UNIVERSITY OF

JUL 2 4 1984

FIELDIANA
Botany

Published by Field Museum of Natural History

Volume 36, No. 4 April 30, 1973

Bignoniaceae of Tropical North America

Louis 0. WILLIAMS

CHAIRMAN, DEPARTMENT OF BOTANY
FIELD MUSEUM OP NATURAL HISTORY

AMPHITECNA AND ENALLAGMA. The tribe Crescentieae of the
Bignoniaceae in America, according to Schumann in Engler & Prantl,
Pflanzenfamilien IV, 3B; 244. 1895, consists of five genera, the best
known of which is Crescentia. The genus Schlegelia is an anomaly in
the Bignoniaceae and I have transferred it to Scrophulariaceae for the
"Flora of Guatemala." Parmentiera is easily distinguished from the
other genera of the tribe but the three remaining, Crescentia, Am-
phitecna, and Enallagma, are closely related. Crescentias are fairly
distinctive trees and the large globose or ovoid fruits are 1-celled.
The fruits of Amphitecna are said to be 2-celled while those of
Enallagma are said to be 1-celled or incompletely 2-celled. There are
certainly more species of the Amphitecna-Enallagma complex than
had been previously distinguished. Several of these are without
known fruits and even of those for which I have good fruits it is diffi-
cult if not impossible to be sure whether the fruit is 1- or 2-celled.

With the several species now distinguished in the Amphitecna-
Enallagma complex and the greater number of collections available I
find that the characters presumed to distinguish the two genera break
down completely and I am obliged to combine them. Dr. Seibert in
his "Bignoniaceae of the Maya Area" (Carnegie Inst. Wash. Publ.
522: 380-381. 1940) calls attention to this possibility, but had in-
sufficient material for study. Our material is still insufficient, and
probably always will be, but there is now three times as much mate-
rial as Dr. Seibert saw before 1940.

There are, at least, one or two more species in the herbaria which
are not accounted for here. Better material for them is desirable.

THE LIBRARY OF THE

Library of Congress Catalog Card Number: 73-806^9

Publication 1164 21 AUG 2 7 1973

UNIVERSITY OF ILLINOIS

AT

22 FIELDIANA: BOTANY, VOLUME 36

Amphitecna donnell-smithii (Sprague) L. Wms. comb. nov.
Crescentia donnell-smithii Sprague Bull. Herb. Boiss. II. 6: 376. 1906.
Enallagma donnell-smithii Standl. Field Mus. Bot. 12: 361. 1936.

The smallest flowered species of Amphitecna, known only from the
Caribbean lowlands of Guatemala and British Honduras.

Amphitecna molinae L. Wms. sp. nov.

Frutices aut arbusculae usque ad 6 m. Folia alterna vel subfasciculata ob-
lanceolata acuminata breviter petiolata chartacea vel coriacea; inflorescentia ter-
minalis fasculo bifloro, pediceli 2.5-3 cm. longi basi bibracteolati; calyx anguste
campanulatus trilobatus lobi ovati acuti; corolla alba infundibuliformis lobato-
crenata; fructus cortice crasso.

Shrub or weak tree 2-6 m. tall, the young branches striate, becoming terete,
grayish-white, glabrous. Leaves alternate or clustered toward the apices of young
branches, oblanceolate, acuminate, short-petiolate, lateral nerves 15-25 pairs at
about 80° angle from mid-nerve, glabrous, the blades chartaceous becoming
coriaceous with age, 8-17 cm. long and 2-6.5 cm. broad, the petiole to 1 cm. long;
inflorescence terminal, a fascicle of 2 flowers (on type); pedicels about 2.5-3 cm.
long, bibracteolate to near the base; calyx narrowly campanulate, divided to
about the middle and trilobate, about 2.5 cm. long at anthesis, the lobes ovate,
acute, about 1 cm. long; the corolla white, infundibuliform, lobate-crenate, to
about 5 cm. long; fruits with heavy shell 2-5 mm. thick, elliptic-oblong to ovate,
attenuate-acuminate to the apex, smooth, to 15 cm. long and 5.5-6.5 cm. in diam-
eter.

