconspicuously longitudinally

Mostly occuring in tropical moist forest, also in premontane wet forest,

lepidote, the ovules 2-seriate in each locule; disc pulvinate, 1 mm long 2 mm

An uncommon vine of the premontane moist forest and edaphically dry

8. Arrabidaea patellifera (Schlecht.) Sandw., Kew Bull. 22: 413. 1968.

9. Arrabidaea pubescens (L.) A. Gentry. Brittonia 25: 239. 1973.

Liana to 2.5 cm in diameter, stem with 4 phloem arms in cross section;

A plant mainly of the tropical dry forest, A. pubescens is widely distributed

One species ranging from Mexico to Brazil.

Callichlamys latifolia (L. C. Rich.) K. Schum. in Engler & Prantl, Nat.

FIGURE 5. Callichiamys latifolia (L. Rich.) K. Schlum.-A. Habit ( X 1/2). [After

pubescent on the undersurface, with plate-shaped glands along the midvein,

Inflorescence a short terminal or axillary, 1-12-flowered raceme, the pedicels

One species ranging from Mexico to Trinidad and Guyana.

Ceratophytum tetragonolobum (Jacq.) Sprague & Sandw., Kew Bull. 1934:

Liana to at least 7 cm in diameter, stem often indistinctly 4-parted and

.. .... ....~ ~ ~ ~~~~~~~~~~~~~~~~. ~

FIGURE 6. Ceratophytum tetragonolobum (Jacq.) Sprague & Sandw.-A. Habit (x 3/8).

A vine characteristic of the tropical moist forest, it is seldom common and