Honduras: flowers white, weak tree to 6 m. along river, open forest
of Quebrada Lejarsia between kms. 9-11 on road La Esperanza-
Marcala, dept. Intibuca, alt. 1,600 m., March 21, 1969, Molina &
Molina 24295 (type F, EAP) ; with the same data, Molina & Molina
24299 (F, EAP).

This small tree is easily distinguished from others of the genus by
the trilobate calcyes, the thick and durable shell of the fruit, the rela-
tively small leaves with numerous lateral nerves. The species memo-
rializes the Molinas who have collected assiduously in Central Amer-
ica for some 25 years. It is a highland species of the montane forest.

Amphitecna montana L. Wms. sp. nov.

Arbores usque ad 15m. altae, ramuli angulati vel teres glabri. Folia alterna
oblanceolata vel anguste obovato-elliptica acuminata basi cuneata longiusculo-
petiolata glabra subcoriacea; inflorescentiae uniflorae; flores pendentes pediceli
graciles; calyx carnosus coriaceus bilobatus aut trilobatus basi circumscissilis;
corolla campanulata leviter crenulato-lobata leviter zygomorpha; fructus elliptico-
ovoideus fragilis; semina crassa carnosa.

Small forest trees 10-15 m. tall the branches angulate at first becoming terete
with age, glabrous. Leaves alternate and scattered along the branches or some-

WILLIAMS: BIGNONIACEAE 23

times clustered toward the apex, oblanceolate to narrowly obovate-elliptic, acum-
inate, cuneate to the base and rather long petiolate (for genus), with 15-20 prom-
inent lateral nerves anastomosing toward the margins, blades subcoriaceous, 12-
30 cm. long and 3-10 cm. broad, the petioles mostly 2-4 cm. long; inflorescences of
single flowers in axils of leaves; flowers pendent, the pedicel slender, to about 10
cm. long; calyx fleshy, coriaceous, bilobate or possibly also trilobate to the mid-
dle, about 3 cm. long, circumscissile at the base and falling away leaving the pedi-
cel and young ovary; corolla light green, narrowly campanulate, shallowly crenate-
lobate, slightly zygomorphic, about 4.5 cm. long and about 2.5 cm. broad at throat;
fruits oblongoid or elliptic-ovoid, to 15 cm. long and 8 cm. in diameter, shell thin
and fragile, the flesh hard and slightly bitter; seeds thick and the cotyledons fleshy,
notched at both top and bottom, about 18 mm. broad, 12 mm. high, and 8-10 mm.
thick.

Guatemala: tree 50 ft. tall, flowers pendent, easily separating
from pedicel which likewise easily separates from stem pedicel or
branch, corolla pale greenish-yellow, anthers dull blackish brown;
mixed hardwood pine woods bordering barranco half way up slopes of
barranco, barrancos 6 miles south and west of town of Tajamulco,
northwestern slopes of Volcan Tajamulco, dept. San Marcos, alt.
2,300-2,800 m., Feb. 26, 1940, Steyermark 3670^ (type two sheets, F) ;
tree 12 m. tall, large fruits elliptic-ovate, about 15 cm. long and 8 cm.
in diameter, pulp surrounding seeds white, flowers white; montane
cloud forest area on outer slopes of Tajamulco Volcano, Sierra Madre
Mountains about 8-10 km. west of San Marcos, dept. San Marcos,
alt. ± 2,300 m., Dec. 31, 1964— Jan. 1, 1965, Williams et. al. 26915.

Mexico: Sil tepee, Chiapas, enero 1938, Matuda 19^7; Mt.
Ovando, Escuintla, Chiapas, 1-16 July 1940, Matuda ^206.

Sterile collections from Dept. San Marcos, Guatemala probably
this species are: Steyermark 36987, 37012; Standley 86200.

This is a montane species distinguished by the large petiolate
leaves, the long slender pedicels of the flowers and the curious calyx
circumscissile at the base. The fruit is perhaps the largest among the
species of Guatemala and the shell very fragile. The species has been
determined as "Enallagma sessilifolia" in all cases, a montane species
of Costa Rica to which it is allied.

Amphitecna oblanceolata L. Wms. sp. nov.

Frutices vel arbusculae usque ad 10 m., ramuli graciles, teretes vel angulati
albidi. Folia oblanceolata vel elliptico-oblanceolata acuminata basi cuneata
glabra petioli perbreves; inflorescentia terminalis vulgo uniflora; calyx bilabiatus,
ca. 2 cm. longus, lobi oblongo-ovati vel anguste obovati obtusi; corolla campanu-
lato-infundibuliformis zygomorpha 5-lobata; fructus anguste ovato-cylindricus.

24 FIELDIANA: BOTANY, VOLUME 36

Shrubs or small trees to 10 m. tall. Branches slender, terete or angulate, whit-
ish, glabrous. Leaves oblanceolate or elliptic-oblanceolate, acuminate, cuneate to
the base, glabrous, secondary nerves 10-15 pairs at nearly a right angle to mid-
nerves, the tips anastomosing, the blade often with small plate-shaped glands, 9-

25 cm. long and 2-7 cm. broad, petiole very short, 2-8 mm. long; inflorescence
terminal on young branches, mostly a single pedicellate flower, the pedicel 2-3 cm.
long; calyx bilabiate at an thesis, 1.5-2 cm. long and 0.5-0.8 cm. broad at the base,
the lobes broadly oblong-ovate to narrowly obovate, obtuse; corolla campanulate-
funnelform, zygomorphic, 5-lobate, 2.5-4.5 cm. long; fruit narrowly ovate-cylin-
dric, probably more than 15 cm. long and about 3.5-5 cm. in diameter; seeds
(nearly mature) immersed in the whitish pulp, subreniform in outline with the
embryo at the center and with lateral wings thin and obtuse.

Guatemala: Rio Chacon, Izabal, Feb. 10, 1921, Johnson 121+1;
shrub, between Bananera and "La Presa" in Montana del Mico,
Izabal, alt. 40-300 m., March 28, 1940, Steyermark 38210; at base of
waterfall, small tree 30 ft. tall, between Bananera and "La Presa" in
Montana del Mico, Izabal, alt. 50-150 m., April 9, 1940, Steyermark
39199; tree 30 ft. tall, wooded slopes, along Rio Frio, Izabal, alt. 75
m., Dec. 17, 1941, Steyermark 39944', tree 30 ft. tall, corolla creamy-
greenish with green calyx, along Rio Frio and tributaries, Izabal, alt.
75-150 m., Dec. 18, 1941, Steyermark 41587; shrub 8 ft. tall, Finca
Yalpemech near Alta Verapaz-Pete"n boundary line, alt. 100-150 m.,
March 23, 1942, Steyermark 45207; tree 30 ft. tall, lowland forest
along Rio San Roman west of Chinaja, Pete"n, alt. 50 m., March 30,
1942, Steyermark 45508; small tree, Cerro Chinaja, between Finca
Yalpemech and Chinaja, above source of Rio San Diego, Alta Vera-
paz, alt. 150-700 m., April 12, 1942, Steyermark 45575; arbusto 1 m.
alto, 6 cm. diametro, fruto verde, en foresta alta, San Luis en camino
para Poctun a kilometre 118, 3 diciembre de 1970, Tun Ortiz 1459
(type F, two sheets, with flowers and fruits; EAP, US).

These specimens except the holotype and two Steyermark collec-
tions (31999 and 41587) are sterile but they seem to all be the same
species. The fruit is said to have been bright green and soft on the
outside. Most specimens have been determined as "Enallagma
sessilifolia" and "E. donnell-smithii" to which they are not too closely
related.

The species is distinctive because of the subcoriaceous oblanceo-
late or elliptic-oblanceolate leaves with secondary nerves at nearly
right angle to the mid-nerve. The fruit is distinctive but fruits of
some members of this genus are not known to me.

The specimens cited are all from the lowland forests along the
Atlantic plain of Guatemala. The somewhat similar Amphitecna

WILLIAMS: BIGNONIACEAE 25

silvicola is easily distinguished by its cauliflorous inflorescences and its
curious venation underleaf and it is a resident of the mountain
forests.

Dr. B. F. Kukachka has studied the juvenile wood from the type
collection and comments that it agrees very well, anatomically, with
that of Amphitecna macrophylla, the only other species for which
wood is known.

Amphitecna obovata (Bentham) L. Wms., comb. nov. Cres-
centia obovata Bentham, Bot. Voy. Sulph. 130, t. 1+6. 1845. Enallagma
obovata Baill. ex. Schumann in Engler & Prantl, Pflanzenf. IV. 3b:
247. 1895.

A not uncommon shrub or small tree distributed in forests or in
mangrove swamps near or little above sea level from southern Mex-
ico, British Honduras, to Panama and the West Indies and to north-
ern South America. Also in southern Florida. This is the plant
which has been called Enallagma latifolia (Miller) Small in most all
recent works. See "Excluded species," p. 26.

Amphitecna sessilifolia (Donn.-Sm.) L. Wms. comb. nov.
Tabebuia sessilifolia Donn.-Sm. Bot. Gaz. 25: 156. 1898. Neo-
tuerckheimia gonoclada Donn.-Sm. Bot. Gaz. 47: 259. 1909. Enal-
lagma sessilifolia Standl. Field Mus. Bot. 18: 1120. 1938.

A species of the mountains of Costa Rica. It has been attributed
to Guatemala by Seibert and others but the specimens involved are
mostly the lowland A. oblanceolata.

Amphitecna silvicola L. Wms. sp. nov.

Frutex circa 2 m. altus. Folia elliptica vel oblanceolata acuminata breviter
petiolata; inflorescentiae fasciculae 1-3-floribus, pedicellis graciles; flos albus;
calyx sub anthesi bilobatum, lobi obovati cochleati obtusi; corolla campanulato-
infundibuliformis crenulato-lobata; stamina 4 inclusae; staminodeum cylindricum
arcuatum; anthera biloculares cellulae explanatae.

A shrub about 2 m. tall, the branches angulate, becoming terete, glabrous.
Leaves alternate on the new growths, elliptic or oblanceolate, acuminate, short
petiolate with 7-12 lateral nerves, these lighter-marginate on the lower surfaces,
the blades 6-12 cm. long and 2.2-4 cm. broad, petiole to 1 cm. long; inflorescence
fascicles of 1-3 flowers on older wood from an enlarged multi-bracteolate base,
pedicels about 2 cm. long, slender; flowers white; calyx closed in bud, bilaterally
split in anthesis nearly to the base, each lobe obovate, somewhat cochleate, ob-
tuse, glabrous, 15-20 mm. long and 10-12 mm. broad; corolla campanulate-fun-
nelform, crenulate-lobate, muriculate toward the apex outside, smooth and glab-

26 FIELDIANA: BOTANY, VOLUME 36

rous within, 4-5 cm. long when mature; stamens 4, in two unequal pairs, reaching
to the throat of the corolla; staminode cylindric, 3-4 mm. long, arcuate; anthers
2-celled, the cells explanate and joined through apex of anther, each cell about 5
mm. long; disc entire, obscurely lobate; young ovary elongate, smooth; style about
30 mm. long, narrowly truncate; fruits unknown.

Mexico: flowers white, [shrub] 6 ft. tall, heavily wooded slope at
Lago Tsiskaw on the Guatemalan border, 30 miles east of La
Trinitaria, municipio La Trinitaria, Chiapas, elevation 4,500 ft.,

27 May 1965, Breedlove 10064 (type, F).

This montane plant from middle elevations is related to A. macro-
phylla but quite distinct from it in the very much smaller leaves scat-
tered along the new growth instead of clustered on the new growth.
Flower structure seems not greatly different in the two species, and
both share the curious lighter colored margins of the veins and vein-
lets on the lower leaf surface.

EXCLUDED SPECIES

CRESCENTIA LATIFOLIA Miller, Gard. Diet. ed. 8. 1768. C.
cucurbitina L. Mant. PI. 250. 1771. Enallagma cucurbitina Baill. Hist.
PL 10: 24. 1888. E. latifolia Small, Fl. Miami 171. 1913.— Nomina
ambigua.

Crescentia latifolia has been interpreted as the not uncommon low-
land plant that occurs along the coasts of Florida, Mexico, Central
America, the West Indies south to northern South America. Miller's
description of Crescentia latifolia does not fit the plant which it has
generally been assumed to describe. Miller says that he had the plant
under cultivation and mentions that Dr. Houstoun sent fruits from
Campeachy. Whether Miller's description is based on plants grown
from this Mexican seed I have no way of knowing, but I am aware of
no plant from Campeche that fits Miller's description — among others
the leaves described are much too small and of the wrong shape.

Dr. William T. Steam has been kind enough to look into the
herbarium of British Museum for Miller's specimen and writes that:
"There is no specimen either from Miller or from Houstoun in our
herbarium to typify the name Crescentia latifolia Miller and the
annotated copy of Miller's Dictionary, 8th edition, indicates that
Banks never received one. Hence the name must be interpreted
purely from Miller's description, which is undoubtedly puzzling, as
you note."

Since Miller's description fits no plant of this relationship known
to me from Campeche and, from the description, it is not the plant

WILLIAMS: BIGNONIACEAE 27

which has gone under this name, I have taken up, as Amphitecna
obovata, the next available name for the plant Bentham's beautifully
illustrated and described Crescentia obovata. See above.

PARMENTIERA. There are three species of Parmentiera in Mexico
and Guatemala and four additional species have been described from
Nicaragua, Costa Rica, and Panama, but these do not affect the
treatment of the genus for the "Flora of Guatemala."

Parmentiera aculeata (HBK.) L. Wms. comb. nov. Crescentia
aculeata HBK. Nov. Gen. & Sp. PI. 3: 158. 1819. Parmentiera edulis
DC. Prodr. 9: 244. 1845. Crescentia edulis Moc. ex DC. I.e., nomen
nudum. Parmentiera aculeata Seemann, Bot. Voy. Herald 183. 1854,
nomen nudum.

Crescentia aculeata HBK. was described from Campeche, Mexico.
The type specimen is in Paris, where it was annotated by N. Y. Sand-
with in 1937 and later photographed by Macbride (F 39404). It has
been generally assumed that the combination to Parmentiera was
made by Seemann in Bot. Voy. Herald 183. 1854. The name given by
Seemann, in text, is a nomen nudum, for he cited no basionym and
gave no description. He did say "The P. aculeata Seem., is founded
on a specimen in Hooker's Herbarium." Therefore Parmentiera
aculeata Seem, is not based on the type of Crescentia aculeata and is
nothing more than an unfortunate nomen nudum, not a new combina-
tion. It does not prevent the proper combination to Parmentiera
being made.

Parmentiera edulis DC. Prodr. 9: 244. 1845, is based upon an un-
published Mocino (Sesse" & Mocifio) drawing in the De Candolle
herbarium (phototype F, 30770) for De Candolle cites an unpub-
lished name "Crescentia edulis Moc. fl. Mex. ic. ined. non Desv." and
used this specific name for his Parmentiera edulis. The illustration is
the type for both the generic and the specific names.

A comparison of these type photographs of Crescentia aculeata
HBK. and Parmentiera edulis DC. leaves no doubt that they repre-
sent the same species, one that doubtless has been carried about by
man for its edible (hardly) fruits. Pringle's No. 7524 from Oaxaca,
Mexico, is an excellent representation of this species, showing the
large corollas as on the type of P. aculeata HBK. (which shows no
fruits) and the thick costate fruits as on the type illustration of P.
edulis. The leaves are so nearly alike on both types that there can be
no question that only one species is involved.

28 FIELDIANA: BOTANY, VOLUME 36

Parmentiera millspaughiana L. Wms. sp. nov.

Frutices vel arbores parvae aculeatae usque ad 6 m. altae. Folia trifoliolata
vel Integra, foliolis late orbiculari-spathulatis acutis vel obtusis vel rotundatis,
basi cuneata; petioli alati vel exalati; inflorescentia terminalis vulgo uniflora;
calyx spathatheus; corolla infundibuliformis; fructus teretes elongati laeves
obscure costati.

Shrubs or small trees to 6 m. tall; the branches slender terete, grayish, with
alternate or paired spines at the nodes, these sometimes (the upper ones) bearing
small leaves at their apices; leaves trifoliolate or sometimes simple, obscurely glan-
dular-lepidote and sparsely puberulent along the veins below, glabrous above, the
leaflets broadly orbicular-spathulate, acute or usually obtuse or rounded at the
apex, cuneate to the base, the terminal leaflet or simple leaf 1-2.5 cm. long and
0.6-2 cm. broad, the lateral leaflets similar but smaller, petioles to 3 cm. long,
alate or not, usually puberulent, those of simple leaves almost lacking; inflorescence
terminal, mostly 1-flowered, pedicels about 1-2 cm. long, slender bibracteolate
near the base; calyx spathaceous, up to 2.5 cm. long; corolla infundibuliform, about
4-4.5 cm. long; fruits terete, elongated, smooth or only obscurely costate, the apex
apiculate, 13-25 cm. long and about 1-1.5 cm. in diameter, yellow when ripe.

Mexico: shrub 12 ft. high, common about Izamal, producing its
neutral green flowers throughout the year, 1895-96, Gaumer 338
(type, F; AA; MO; NY; US); San Anselmo, Gaumer 1613; Catcuuc;
cat; shrub 15 ft. high and from 1-2 in. in diameter, it blooms in May,
forests of Suitun, Gaumer & Sons 23289; shrub 8 ft. tall, sometimes
higher, seldom has more than one flower on at a time, its fruit when
ripe is yellow, in Suitun woods, October, 1916, Gaumer & Sons 23439;
small 3 m. tree or bush, Chan Laguna, Campeche, Dec. 5, 1931,
Lundell 1021; MeYida, May 25, 1865, Schott 382; Merida, June 28,
1865, Schott 382 (sic).

Related to Parmentiera parviflora Lundell of the Yucatan Penin-
sula in Mexico and Guatemala and to P. cerifera Seemann of Panama.
It is easily distinguished from both by its orbicular-spathulate, ob-
tuse leaflets; from P. parviflora by its much larger flowers and larger
and relatively thinner fruit; from P. cerifera by its terminal, not
cauliflorous, inflorescence, much smaller fruits, and great geographical
separations.

I am pleased to name this plant for Dr. Charles F. Millspaugh,
first scientific curator at Field Museum and for many years head of
the Botany Department; first American botanist with an especial
interest in the flora of Yucatan and the author of the first serious
contributions to the Yucatecan flora; my predecessor at Field Mu-
seum by 65 years. It was Dr. Millspaugh who brought all the speci-
mens cited above, except one, to the herbarium which he founded.

WILLIAMS: BIGNONIACEAE 29

Dr. Siebert (in Carnegie Inst. Wash. Publ. 552: 385. 1940) used
the name "P. aculeata (HBK.) Seem." for this plant in exactly the
same circumscription. It is shown above that P. aculeata (HBK.)
L. Wms. includes P. edulis DC. and must apply to the more common
and widespread species. No name has been found which applies to
this long recognized taxon.

Parmentiera parviflora Lundell, Lloydia 3: 211. 1940 is a little
known species from the Yucatan Peninsula of Mexico and Guatemala
that is easily distinguished from all species of the genus by its rela-
tively small flowers. It has been found for the first time in Guatemala
near Santa Elena, Dept. Pete"n, Tun 835.

UNIVERSITY OF ILLINOIS-URBANA